Zootaxa, Revision of Oriental Phasmatodea: The tribe ...

ZOOTAXA 1906

Revision of Oriental Phasmatodea: The tribe Pharnaciini Günther, 1953, including the description of the world's longest insect, and a survey of the family Phasmatidae Gray, 1835 with keys to the subfamilies and tribes (Phasmatodea: "Anareolatae": Phasmatidae) FRANK H. HENNEMANN & OSKAR V. CONLE

Magnolia Press Auckland, New Zealand

Frank H. Hennemann & Oskar V. Conle Revision of Oriental Phasmatodea: The tribe Pharnaciini Günther, 1953, including the description of the world's longest insect, and a survey of the family Phasmatidae Gray, 1835 with keys to the subfamilies and tribes (Phasmatodea: "Anareolatae": Phasmatidae) (Zootaxa 1906) 316 pp.; 30 cm. 15 Oct. 2008 ISBN 978-1-86977-271-0 (paperback) ISBN 978-1-86977-272-7 (Online edition)

FIRST PUBLISHED IN 2008 BY Magnolia Press P.O. Box 41-383 Auckland 1346 New Zealand e-mail: [email protected] http://www.mapress.com/zootaxa/

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2 · Zootaxa 1906 © 2008 Magnolia Press

HENNEMANN & CONLE

Zootaxa 1906: 1–316 (2008) www.mapress.com / zootaxa/

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Copyright © 2008 · Magnolia Press

ISSN 1175-5334 (online edition)

ZOOTAXA

Revision of Oriental Phasmatodea: The tribe Pharnaciini Günther, 1953, including the description of the world’s longest insect, and a survey of the family Phasmatidae Gray, 1835 with keys to the subfamilies and tribes* (Phasmatodea: “Anareolatae”: Phasmatidae) FRANK H. HENNEMANN¹ & OSKAR V. CONLE² ¹ Kirchdell 24, 67661 KL-Hohenecken, Germany. E-mail: [email protected] ² Goldbachweg 24, 87538 Bolsterlang, Germany. E-mail: [email protected] Website: www.Phasmatodea.com

Table of contents Abstract ............................................................................................................................................................................... 5 1 Introduction ...................................................................................................................................................................... 8 2 Material & methods .......................................................................................................................................................... 8 2.1 Abbreviations ......................................................................................................................................................... 9 3 Basic background ........................................................................................................................................................... 11 3.1 The present classification ..................................................................................................................................... 11 3.2 Characters and evolutionary trends ...................................................................................................................... 12 3.3 Ootaxonomy ......................................................................................................................................................... 16 4 Family Phasmatidae Gray, 1835 S. STR. (= Lanceocercata Bradler, 2001) .................................................................. 17 4.1 The Phasmatidae and Phasmatinae sensu Bradley & Galil, 1977 ........................................................................ 20 4.2 Keys to the subfamilies and tribes of Phasmatidae s. str. .................................................................................... 24 4.3 Phasmatinae Gray, 1835 ....................................................................................................................................... 26 4.3.1 Phasmatini Gray, 1835 ............................................................................................................................... 28 4.3.2 Acanthomimini Günther, 1953 .................................................................................................................. 32 4.3.3 Acanthoxylini Bradley & Galil, 1977 ........................................................................................................ 35 4.4 Tropidoderinae Brunner v. Wattenwyl, 1893 ....................................................................................................... 37 4.4.1 Tropidoderini Brunner v. Wattenwyl, 1893 ............................................................................................... 39 4.4.2 Monandropterini Brunner v. Wattenwyl, 1893 .......................................................................................... 43 4.4.3 Gigantophasmatini trib. nov. ...................................................................................................................... 44 4.5 Extatosomatinae Clark-Sellick, 1997 stat. nov. ................................................................................................... 46 4.6 Problematic taxa ................................................................................................................................................... 48 4.6.1 Subfamily Pachymorphinae Brunner v. Wattenwyl, 1893 ......................................................................... 48 4.6.2 Subfamily Xeroderinae Günther, 1953 ...................................................................................................... 52 4.7 Tribes excluded from the family Phasmatidae s. str. ............................................................................................ 53 4.7.1 Achriopterini Günther, 1953 ...................................................................................................................... 54 4.7.2 Stephanacridini Günther, 1953 .................................................................................................................. 54 5 Subfamily Clitumninae Brunner V. Wattenwyl, 1893 .................................................................................................... 62 5.1 Pharnaciini Günther, 1953 .................................................................................................................................... 66 5.2 Clitumnini Brunner v. Wattenwyl, 1893 .............................................................................................................. 67 5.3 Medaurini trib. nov. .............................................................................................................................................. 72 * Dedicated to the authors’ parents Gisela & Hans-Peter Hennemann and Xenia & Matthäus Conle

Accepted by B. Mantovani: 29 Jul. 2008; published: 15 Oct. 2008

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5.4 Subfamily Lonchodinae Brunner v. Wattenwyl, 1893 ......................................................................................... 73 5.4.1 Lonchodini Brunner v. Wattenwyl, 1893 ........................................................................................................... 75 5.4.2 Neohiraseini trib. nov. ............................................................................................................................... 78 5.5 Keys to the tribes of Clitumninae and Lonchodinae ............................................................................................ 79 6 Revision of the tribe Pharnaciini Günther, 1953 ............................................................................................................ 80 6.1 Genus Baculonistria gen. nov. ............................................................................................................................. 85 6.2 Genus Pharnacia Stål, 1877 ................................................................................................................................ 95 6.3 Genus Phobaeticus Brunner v. Wattenwyl, 1907............................................................................................... 129 6.4 Genus Phryganistria Stål, 1875 ......................................................................................................................... 199 6.5 Genus Tirachoidea Brunner v. Wattenwyl , 1893 stat. rev. ................................................................................ 218 7 Species incertus sedis ................................................................................................................................................... 252 8 Discussion of Pharnaciini Günther, 1953 ..................................................................................................................... 252 8.1 Biogeography ..................................................................................................................................................... 257 8.2 Habitats & ecology ............................................................................................................................................. 261 9 Taxonomic summary .................................................................................................................................................... 262 9.1 New taxa and new names ................................................................................................................................... 262 9.2 List of taxonomic changes ................................................................................................................................. 262 9.3 Taxonomic catalogue of Pharnaciini Günther, 1953 .......................................................................................... 266 10 Conclusion .................................................................................................................................................................. 268 11 Acknowledgements .................................................................................................................................................... 269 12 Addendum .................................................................................................................................................................. 311 13 Index of scientific names ........................................................................................................................................... 315


HENNEMANN & CONLE

Abstract The family Phasmatidae Gray, 1835 is reviewed and the subfamily Phasmatinae shown to be polyphyletic. Based on features of the exosceleton of the insects, egg-morphology and copulation habits a new arrangement of Phasmatidae is proposed. The monophyly of Lanceocercata Bradler, 2001 is confirmed but this name shown to be a synonym of Phasmatidae, hence Lanceocercata is here referred to as Phasmatidae sensu stricto. Six subfamilies belong in Phasmatidae sensu stricto all of which share several common and supposedly apomorphic characters: Phasmatinae, Tropidoderinae, Extatosomatinae (stat. nov.), Xeroderinae, Pachymorphinae and “Platycraninae”. The other two subfamilies contained in Phasmatidae sensu Bradley & Galil, 1977 (Eurycanthinae and Cladomorphinae) are not cosely related and here regarded as subfamilies of Phasmatidae sensu lato. The subfamily Phasmatinae sensu Bradley & Galil, 1977 is shown to be polyphyletic. The two tribes Pharnaciini and Clitumnini (= Baculini Günther, 1953) are removed from Phasmatinae and shown to be closely related to each other. They are transferred to the here established subfamily Clitumninae, a subordinate clade of Phasmatidae sensu lato. The subfamily Lonchodinae is closely related to Clitumninae, hence removed from Diapheromeridae and transferred to Phasmatidae sensu lato. The tribes Achriopterini and Stephanacridini (formerly in Phasmatinae) are shown to be not closely related to either Phasmatinae sensu stricto, Clitumninae or Lonchodinae, and provisionally must be treated as tribes of Phasmatidae sensu lato (incerte sedis). A re-arrangement of Phasmatidae sensu stricto is proposed along with determinating keys to all subfamilies and their tribes. The subfamilies Phasmatinae, Tropidoderinae and Extatosomatinae stat. nov. are re-described and discussed in detail. Full lists of genera are provided for each tribe. Only three of seven tribes formerly in Phasmatinae remain in the subfamily, this is Phasmatini, Acanthomimini and Acanthoxylini. The subfamily Tropidoderinae contains three tribes: Tropidoderini, Monandropterini and Gigantophasmatini trib. nov. The tribe Extatosomatini Clark-Sellick, 1997 is removed from Tropidoderinae and raised to subfamily level (Extatosomatinae stat. nov.). Several genera are transferred to other tribes or subfamilies. Didymuria Kirby, 1904 is removed from Tropidoderini, since it differs by having a closed internal micropylar plate in the eggs (open in all Tropidoderini). It here remains as a genus incerte sedis of Tropidoderinae and its systematic position clearly deserves further clarification. Gigantophasma Sharp, 1898 from the Loyalty Islands is removed from Pharnaciini, and becomes the type genus of the tribe Gigantophasmatini trib. nov.. Anophelepis Westwood, 1859 is removed from “Platycraninae” and shown to belong in Phasmatinae: Acanthomimini. The two Australian genera Arphax Stål, 1875, and Vasilissa Kirby, 1896 are removed from Acanthoxylini and provisionally transferred to Acanthomimini, but their position remains as yet debatable. Echetlus Stål, 1875 is misplaced in “Platycraninae” and shown to be a likely member of Phasmatinae. The two Brazilian species Echetlus evoneobertii Zompro & Adis, 2001 and Echetlus fulgens Zompro, 2004b are obviously misplaced and belong in the New World Diapheromeridae: Diapheromerinae: Diapheromerini. The subfamily Pachymorphinae is briefly discussed and considered polyphyletic. Two genera of Pachymorphinae: Gratidiini Bragg, 1995 (Parapachymorpha Brunner v. Wattenwyl, 1893 and Cnipsomorpha Hennemann et al., 2008) are transferred to Clitumninae: Medaurini trib. nov. The genus Gongylopus Brunner v. Wattenwyl, 1907 is transferred from Pachymorphinae: Gratidiini to Clitumninae: Clitumnini. The subfamily Xeroderinae is briefly discussed and shown likely to be polyphyletic, due to it contains two fundamentally different types of genitalia in the males. Only the genera Xeroderus Gray, 1835 and perhaps Epicharmus Stål, 1875 clearly belong in Phasmatidae sensu stricto. Both, the Pachymorphinae and Xeroderinae certainly deserve more detailed investigation to clarify their systematic positions with confirmation. Two generic groups are recognized within Clitumnini (subfamily Clitumninae). Due to differing by genital features and egg-morphology Medaura Stål, 1875 and Medauroidea Zompro, 2000 are removed from Clitumnini and transferred to the newly described Medaurini trib. nov.. The new tribe furthermore contains two genera formerly included in Pachymorphinae:Gratidiini and transferred here, Cnipsomorpha Hennemann et al., 2008 and Parapachymorpha Brunner v. Wattenwyl, 1893. Phryganistria Stål, 1875 is removed from Clitumnini and transferred to Pharnaciini. Nesiophasma Günther, 1934 is shown to belong in the tribe Stephanacridini. The Australasian subfamily Lonchodinae Brunner v. Wattenwyl, 1893 has formerly been included in Diapheromeridae Zompro, 2001 (= Heteronemiidae by Bradley & Galil, 1977). However, numerous features of the genitalia and egg morphology show close relation to the Oriental subfamily Clitumninae instead. Thus, Lonchodinae is here transferred to the family Phasmatidae (sensu lato). Within Lonchodinae the new tribe Neohiraseini trib. nov. is recognized and contains the five genera formerly placed in the “Neohirasea-complex” of that subfamily, namely Andropromachus Carl, 1913, Neohirasea Rehn, 1904, Pseudocentema Chen, He & Li, 2002, Qiongphasma Chen, He & Li, 2002 and Spinohirasea Zompro, 2001. It differs from all other Lonchodinae (= tribe Lonchodini) by the well developed vomer of males and the lack of a capitulum in the eggs. The genus Cladomimus Carl, 1915 was previously misplaced in Clitumninae:

REVISION OF PHARNACIINI

Zootaxa 1906 © 2008 Magnolia Press ·

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Pharnaciini and is here transferred to Lonchodinae: Lonchodini. It appears to be close to the Australian Hyrtacus Stål, 1875. Leprocaulinus Uvarov, 1940 and Phenacocephalus Werner, 1930 are removed from the subfamily Necrosciinae and transferred to Lonchodinae: Lonchodini. Extensive research on the genera which belong to the tribe Pharnaciini Günther, 1953 and taking features of the genital exosceleton and egg-morphology into account, has shown this tribe to be polyphyletic. Based on such features two generic groups are easily recognized within Pharnaciini sensu Günther, 1953. Males of the first group have a longitudinally split anal segment, which consists of two separate, more or less elongate semi-tergites and forms a clasping apparatus, the vomer is strongly reduced or lacking, the profemora have a prominent, lamellate medioventral carina which is strongly displaced towards the anteroventral carina and the eggs have an open internal micropylar plate with a clear median line. Only the genera falling into this group remain in Pharnaciini. Males of the second group in contrast have an anal segment which is not split, but possess a clearly visible, well sclerotised, triangular or hook-like external vomer, an indistinct medioventral carina on the profemora and eggs with a closed internal micropylar plate. Most of the genera which fall into the second group are here transferred to the tribe Stephanacridini Günther, 1953, this is Diagoras Stål, 1877b, Eucarcharus Brunner v. Wattenwyl, 1907, Phasmotaenia Návas, 1907 and Sadyattes Stål, 1875. A detailed discussion of the differences between Pharnaciini and Stephanacridini is provided along with distinguishing keys, illustrations and maps showing the distinct geographic distributions. The five genera that belong in Pharnaciini are: Baculonistria gen. nov., Pharnacia Stål, 1877a, Phobaeticus Brunner v. Wattenwyl, 1907 (= Baculolonga Hennemann & Conle, 1997a, = Lobophasma Günther, 1934b syn. nov. , = Nearchus Redtenbacher, 1908 syn. nov. ), Tirachoidea Brunner v. Wattenwyl, 1893 stat. rev. and Phryganistria Stål, 1875. Pharnacia annulata Redtenbacher, 1908 and Pharnacia enganensis Redtenbacher, 1908 were misplaced in Pharnacia Stål, 1877 (tribe Pharnaciini) and are transferred to the genus Sadyattes Stål, 1875 (tribe Stephanacridini, comb. nov.). Phobaeticus kuehni Brunner v. Wattenwyl, 1907 is removed from Phobaeticus Brunner v. Wattenwyl, 1907 (Phasmatinae: Pharnaciini) and shown to belong in Nesiophasma Günther, 1934c (tribe Stephanacridini, comb. nov.). Phobaeticus incertus Brunner v. Wattenwyl, 1907 (= Nearchus grubaueri Redtenbacher, 1908 syn. nov.) is unlikely to belong in Pharnaciini and here only retained in the original genus Phobaeticus Brunner v. Wattenwyl, 1907 with doubt, it may belong in Nesiophasma Günther, 1934c (tribe Stephanacridini). Based on a total of almost 700 examined specimens, the Oriental tribe Pharnaciini Günther, 1953 is revised at the species level. The new genus Baculonistria gen. nov. (Type species Baculonistria alba (Chen & He, 1990) comb. nov.), is described to contain three species from Central and Eastern China. Tirachoidea Brunner v. Wattenwyl, 1893 was erroneously synonymised with Pharnacia Stål, 1877 and is here re-established as a valid genus (stat. rev.). All five genera are re-diagnosed and differentiated, their systematic position within Pharnaciini discussed, and complete synonymic and species-listings as well as distribution maps and determination keys to the insects and eggs are provided. Detailed descriptions, diagnoses, synonymic listings, illustrations, material listings, distribution maps and measurements are provided for all 42 valid species. The type material of a further two species appears to be lost. Seven new species are described: Pharnacia borneensis spec. nov. from Borneo; Pharnacia palawanica spec. nov. from Palawan, Phobaeticus mucrospinosus spec. nov. from Sumatra, Phobaeticus palawanensis spec. nov. from Palawan, Tirachoidea herberti spec. nov. from Borneo, Tirachoidea siamensis spec. nov. from Thailand and S-Vietnam and Phobaeticus chani Bragg spec. nov. from Borneo. Phobaeticus chani Bragg spec. nov. is the world’s longest known insect with a maximum body length of 357 mm and an overall length of 567 mm in the female. Twelve new synonymies were discovered: Bactridium grande Rehn, 1920 = Phobaeticus serratipes (Gray, 1835) syn. nov.; Pharnacia rigida Redtenbacher, 1908 = Phobaeticus sumatranus Brunner v. Wattenwyl, 1907, syn. nov.; Clitumnus irregularis Brunner v. Wattenwyl, 1907 = Phibalosoma tirachus Westwood, 1859, syn. nov.; Pharnacia magdiwang Lit & Eusebio, 2008 = Pharnacia ponderosa Stål, 1877 syn. nov.; Pharnacia spectabilis Redtenbacher, 1908 = Phibalosoma hypharpax Westwood, 1859, syn. nov.; Pharnacia semilunaris Redtenbacher, 1908 = Eucarcharus inversus Brunner v. Wattenwyl, 1907, syn. nov.; Pharnacia chiniensis Seow-Choen, 1998c = Pharnacia biceps Redtenbacher, 1908, syn. nov.; Nearchus grubaueri Redtenbacher, 1908 = Phobaeticus incertus Brunner v. Wattenwyl, 1907, syn. nov.; Phibalosoma maximum Bates, 1865 = Cladoxerus serratipes Gray, 1835, syn. nov.; Phobaeticus lambirica Seow-Choen, 1998a = Eucarcharus rex Günther, 1928, syn. nov.; Phobaeticus sichuanensis Cai & Liu, 1993 = Baculum album Chen & He, 1990, syn. nov. and Phobaeticus beccarianus Brunner v. Wattenwyl, 1907 is shown to represent the previously unknown female of Phobaeticus sobrinus Brunner v. Wattenwyl, 1907 (syn. nov.) Lectotypes are designated for: Nearchus redtenbacheri Dohrn, 1910, Pharnacia biceps Redtenbacher, 1908, Pharnacia ingens Redtenbacher, 1908, Pharnacia heros Redtenbacher, 1908, Phibalosoma westwoodi Wood-Mason, 1875, Phobaeticus sinetyi Brunner v. Wattenwyl, 1907, and Phobaeticus sumatranus Brunner v. Wattenwyl, 1907. A neotype is designated for Nearchus maximus Redtenbacher, 1908 and Phobaeticus magnus nom. nov. introduced as a replacement


HENNEMANN & CONLE

name for Nearchus maximus Redtenbacher, which is a junior homonym of Phibalosoma maximum Bates, 1865. The previously unknown males of Pharnacia heros Redtenbacher, 1908, Phobaeticus ingens (Redtenbacher, 1908), Tirachoidea jianfenglingensis (Bi, 1994), Pharnacia sumatrana (Brunner v. Wattenwyl, 1907), Phryganistria fruhstorferi (Brunner v. Wattenwyl, 1907) and Tirachoidea westwoodii (Wood-Mason, 1875) as well as the females of Pharnacia ponderosa Stål, 1877a and Pharnacia tirachus (Westwood, 1859) are described and illustrated for the first time. A brief description on the basis of colour photos of the so far unknown male of Pharnacia kalag Zompro, 2005 are presented. Detailed descriptions and illustrations are provided for the eggs of 24 species. The eggs of the following 18 species are described and illustrated for the first time: Phobaeticus magnus nom. nov., Pharnacia borneensis spec. nov., Pharnacia palawanica spec. nov., Pharnacia ponderosa Stål, 1877a, Pharnacia sumatrana (Brunner v. Wattenwyl, 1907), Pharnacia tirachus (Westwood, 1859), Phobaeticus hypharpax (Westwood, 1859), Phobaeticus chani Bragg spec. nov., Phobaeticus incertus Brunner v. Wattenwyl, 1907, Phobaeticus magnus nom. nov., Phobaeticus philippinicus (Hennemann & Conle, 1997a), Phobaeticus sinetyi Brunner v. Wattenwyl, 1907, Phryganistria grandis Rehn, 1906, Phryganistria virgea (Westwood, 1848), Tirachoidea biceps (Redtenbacher, 1908), Tirachoidea herberti spec. nov., Tirachoidea jianfenglingensis (Bi, 1994) and Tirachoidea siamensis spec. nov.. Several species were originally placed in or subsequently transferred into wrong genera by various authors. Consequently, numerous taxa are here transferred or re-transferred to other genera, which results in 22 new or revised combinations or status of genera and species (comb. nov. / stat. rev. / stat. nov.). A list of the taxonomic changes made in this revision is provided in the summary ( 9.2), which in all lists 70 nomenclatural changes. Key words: Phasmatodea, Phasmatidae, Lanceocercata, keys, diagnoses, subfamilies, tribes, Extatosomatinae stat. nov., Gigantophasmatini trib. nov., Clitumninae, Medaurini trib. nov., Lonchodinae, Neohiraseini trib. nov., Pharnaciini, Baculonistria gen. nov., Pharnacia, Phobaeticus, Phryganistria, Tirachoidea, classification, ootaxonomy, taxonomic revision, keys, differentiations, descriptions, new genus, new species, new synonyms, new combinations, eggs, biogeography, distribution, maps, habitats, ecology



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1 Introduction Although several of the largest and most spectacular insects belong to the orthopteroid order Phasmatodea, even groups such as the Oriental Pharnaciini Günther, 1953 and Australasian Stephanacridini Günther, 1953, which contain some of the world’s longest known insects, have so far remained only fractionally studied. The lack of useful identification keys at generic level is still one of the most striking problems and due to the strong sexual dimorphism attributing males or females to a species described from just one sex has remained problematic. A further serious problem is the considerable variation of many taxa and consequently, up to now the identification of genera or species is still difficult in various subgroups of the order. So far not enough attention has been paid to the systematic value of ootaxonomy and features of the genitalia for arranging a classification of Phasmatodea, and consequently there is still no satisfactory system based on natural relationships. One of the basic problems is that many genera were insufficiently characterized or not even understood by their describers. Consequently, there have been numerous misinterpretations of genera by subsequent authors which is why some e.g. Pharnacia Stål, 1877 or Hermarchus Stål, 1875 are highly polyphyletic. Since starting studies on the tribe Pharnaciini Günther, 1953 in 1994, and visiting many European museums which contain the necessary type material, several new taxa, undescribed sexes and eggs were discovered; these helped clarify the systematic position or status of certain species. Soon it became obvious that the tribe is polyphyletic and that several taxonomic changes within Pharnaciini were necessary. Studies on the systematic position of Pharnaciini and comparison with related tribes have shown the subfamily Phasmatinae sensu Bradley & Galil, 1977 to be an artificial group as well. The polyphyly of Phasmatinae has already been suggested by various authors (Bradler, 2000; Tilgner, 2002; Whiting et al., 2003). Bradler (2001) identified a supposedly monophyletic group of predominantly australasian taxa, the Lanceocercata, which contains sections of several taxa traditionally referred to as subfamilies, hence suggesting these to be polyphyletic. For clarifying the systematic position of Pharnaciini within Phasmatidae a detailed, critical review of the complete family with new determination keys to the tribes was urgently needed. The main subject of the present work is a detailed taxonomic revision of the tribe Pharnaciini Günther, 1953 at specific level, including the diagnosis, description, differentiation and illustration of all known genera and species, the description of new taxa, newly discovered sexes and eggs as well as providing detailed information on the geographic distribution of each taxon. Furthermore, in order to explain the systematic position of Pharnaciini Günther, 1953 a new a critical review of the family Phasmatidae Gray, 1833 is provided along with new diagnoses, list of species and determinating keys to the subfamilies and tribes of Phasmatidae sensu stricto. It is hoped that this work will be a good basis for future studies on these fascinating insect-giants and a useful tool for identifying the genera and species of Pharnaciini.

2 Material & methods Whenever possible, this study was based on examination of the necessary type material and received much support from most curators of the corresponding European museums and institutions containing type specimens. In a very few cases the examination of the type specimens was not possible, but good photographs, data, measurements and non-type material were available. The type material of four species was not traced in any of the museums visited and inquiries to other institutions in order to trace the specimens concerned have not received any reply. If the type material was not directly examined this is clearly mentioned in the comments of the corresponding taxon. In addition to the types, all other material available from various public or private collections was examined and determined, making a total of more than 650 specimens of Pharnaciini Günther, 1953 examined for this work.


HENNEMANN & CONLE

Breeding of several species and examination of large series of specimens confirmed the genitalia and the external and internal egg morphology to show the smallest range of variation, hence these are the best characters for the differentiation of taxa and emphasised in the present study. Additionally, the relative length ratio of the median segment and metanotum has proven to be a helpful feature for the differentiation of genera. The armature of the extremities, which has been used as the main distinguishing feature by most former authors, is subject to strong intraspecific variation in most species and therefore of little use for the distinction of taxa. The material used for this study is invariably dried and pinned. Captive reared material in the first author’s collection (FH) was preserved as described by Zompro (1996) or with silica-gel, material in the second author’s collection (OC) with acetone or silica-gel. Insects and eggs were examined using an magnifying glass with 4x magnification and a Russian MBC-10 binocular microscope. Measurements were taken using a long ruler or a digital calliper and are given to 0.1 mm. If not differently cited, measurements include all examined and listed material as well as the type specimens, although measurements of type specimens are in most cases listed separately. If more than one egg has been examined, average measurements are given. The body-lengths cited in the determinating keys for && are including the subgenital plate. If not differently cited and no live material was available, the colouration is described from dried specimens, to which attention needs to be drawn, as the insects may have changed colour due to preservation. All eggs examined were already laid or removed from the female’s ovipositor and were thus fully developed. If eggs were extracted from the abdomen this is clearly mentioned. The terminology used for the descriptions of external and internal egg structures follows that of Clark-Sellick (1997a). Lectotypes are designated for nine taxa. The revision of the tribe Pharnaciini on specific level is reason enough for the designation of lectotypes and clearly justifies these selections according to Article 74 of the Code (ICZN 1999). For the discussion of taxa, the following format is used. Genera and species are listed in alphabetical order. The data of type material is included in the synonymic lists and follows the corresponding citation. The section “Further material examined” lists the complete examined non-type material, arranged in geographical order. The differential diagnosis is followd by descriptions of the adults and eggs, and comments which provide information on the synonymy, variation, distribution, biology, foodplants or culture status of the species. Finally the complete number of specimens examined for this work (including the types) is provided at the end of each section, followd by tables with measurements.

2.1 Abbreviations ANIC ANSP BFU BMNH CAU DBNU ETHZ FRIM FRCS HLDH HNHM ISNB IZCAS KNP

Australian National Insect Collection, Canberra / Australia. Academy of Natural Sciences, Philadelphia / U.S.A. Beijing Forestry University, Beijing / China. British Museum of Natural History, London / England. China Agricultural University, Beijing / China. Department of Biology, Nankai University, Tianjin / China. Eidgenössische Technische Hochschule, Zürich / Switzerland. Forest Research Institute, Kepong, Kuala Lumpur / Malaysia. Forest Research Centre, Sandakan, Sabah / Malaysia. Hessisches Landesmuseum Darmstadt Hessen / Germany. Hungarian Natural History Museum, Budapest / Hungary. Institut Royal des Sciences Naturelles de Belgique, Brüssel / Belgium. Institute of Zoology, Chinese Academy of Sciences, Beijing / China. Kinabalu National Park Collection, Sabah / Malaysia.



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MCSN MHNG MLUH MNCN MNHN MNHU MRSN MUME NHMB NHMW NHRS NZSI OXUM RMNH SFDK SIES SMFM SMSM SMTD SMNS SNMB UPPC UKM ZMAS ZMPA ZMUH ZMUK ZRCS ZSMC FH CLC FSC JB MG MH OC OZ PEB HT AT NT PT ST LT PLT

Museo Civico di Storia Naturale „Giacomo Doria“, Genua / Italy. Muséum d’Histoire Naturelle, Genève / Switzerland. Wissenschaftsbereich Zoologie, Martin-Luther-Universität, Halle (Saale) / Germany. Museo Nacional de Ciencas Naturales, Madrid / Spain. Muséum National d´Histoire Naturelle, Paris / France. Museum für Naturkunde der Humboldt-Universität, Berlin / Germany. Museo Regionale di Scienze Naturali, Torino / Italy. Manchester Museum, University of Manchester, Manchester / England. Naturhistorisches Museum, Basel / Switzerland. Naturhistorisches Museum, Vienna / Austria. Naturhistoriska Riksmuseet, Stockholm / Sweden. National Zoological Survey of India, Calcutta / India. Oxford University Museum, Oxford / England. Rijksmuseum van Natuurlijke Historie, Leiden / Netherlands. Sarawak Forestry Department, Kuching, Sarawak / Malaysia. Shanghai Institute of Entomology, Academia Sinica, Shanghai / China. Senckenbergmuseum, Frankfurt a. M. / Germany. Sarawak Museum, Kuching, Sarawak / Malaysia. Staatliches Museum für Tierkunde, Dresden / Germany. Staatliches Museum für Naturkunde, Stuttgart / Germany. Staatliches Naturhistorisches Museum, Braunschweig / Germany. Museum of Natural History, University of the Philippines, Los Baños, Laguna / Philippines. Centre for Insect Systematics, Bangi / Malaysia. Zoological Museum, Academy of Science, St. Petersburg / Russia. Polish Academy of Sciences, Warshaw / Poland. Zoologisches Museum und Institut, Hamburg / Germany. Zoologisches Museum der Christian-Albrechts-Universität, Kiel / Germany. Zoological Reference Collection, National University of Singapore / Singapore. Zoologische Staatsammlung München, Munich / Germany. Private collection of Frank H. Hennemann, Kaiserslautern / Germany. Private collection of C. L. Chan, Kota Kinabalu / Malaysia. Private collection of Francis Seow-Choen / Singapore. Private collection of Joachim Bresseel, Meise / Belgium Private collection of Marco Gottardo, Rovigo / Italy. Private collection of Mel Herbert, Hütschenhausen / Germany. Private collection of Oskar V. Conle, Bolsterlang / Germany. Private collection of Oliver Zompro, Kiel / Germany (affiliated with ZMUK). Private collection of Phil E. Bragg, Nottingham / England. Holotype Allotype Neotype Paratype Syntype Lectotype Paralectotype


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3 Basic background 3.1 The present classification Stål (1875) divided the Phasmatodea into two large groups according to whether the insects possess an “areola apicali tibiarum” (= area apicalis) or not, but did not name these groups. In their well known monograph Brunner v. Wattenwyl (1907) & Redtenbacher (1906, 1908) termed these groups “Areolatae” and “Anareolatae” and arranged the order in a number of equally placed “tribes”, which were mainly differentiated by the length of the median segment and its proportion to the metanotum. Karny (1923) united these “tribes” to a few subfamilies and recognized the Areolatae and Anareolatae as families and renamed them Phylliidae and Phasmatidae. Redtenbacher (1908: 436) who worked out the Anareolate (= Phasmatidae Gray sensu Karny, 1923; = Phasmidae Gray sensu Günther, 1953), which included Acrophyllini, characterized this tribe by the triangular cross-section of the profemora in which the anterodorsal carina is distinctly raised and serrate, while the posterodorsal carina is strongly reduced, as well as the elongate and broadened cerci of certain genera. Redtenbacher’s Acrophyllini is however an artificial group and was subsequently raised to subfamily level by Karny (1923: 240) who used the name Phasminae. Bradley & Galil (1977) corrected the spelling into Phasmatinae. Various authors have already shown Phasmatinae sensu Bradley & Galil as well as the suborder “Anareolatae” to be polyphyletic (Bradler, 2001; Tilgner, 2002; Whiting et al., 2003). The next basic changes on the order’s classification were undertaken by Günther (1953), who provided keys to all subfamilies and most of the newly established tribes. Günther accepted Karny’s families Phylliidae and Phasmidae and sub-divided the subfamily Phasminae sensu Karny, 1923 into seven tribes. Furthermore, Günther (1953: 554) was the first author to recognize the feature that separates great parts of his Phasminae (= Phasmatinae Bradley & Galil, 1977) from the other subfamilies of Phasmidae: the split anal segment and reduced vomer of the %%. The seven tribes that Günther (1953) placed in Phasminae (= Phasmatinae Gray, 1835) are: Phasmini (= Phasmatini Gray, 1835), Stephanacridini, Acanthomimini, Achriopterini, Pharnaciini, Baculini (= Clitumnini Brunner v. Wattenwyl, 1907) and Macracanthini (= Acanthoxylini Bradley & Galil, 1977). Günther (1953: 555) himself had recognized that his Phasminae might be polyphyletic and wrote: “Von diesen Tribus sind wahrscheinlich manche keine natürlichen Einheiten oder noch zu weit gefaßt. Besonders die Pharnaciini und die Baculini würden vielleicht weiter, und unter Berücksichtigung des Baues der Analsegmente der %% auch anders aufzuteilen sein.“[Some of these tribes may not represent natural groups and are probably still too broadly established. Especially Pharnaciini and Baculini may have to be arranged differently, if the morphology of the anal segments of the %% is taken into account.]. Although Günther mentioned the split anal segment of %% to be a characteristic feature of his Phasminae, he did not recognize that certain genera which he included in Pharnaciini and Macracanthini (= Acanthoxylini) as well as the tribes Achriopterini and Stephanacridini lack this feature. Actually, it is only present in Günther’s Phasmini (= Phasmatini Bradley & Galil, 1977), Baculini (= Clitumnini) and certain genera of Pharnaciini. Bradley & Galil (1977) presented the most current and widely accepted arrangement of the Phasmatodea but generally translated Günther’s 1953 publication into English. These authors also took over Günther’s seven tribes of Phasminae. Taxonomic changes concerning taxa discussed here were limited to correcting Günther’s spelling of Phasminae into Phasmatinae and re-naming his tribes Phasmini and Macracanthini to Phasmatini and Acanthoxylini. The keys to the subfamilies and tribes were mostly based on literature sources and do not contribute any new features. Instead the authors re-used the key features that had already been established by Redtenbacher (1908) and Günther (1953). Bradley & Galil (1977: 186) characterized Phasmatinae as follows: “Anterior femora approximately triangular in cross-section, serrate at least dorsobasally; either with wing rudiments or with mesonotum longer than metanotum; [...], and anal segment of male cleft”.



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The Pharnaciini they distinguished by the following combination of characters (Bradley & Galil, 1977: 192): “Both sexes or at least female, apterous; Male alate; median segment of female almost or completely as long as metanotum”. Günther’s (1953: 554) comments on the genital morphology of %% in Phasmatinae have remained mostly unrecognized by subsequent authors and it was until Bradler (2001) that Günther’s suggestions were picked up again and examined in more detail. Bradler (2001) showed that the genitalia of %% in particular are indeed of great importance for the taxonomic arrangement of Phasmatinae and related subgroubs of Old World Phasmatodea. Based on morphological features, such as laterally flattened and leaf-like or lanceolate cerci, a tectiform anal segment (= abdominal tergite X) and reduced vomer (= a sclerotised derivate of sternum X) of %% this author recognized a supposedly monophyletic group of phasmatodeans in the Australian Region, the Lanceocercata. According to Bradler (2001: 183) the following taxa belong in Lanceocercata: Tropidoderinae, Xeroderinae, sections of Phasmatinae (tribes Phasmatini and Acanthoxylini), sections of Pachymorphinae (tribe Pachymorphini) and the subfamily Platycraninae Redtenbacher, 1908, except its type genus Platycrana Gray, 1835. This shows several groups traditionally referred to as subfamilies, e. g. Phasmatinae and Pachymorphinae, to be polyphyletic and is generally supported by subsequent cladistic and molecular studies (Tilgner, 2002; Whiting et al., 2003). The generally presumed monophyly of Bradler’s Lanceocercata clearly excludes two tribes from former Phasmatinae, since they do not show foliaceous cerci nor the typical anal segment of %%, namely the tribes Pharnaciini and Clitumnini (= Baculini sensu Bradley & Galil, 1977). These two tribes are apparently closely related to each other and obviously represent a subfamiliar unit distinct from what actually must be regarded as Phasmatinae. Furthermore, Lanceocercata and mentioned recent cladistic and molecular studies claim Tropidoderinae and Phasmatinae to be closely related, but so far no detailed study, comprehensive comparison or new arrangement with updated diagnoses of the taxa included in Lanceocercata has been conducted. Some tribes (e.g. Pharnaciini and Clitumnini) are presently of unknown systematic position within the family Phasmatidae Gray and require clarification. Clark-Sellick published three extensive papers dealing with the eggs of Phasmatodea (1997a, 1997b, 1998). In the third paper (Clark-Sellick, 1998) the author studied the shape and structure of the micropylar plate and discussed its importance for the systematics of the order. This publication lists numerous interesting observations and suggestions but only a few conclusions relevant for the classification were drawn. Clark-Sellick (1998: 226) had already noted that the plate and capsule morphology within certain groups of former Phasmatinae, e.g. Pharnaciini, showed so much variation that it could be considered polyphyletic.

3.2 Characters and evolutionary trends All former arrangements of the Phasmatodea in which the winged forms are strictly separated from the apterous forms are a long way from a classification based on natural relationships, and it is obviously seen that close phylogenetic relationships exist between the winged and the apterous forms. As phasmids are masters of camouflage many external structures are merely adaptations and subject to considerable variation, thus being of doubtul systematic value. Not closely related but sympatric taxa are often seen to show convergent adaptions caused by similar ecological pressure in their particular habitat. Reconstruction of the phasmatodean ground plan certainly still deserves more research and confirmed determination of the sister group of Phasmatodea. While significant progress was recently achieved in the basal phylogeny of recent Phasmatodea (Bradler, 2003; Klug & Bradler, 2006), the phylogenetic position of the crown-group Phasmatodea is still very controversial (Delclòs et al., 2008: 370). So, opinions concerning the sister group of Phasmatodea still contradict amongst different authors, but Orthoptera and Embioptera are generally considered the most likely candidates (Kristensen, 1975). Rähle (1970), Tilgner (2001), Whiting et al. (2003) and Terry & Whiting (2005)


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interpreted Embioptera as the sister group, while e.g. Wheeler et al. (2001) and Gorochov (2001) came to the conclusion that Orthoptera must be the sister taxon of the Phasmatodea. Terry & Whiting (2005: 249) introduced the term Eukinolabia to contain Phasmatodea and Embioptera. Zompro (2004a) recognized the Phasmatodea as the sister-taxon of one group “Timetatodea” sister to the Embioptera, together representing the sister-group of the Plecoptera, and the whole set [Plecoptera + (Timetatodea + Embioptera)]. Based on sperm structures Dallai et al. (2005) considered the Phasmatodea to be the sister-taxon of all other Polyneoptera. The basal diversification of the Phasmatodea still remains debatable (Bradler et al., 2003) although significant progress was achieved recently. Within the order Phasmatodea Kristensen (1975), Tilgner et al. (1999); Tilgner (2001), Wheeler et al. (2001), Bradler (2000 & 2003) and Whiting et al. (2003) all interpreted Timema as the sister taxon of all remaining Phasmatodea, the Euphasmatodea. Based on a cladistic analysis of morphological features of the adult && and eggs Tilgner (2001) has presented reasonable evidence that the monophyletic Old World Aschiphasmatinae represent the most basal branch of extant Euphasmatodea. This contradicts to Bradler (2000 & 2003) who, due to supposed plesimorphic characters such as the orthognathous head and short thorax and abdominal segments with longitudinal musculature spanning the entire segments as in Timema, interpreted the wingless highly mountainous Neotropical genus Agathemera Stål, 1875 as the basalmost branch of Euphasmatodea (Bradler et al., 2003). Subsequent examination has however shown at least the elongated longitudinal musculature to be not the plesiomorphic condition in Phasmatodea, but might be a result of secondary elongation of the external ventral longitudinal muscles (see Klug & Bradler, 2005). Zompro (2004a) raised Timema to ordinal rank, interpreted it as the sister group to all Phasmatodea, and recognized Agathemera as the sister taxon to all remaining Euphasmatodea (= Phasmatodea sensu Zompro), as did Bradler et al. (2003). However, a paraphyly of Phasmatodea (= Timematodea + Euphasmatodea) as proposed by Zompro (2004a) is clearly contradicting to the results of Whiting et al. (2003), and Terry & Whiting (2005). Area apicalis To date, the suborders “Areolatae” and “Anareolatae” of Phasmatodea have been based on the presence or absence of an impressed area on the apices of the tibiae, the area apicalis (→ see above). A similar structure is however part of the ground pattern of Timema, the sister-group of Euphasmatodea (Tilgner et al., 1999; Tilgner, 2001; Bradler, 2000 & 2003; Whiting et al., 2003), and Embioptera, a closely related order of lower Neoptera and supposed sister-taxon of the Phasmatodea (see Rähle, 1970; Kristensen, 1975; Tilgner, 2001; Whiting et al., 2003; Terry & Whiting, 2005). This indicates that the presence of an area apicalis might also be in the ground pattern of Phasmatodea (Zompro, 2004a) and can thus not hold to distinguish two distinct “suborders” within the Phasmatodea or Euphasmatodea respectively. In fact, the area apicalis rather appears to have been reduced at least twice within the Phasmatodea, why its lack can be regarded as apomorphic (Bradler, 1999: 43, 46). The plesiomorphy of the area apicalis for the Phasmatodea however still needs further evaluation and clarification of its function. For instance Baccetti (1987) has, based on sperm ultrastructure, concluded the areolate genus Bacillus St. Fargeau & Audinet-Serville, 1825 is closely related to three taxa of “Anareolatae” that were placed in Phasmatinae, Lonchodinae and Pachymorphinae by Bradley & Galil (1977). Bradler (1999) supported this assumption by the presence of a gula, which is not part of the phasmatodean ground plan (Kristensen, 1975), in more derived representatives of both the “Areolatae” and “Anareolatae”. As a result, “Anareolatae” are regarded either paraphyletic or polyphyletic (e.g. Kristensen, 1975; Baccetti, 1987; Bradler, 1999; Whiting et al., 2003). Vomer The vomer is a hook-shaped or papillate, sclerotised derivative of abdominal sternum X of %% (Pantel, 1915) that clasps the posterior margin of sternum VII of the & during copulation, by being inserted into a special bend, hole, slit or appendage on sternum VII, the so-called praeopercular organ, or it is inserted laterally



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into the & ovipositor. Key (1974 & 1991) interpreted the vomer as a family feature and suggested re-establishing the taxon Phylliidae Günther, 1953 based on the presence of a well sclerotised vomer, instead of having an area apicalis. According to this author the vomer occurs in all “Areolatae”, Necrosciinae and in certain taxa of Lonchodinae and Bacteriinae (= Diapheromerinae, = Cladomorphinae). However, the vomer is much more widely distributed amongst Phasmatodea and occurs in almost all subgroups. Certain rather basal and predominantly Australian Euphasmatodea lack an external vomer and were stated to form a monophyletic group, the Lanceocercata (Bradler, 2001: 181). The only other Euphasmatodea that lack an external vomer are most Old World Lonchodinae and Clitumninae, as well as certain genera of the New World Diapheromerinae: Diapheromerini Kirby, 1904. In all these cases a rudimentary internal vomer is present and the external vomer is functionally replaced by elongated hook-like cerci to clasp the && abdomen (sections of Diapheromerinae: Diapheromerini), or an anal segment split into two movable and internally dentate semi-tergites, which enable the % to hold fast to appendages of the & praeopercular organ on sternum VII (Clitumninae and sections of Lonchodinae) or a keel of the subgenital plate (Lanceocercata). Bradler (1999) has shown a sclerotised vomer to be present in Timema, the widely accepted sister-group of the Euphasmatodea, and hence concluded it must have already been present in the common stem species of all Phasmatodea. Consequently, the wide distribution of the vomer amongst Phasmatodea and presence in almost all subgroups of former “Areolatae” and “Anareolatae” indicate the vomer is part of the phasmatodean ground plan, hence representing a plesiomorphy of all its taxa (Bradler, 1999: 44). Wings Phasmatodea are generally believed to have lost their wings during the course of their evolution, and indeed thousands of independent transitions from full-sized wings (macroptery) to winglessness have occured in the evolution of insects. An evolutionary reversal had never been demonstrated and was generally considered to be very unlikely because functional wings require complex interactions among multiple structures, and the associated genes would be free to accumulate mutations in wingless lineages, effectively blocking the path for any future wing reacquisition (Whiting et al., 2003: 266). Whiting et al. (2003) proposed a new phylogeny for the Phasmatodea, based on molecular data obtained fom 18S rDNA, 28S rDNA and histone 3. In contrast to all former assumptions the result hypothesizes an apterous stem species for the whole Phasmatodea and a secondary recovery of wings at least four times independently in different subgroups. These authors emphasized the detailed homology in the wing features shared among phasmids and other insects and suggested that wings did not re-evolve de novo in Phasmatodea, but rather represented a “re-expression” of the basic wing which was lost in ancestral stick-insects and that the wing development pathways are conserved in wingless taxa (Whiting et al., 2003: 266). They based this assumption on the fact that, in Drosophila (Diptera) and other insects, leg and wing imaginal discs have a common origin from a single group of cells and hence suggested the basic genetic instructions for wing formation were possibly conserved in wingless (apterous) Phasmatodea. Their hypothesis that the ancestral condition in Phasmatodea is apterous and that wings were derived secondarily is supported by a review of fossil Phasmatodea (Tilgner, 2001), a cladistic study of morphological features (Tilgner, 2002) and other studies on the basal diversification of Phasmatodea (e.g. Bradler, 1999 & 2003; Zompro, 2004a), which have all shown the most basal branches of recent Phasmatodea to be wingless. However, the newly proposed phylogeny of Whiting et al. (2003) itself, which suggests several clades of presumably non-monophyletic “Diapheromerinae” to form the basal lineages of extant Euphasmatodea, surely needs to be recognized with caution. Trueman et al. (2004: 138) argued that Whiting et al. (2003) had overstated significantly the probability of wing re-evolution in stick-insects and that reconstruction of the phasmid ancestor was not the relevant issue. Using easy math Trueman et al. (2004) stated the ratio of wing loss to gain was 2.5:1 (or 2:1 if nodes are “alternatively resolved” by an ad hoc rearrangement of taxa to minimize character transformations), a hypothesis rejected by Whiting & Whiting (2004). Zompro (2004a: 323)


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as well doubted the result of Whiting et al. (2003) and argumented that it partially sufferd from the unfavourable selection of mainly apterous species or genera from taxa in which winged and apterous forms coexist, which seems to have been influenced by availability rather than necessity. Furthermore, Zompro (2004a: 323) stated the presented tree to be misinterpreted and pointed out fossil records of Phasmatodea to reveal exclusively winged taxa. However, although generally resembling winged extant Phasmatodea the true relationships of many winged fossils of Phasmatodea-like Neoptera with extant Phasmatodea are still enigmatic and doubted by various authors (e.g. Kristensen, 1975; Gorochov, 1994; Tilgner, 2001; Nel et al., 2004; Delclòs et al., 2008). The oldest documented Euphasmatodea fossils date to 44–49 million years ago (Lutz, 1990; Tilgner, 2001), one compression fossil from the Eocene of Grube Messel (Darmstadt, Germany) having a strikingly Euphasmatodea-like habitus with half-sized forewings. Several fossil insects from the Tertiary strongly resemble Recent Phasmatodea in various aspects e.g. the families Aeroplanidae Tillyard, 1918 from the upper and lower Triassic of Australia and Russia, and Hagiphasmatidae Ren, 1997 from the the late Jurassic (Ren, 1997) or upper Cretaceous (Barrett, 2000) of China respectively. Indeed, representatives of the Tertiary Hagiphasmatidae show obvious similarity to some of the very basal extant Euphasmatodea (e.g. Podacanthus Gray, 1833; Phasmtidae: Tropidoderinae), having long alae, a rather short mesothorax, a fused median segment, anareolate tibiae, 5-segmented tarsi, lanceolate not segmented cerci and in && a subgenital plate that projects over the apex of the anal segment. However, there appear to be differences concerning the wing venation and the organization of the cubito-median veins in particular between these and extant Euphasmatodea (Tilgner, 2001; Nel et al., 2004). Hence, and due to %% of Hagiphasmatidae appear to lack a vomer, an autapomorphy of the Phasmatodea, the phylogenetic relationship of these Tertiary orthopteroid insects with Recent and Mesozoic Phasmatodea still remains uncertain (Tilgner, 2001). The taxonomic placement of the enigmatic Mesozoic insect taxon Chresmodidae Handlirsch, 1906 was subject of many debates and dramatically reinterpreted several times. Handlirsch (1906–1908) originally discussed its inclusion in either Phasmatodea or Orthoptera and subsequently (Handlirsch, 1925; 1926–1930) considered them as aqatic phasmatodeans. Also Martynov (1928) and Sharov (1968) included Chresmodidae in the Phasmatodea. But based on a detailed restudy of the complete venation of both fore- and hind wings Delclòs et al. (2008) showed the “water-strider-like” Chresmodidae to belong in the Archaeorthoptera (= Orthopterida), thus in the orthopteroid lineage rather than the crown-group Phasmatodea. The widely supposed wingless sister taxon of the Euphasmatodea, namely Timema, the wingless Agathemera as the most basal branch amongst extant Euphasmatodea, and Embioptera as the possible sister group of Phasmatodea (see Kristensen, 1975; Terry & Whiting, 2005) however all support the main hypothesis of Whiting et al., (2003: 267) that wing loss does not seem to be a an evolutionary dead end in the Phasmatodea which also does not principally exclude the possibility of a secondary loss. Zompro (2004a) erroneously homologized the posterior mesonotal lobes of Agathemera as rudiments of alae (possibly meaning tegmina). Camousseight (1995) has however shown that these lobes are not homologous with tegmina, since they are positioned dorsally on the hind margin of the mesonotum and lack venation or articulation. The fact that Agathemera is truely apterous and considered the most basal lineage of Euphasmatodea supports the hypothesis of wing recovery in phasmid evolution (Bradler et al., 2003: 133). The fact that certain species exist which occur in a fully winged and brachypterous or completely apterous form (e.g. Asceles margaritatus Redtenbacher, 1908 in Borneo & Rhamphosipyloidea philippa Redtenbacher, 1908 in the Philippines, both currently in Necrosciinae), in particular shows that wings can be lost or gained very rapidly. There are several genera amongst various subgroups of Euphasmatodea which possess considerable intrageneric variation relating to the length of the alae, e.g. Lopaphus Westwood, 1859 (Necrosciinae), Malacomorpha Rehn, 1906 (Pseudophasmatinae) or && of Phyllium Illiger, 1798 (Phylliinae). For instance, Phobaeticus Brunner v. Wattenwyl, 1907 (Clitumninae: Pharnaciini) contains fully alate %%, ones with considerably shortened alae and %% that are completely apterous.



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Certainly, more research is necessary for reconstruction of the phasmatodean ground plan but to date it is considered to contain at least the following features (according to: Kristensen, 1975; Clark-Sellick, 1998; Bradler, 1999; Tilgner et al., 1999; Tilgner, 2001; Zompro, 2004a): 1. Mesothorax short, hardly longer than the pronotum 2. Pronotum with a pair of defensive glands 3. Abdominal segment I (median segment) fused with the metanotum 4. Ten abdominal segments 5. Sclerotised internal and external vomer present in %% 6. && with a praeopercular organ on abdominal sternum VII 7. Cerci consisting of one article 8. Costal vein of tegmina abbreviated 9. Area apicalis present on all tibiae 10. Eggs with a micropylar plate system 11. Spermatozoa lacking mitochondria Several evolutionary trends are frequently observed independently in various subgroups of extant Euphasmatodea, e.g. an increasing elongation of the mesothorax, a reduction of the area apicalis, and a reduction or recovery of the wings. Such features are generally regarded as apomorphic for the Euphasmatodea, although the fact wether the wings were reduced or recovered is still controversely discussed (see Whiting et al., 2003; Trueman et al., 2004; Whiting & Whiting, 2004). Numerous examples prove the presence or absence of ocelli is always in correspondence with a presence or absence of wings, alate taxa capable of active flight usually having evolved ocelli, while apterous taxa lack these additional “flight eyes”. A sclerotised vomer is present in the ground plan of Phasmatodea, but several subgroups appear to have reduced it secondarily (e.g. Lanceocercata, Lonchodinae & Clitumninae). According to Zompro (2004a: 24) the vomer consists of an internal and an external part, but in fact only the sclerotised, usually triangularly hook-like external part was originally referred to as the vomer by Pantel (1915). Hence, the internal part cannot deserve the term “vomer” but is still traced as a thick, desclerotized structure concealed by sternum IX in taxa lacking the sclerotised external vomer (Zompro, 2004a: 24). The area apicalis is believed to have been reduced secondarily at least twice during the course of the evolution of Phasmatodea (Bradler, 1999: 43). Features such as the specialized anal segments of %% of certain taxa (e.g. Lanceocercata, Clitumninae and Lonchodinae), which are longitudinally split and form two interiorly dentate semi-tergites to replace the function of the reduced external vomer, or enlarged cerci of the australasian Lanceocercata are believed to be apomorphic (Bradler, 2001). This would in correspondence with the %% genitalia also concern to the lack of a praeopercular organ on sternum VII of && of the mentioned groups.

3.3 Ootaxonomy The morphology of the eggs is often less variable than the that of the insects and exhibits numerous characters that serve very well for the distinction of genera or species. Kaup (1871a) was the first author who drew attention to the eggs of phasmids and provided descriptions and illustrations of several eggs that he had extracted from the abdomens of preserved specimens in his collection (now in HLDH). Kaup (1871a: 2) stated: “Vielleicht wird man später die Arten durch die Eier schneller unterscheiden lernen als durch die Thiere selbst…[Perhaps the eggs will one day allow distinction between species more easily, than by examination of the insects themselves…]. Unfortunately, Kaup’s important publication and correct suggestions were ignored for more than 100 years. It was more than a century later, that Clark (1976, 1979; later referred to as Clark-


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Sellick) standardized the terminology for describing egg structures. Several of Clark-Sellick’s subsequent publications (e.g. 1997a, 1997b & 1998) drew attention to the relevance of egg structures for the taxonomy and classification of the Phasmatodea, but the author rarely drew conclusions from his observations. The first papers in which the eggs were strictly considered in the higher classification were published by Zompro (2001a, 2004a). Evolutionary trends of the egg morphology are however to-date still difficult to define. Clark-Sellick (1998) discussed the micropylar plate of the phasmid egg and provided a survey of the range of plate form within the order. He drew particular attention to the shape and structure of the internal micropylar plate. Clark-Sellick distinguished four distinct types of internal plates, based on combinations of either being “open” or “closed” and having or lacking a median line. Closed internal plates (Figs. 60–67) have the micropylar cup and micropylar stalk placed within the boarders of the plate, which has no gap posteriorly. Closed plates never possess a median line. Open internal plates (Figs. 48–59) have a posterior gap or slit, which opens towards the micropylar cup and micropylar stalk. This is usually displaced towards the posterior end of the plate and not fully included within the plate itself. Eggs with open internal plates may or may not have a median line. In discussing the importance of the internal micropylar plate for the classification of Phasmatodea, ClarkSellick (1998: 204) stated “Although all three types [of internal plates] may occur within one family, it is frequently true that an individual tribe has a uniform type of plate”. Thus, Clark-Sellick interpreted an open or closed internal plate as a character valid for tribal distinction of taxa. Although the hypothesis of Clark-Sellick is well supported, the value of the micropylar plate for the higher classification of the Phasmatodea is still unknown. In several cases the type of internal micropylar plate is seen to be uniform even within entire subfamilies currently believed to be monophyletic (e.g. Aschiphasmatinae, Heteropteryginae and Pseudophasmatinae).

4 Family Phasmatidae Gray, 1835 s. str. (= Lanceocercata Bradler, 2001) Phasmatidae Gray, 1835 sensu stricto Type genus: Phasma Lichtenstein, 1796: 77. Phasmidae Gray, 1835: 12. Phasmidae Karny, 1923: 235. Günther, 1953: 546. Phasmatidae, Bradley & Galil, 1977: 186. [Corrected spelling of Phasmidae Karny, 1923: 235] Otte & Brock, 2005: 31. Lanceocercata Bradler, 2001: 179. syn. nov.

Sensu Bradley & Galil (1977: 180) the family Phasmatidae Gray, 1835 contained six distinct subfamilies, five of them distributed througout the Old World (Phasmatinae, Tropidoderinae, Xeroderinae, Platycraninae and Eurycanthinae) and one, the Cladomorphinae, inhabiting the Neotropical Region. Close relationship between the latter and the five Old World subfamilies is rather unlikely (→ 4.1). Apparently, the Phasmatidae as interpreted by Bradly & Galil (1977) form an artificial group, which is not only seen in containing a single not closely related Neotropical taxon, but also in several morphological features of the exosceleton of the insects such as the genitalia, structure of the profemora or antennae. Based on morphological examination of the genital exosceleton and copulation habits Bradler (2001) recognized a predominantly Australian and supposedly monophyletic group of anareolate phasmatodeans he termed Lanceocercata. Subsequent, cladistic studies (Tilgner, 2002) and molecular studies (Whiting et al., 2003) support the monophyly of this group. Bradler (2001: 183) interpreted the rather basal ground pattern of Lanceocercata, like the just slightly prognathous head, presence of ocelli, short thorax and long tegmina, as an REVISION OF PHARNACIINI


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indication that this group must have splitted off rather early from the remaing Phasmatodea. According to Bradler (2001: 183), Lanceocercata includes the subfamilies Tropidoderinae and Xeroderinae, sections of Phasmatinae (tribes Phasmatini and Acanthoxylini) and Pachymorphinae (tribe Pachymorphini), as well as the subfamily Platycraninae, except its type genus Platycrana Gray, 1835. Consequently, large traditional groups like the subfamilies Phasmatinae, Pachymorphinae and Platycraninae are currently believed to be polyphyletic (Bradler, 2001 & 2003). Bradler (2001) based the monophyly of Lanceocercata on the following five supposedly apomorphic characters: 1. Cerci strongly laterally flattened and ± enlarged, foliaceous or lanceolate. 2. Vomer of %% strongly reduced and concealed by sternum XI (= paraproct). Functionally replaced by a clasping apparatus of the anal segment (= tergite X). 3. Anal segment (= tergum X) of %% specialized and forming a clasping apparatus to hold fast to the & abdomen during copulation. Tergite strongly tectiform, longitudinally split and forming two flexible semi-tergites, only connected by a thin membrane dorsally. Interior surfaces of anal semi-tergites posteroventrally armed with a variable number of distinct incurving teeth (= “Dornenfeld”). 4. Subgenital plate of && (= sternum VIII) with a prominent, ± ledge-like longitudinal keel in basal portion. 5. Derived copulation position. By making use of the clasping apparatus of their anal segment %% grasp the & along the longitudinal keel of the subgenital plate (= sternum VIII). Detailed examination and extensive comparison of genital structures and other morphological characters (e.g. structure of the profemora and antennae) or copulation habits here undertaken amongst the concerned taxa and other related subgroups, support a monophyly of Lanceocercata. Certain subfamilies included in the Lanceocercata by Bradler (e.g. Xeroderinae and Pachymorphini) however are rather insufficiently defined and most certainly artificial groups, which clearly warrant better recognition (→ 4.6). This is in particular indicated by containing very differently structured types of genitalia, some of which do not at all match with the characteristics listed by Bradler. Indeed however, the five supposedly apomorphic features that Bradler (2001) listed to characterize Lanceocercata are true for almost all of the taxa originally included, and apparently such correlating specialisations are rather unlikely to have been evolved independently in separate subgroups, thus supporting Bradler’s hypothesis. The concerned features and their importance for the arrangement of Old World Phasmatodea are discussed in some more detail below. 1) The conspicuous lanceolate or foliaceous cerci are frequently found throughout anareolatae Old World Phasmatodea but can with quite some certainty be put down to a single genealogical lineage, which must have separated from the stem of Phasmatodea rather early in the evolution of the order (Bradler, 2001: 183). Laterally compressed cerci are also found in other subgroups, but they are never structured in the typical manner of Lanceocercata. In Lanceocercata they are usually formed by flattened, leaf-like extensions of the dorsal and ventral surfaces and the apex of the cercus (Figs. 2–19). They vary considerably in size and shape amongst the taxa of Lanceocercata and even show variation within individual tribes here interpreted as monophyletic (e.g. Phasmatini & Monandropterini). The enlarged and specialized cerci do not serve as forceps to grasp the & abdomen during copulation like, for instance, in certain members of the New World Diapheromerinae: Diapheromerini Kirby, 1904 (e.g. Diapheromera Gray, 1835 or Pseudosermyle Rehn, 1906) but they do in certain genera of Phasmatinae and Tropidoderinae have the basal portion set with minute denticles. During copulation these serve for the % as an additional anchorage to hold fast by being pressed tightly against the lateral surfaces of the & subgenital plate (Figs. 432 & 434). The conspicuous dorsal tooth of the cerci in %% of Monandropterini and Gigantophasmatini trib nov. (subfamily Tropidoderinae) may have a similar function (Figs. 11–16). 2) Bradler (2001: 181) stated the vomer of %% to be completely extinct in Lanceocercata which however


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only concerns to the external part of that organ (Figs. 38, 39 & 42; → 3.2). The vomer is part of the phasmatodean ground plan and in Phasmatodea consists of two parts, an internal part and an often well sclerotised external part (Zompro, 2004: 24). The external part is often increasingly reduced secondarily in taxa which have it functionally replaced by a split anal segment that serves as a clasping apparatus to hold fast to the & abdomen during copulation. But although the external part is reduced or lacking the internal part is still present as a desclerotized structure concealed by sternum XI (= paraproct). Indeed, the vomer is not visible without internal examination and removing of the paraprocts in all of the taxa which Bradler (2001) placed in Lanceocercata. However, an increasingly reduced vomer is also true for other subgroups e.g. the related subfamilies Clitumninae and Lonchodinae (Figs. 35–36, 40–41), both of which also have it functionally replaced by a clasping apparatus formed by the anal segment. Hence, the lack of an external vomer in general can well be presumed as apomorphic but it can hardly be interpreted an autapomorphy of Lanceocercata. 3) The function of an anchorage to hold fast to the & abdomen during the copulation is replaced by a conspiuous clasping apparatus of the %% anal segment. This is of a very characteristic shape and structure and found in almost all the taxa that Bradler (2001) included in Lanceocercata. The anal segment (= tergite X) is strongly tectiform, longitudinally split and consists of two movable semi-tergites which are merely connected by a thin membrane dorsally (Figs. 2–19, 32–34). The ventro-apical angles are ± distinctly elevated and in some taxa even form a finger-like, in-curving process (e.g. Podacanthus Gray, 1833 or Parapodacanthus Brock, 2003). This processs either bears several terminal teeth, or the internal surfaces of the semi-tergites ventro-apically bear a longitudinal bulge or ledge, which is armed with a variable number of back-curving teeth (Figs. 32–34). A split anal segment which consists of two movable semi-tergites to form a clasping apparatus is also frequently found in %% of the subfamilies Clitumninae (→ 5.) and Lonchodinae (→ 5.5). These are however of a rather distinct structure, having the complete apex ± elongated, finger or spatula-like and with the interior surfaces almost entirely or at least along the outer margins set with small teeth (Figs. 35–36, 20–25, 29–31). Most members of these two subfamilies have the external vomer increasingly reduced and often lacking, but it is in some genera present as a rather small papillate or filiform but well sclerotized organ (→ 5.3). Another example for a strongly reduced external vomer and a specialized a clasping apparatus of the anal segment is the Old World subfamily Eurycanthinae (→ 4.1). As a result, it is not the split anal segment itself but the particular shape and structure of the clasping apparatus which might be regarded an autapomorphy of Lanceocercata. A split anal segment which consists of two movable semi-tergites may rather be interpreted a synapomorphy of Lannceocercata + Clitumninae + Lonchodinae + Eurycanthinae, supposing relationship between these groups. 4) Already Bradler (2001: 183) himself doubted the true apomorphic nature of the sharply keeled and boat-shaped subgenital plate of && of Lanceocercata. Indeed longitudinally keeled subgenital plates are frequently found in most other subgroups of Phasmatodea, e.g. the related Clitumninae, Eurycanthinae or sections of Lonchodinae. As Bradler stated it is a supposition for the derived copulation position of Lanceocercata in which %% grasp the longitudinal keel close to the base of the subgenital plate and true for all taxa contained in Lanceocercata. Its presence in other not closely subgroups, none of which exhibit the same conspicuous copulation habit and often have a praeopercular organ and/or a well developed external vomer in %%, suggests it must not necessarily be a specialization dependent on the copulation behaviours. It can thus hardly be an autapomorphy of Lanceocercata. 5) The derived copulation position observed in Lanceocercata is indeed remarkable and differs from all other extant Phasmatodea. In contrast to the remaining Phasmatodea, even ones without an external vomer, %% of all taxa of Lanceocercata grasp the & abdomen along the distinct longitudinal median keel of the subgenital plate (= sternum VIII), which is well supported by the conspicuous longitudinal interior row of teeth or terminal teeth on the ventro-apical elevation of the anal semi-tergites of the (Figs. 431–435). Other groups which lack an external vomer (e.g. Clitumninae or Lonchodinae) and have an anal segment that forms a clasp-



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ing apparatus have && with a ± well developed praeopercular organ on sternum VII, formed by a pair of humps, laterally compressed spines or lobes (→ 5.). These appendages of the && praeopercular organ are then grasped and used as an anchorage by the %%, the shape of the appendages usually correlating with the shape of the anal semi-tergites of the %% (Figs. 436–437). Other groups, like sections of the New World subfamily Diapheromerinae, have the vomer functionally replaced by elongated, forceps-like cerci to enable the %% to grasp the & abdomen during the copulation. In all those taxa with a sclerotised external vomer in %%, these hold fast to the & abdomen by inserting this hook-like organ into a hole, slit or special external structure (= praeopercular organ) on sternum VII of &&. Consequently, the conspicuous copulation position can obviously be regarded an autapomorphy of the Lanceocercata. Another apomorphic feature of Lanceocercata not mentioned by Bradler (2001) is the lack of a praeopercular organ in && which correlates with the lack of an external vomer in %%. The && of neither taxon of Lanceocercata have any hint of a praeopercular organ on sternum VII, hence this structure can be regarded as lacking in this particular group of Phasmatodea. The lack may be explained by the derived copulation position for which a praeopercular organ has apparently become unnecessary. A common feature of the eggs of Lanceocercata is the presence of a capitulum which however may be of various shapes and structures. All have closed capitula (for a definition see: Clark-Sellick, 1988: 275), which are either hat-like with a ± stalk, knob-like and closely appressed to the operculum, or conical without a stalk. However, despite these common and obviously homologous morphological features of the insects, two fundamentally distinct types of internal micropylar plates are observed amongst the eggs of the taxa contained in Lanceocercata—open and closed plates. The structure of the internal micropylar plate may well serve for an arrangement of the subgroups of Lanceocercata and should be taken into account, since its type of shape (open or closed), and presence or absence of a median line, are obviously uniform within an individual tribe (ClarkSellick, 1998: 204, → 4.3). Amongst Lanceocercata the internal micropylar plate is open in sections of Phasmatinae (Phasmatini and Acanthomimini), sections of Tropidoderinae (tribes Tropidoderini and Monandropterini), but closed in Extatosomatinae, Gigantophasmatini trib. nov. (subfamily Tropidoderinae), Acanthoxylini (subfamily Phasmatinae) and “Platycraninae”. The internal plates of Pachymorphini (szubfamily Pachymorphinae) are not known. The presence of both types of plates in Xeroderinae sensu Bradley & Galil, 1977 (closed in Cotylosoma Wood-Mason, 1878 and Nisyrus Stål, 1877 but open in Dimorphodes Westwood, 1859), suggests this latter subfamily is polyphyletic or deserves splitting into two separate tribes (→ 4.6.2). As a result, the monophyly of Lanceocercata is well supported by morphological features of the genital exosceleton (see also: Tilgner, 2002) and indeed some of the features emphasized by Bradler (2001) appear to be autapomorphies of a supposedly monophyletic group. But, due to Lanceocercata contains the subfamily Phasmatinae with the subordinate taxon Phasmatini and the type genus Phasma, Lanceocercata represent what must actually be regarded the family Phasmatidae Gray, 1835 (sensu stricto). Consequently, the term “Lanceocercata” has to be interpreted as a synonym of the available family name Phasmatidae. For a list of the subfamilies and tribes contained in Phasmatidae s. str. see below.

4.1 The Phasmatidae and Phasmatinae sensu Bradley & Galil, 1977 Sensu Bradley & Galil (1977: 186) the family Phasmatidae contained six distinct subfamilies, five from the Old World Region (Phasmatinae, Tropidoderinae, Xeroderinae, Platycraninae and Eurycanthinae), and one, the Cladomorphinae, from the Neotropical Region. The more strict definition of Phasmatidae, suggested by Bradler (2001) and termed Lanceocercata by that author, excludes sections of former Phasmatinae, Xeroderinae and Platycraninae as well as the subfamilies Eurycanthinae and Cladomorphinae from the family, since they do not match with the diagnostic features like the laterally compressed, foliaceous cerci or tectiform anal


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segment of %% which forms a conspicuous clasping apparatus. Indeed, certain tribes of Phasmatinae (→ 4.3), some genera of Xeroderinae (→ 4.6.2) and the type genus of Platycraninae (→ 4.) strongly differ in genital features and do obviously not belong in Phasmatidae s. str. (= Lanceocercata). Eurycanthinae Brunner v. Wattenwyl, 1893 Relation between Eurycanthinae and Phasmatidae s. str. certainly deserves further evaluation, and as presently treated the subfamily appears to be paraphyletic (Hennemann & Conle, 2006c: 43 ff.). A review of Eurycanthinae Brunner v. Wattenwyl, 1893 with a key to the genera was provided by Zompro (2001b). The subfamily has its centre of distribution in New Guinea and surrounding islands, but eastward and southward extends as far as the Solomon Islands, New Caledonia, the Loyalty Islands or Ball’s Pyramid (near Lord Howe Island). Members are typical for a beak-like ovipositor in &&, which is formed by elongation of the anal segment and subgenital plate (Bradler, 2003; Hennemann & Conle, 2006c), but may be secondarly reduced in certain genera (e.g. Papuacocelus Hennemann & Conle, 2006 or Thaumatobactron Günther, 1929; Zompro, 2001b: 20). The eggs are characteristic for their bullet-shaped capsule, rather small and cordiform micropylar plate which is open internally, and the flat operculum which lacks a capitulum (Zompro, 2001b: 19). %% of certain more specialised members have the metafemora increasingly broadened and often heavily armed ventrally (e.g. Eurycantha Boisduval, 1835; Dryococelus Gurney, 1947 or Papuacocelus Hennemann & Conle, 2006). Although %% lack an external vomer and have the anal segment tectiform and specialized to form a clasping apparatus, neither genus has the typically structured anal segment found throughout Phasmatidae s. str. (= Lanceocercata). It usually has the posterior margin incised medially and on both sides of the incision elongated to form a short and slender ± conspicuous finger-like process which has the internal surfaces dentate. A feature analogous to Phasmatidae s. str. is the lack of a praeopercular organ on abdominal sternum VII of &&, which is not neccessary since %% lack an external vomer. However, although %% don’t have an external vomer and && lack a praeopercular organ as an anchorage for the %% during copulation, these grasp the && along the posterior margin of sternum VII, and not along the longitudinal keel of sternum VIII (= subgenital plate) as do Lanceocercata. Due to this copulation position the claspers of the %% anal segment usually leave a wide space inbetween them, to allow grasping the posterior margin of sternum VII of && on the lateral surfaces. Although the cerci may be laterally flattened in certain genera of Eurycanthinae (e.g. Eurycantha), none of them has them conspicuously foliaceous with the dorsal and ventral carinae lamellate as in members of Phasmatidae s. str. (= Lanceocercata). Hennemann & Conle (2006c: 44 ff.) have shown external features of the && genitalia and egg-morphology to indicate close relation to Old World genera currently contained in the subfamilies Lonchodinae (Phasmatidae s. l. → 5.4) and Necrosciinae (family Diapheromeridae). This also concerns to the genitalia of %%, which are rather similar in structure to those of certain Lonchodinae, and indeed some of the more basal genera of Eurycanthinae (e.g. Brachyrtacus Sharp, 1898 or Neopromachus Giglio-Tos, 1912) strikingly resemble members of Lonchodinae with a beak-like ovipositor in && (e.g. Manduria Stål, 1877). Provisionally, as Eurycanthinae obviously does not belong in Phasmatidae s. str. (= Lanceocercata), it may be regarded a subfamily of Phasmatidae sensu lato. Cladomorphinae Brunner v. Wattenwyl, 1893 The Neotropical subfamily Cladomorphinae Brunner v. Wattenwyl, 1893 is apparently not closely related to Phasmatidae s. l. which is, apart from the distribution in the Neotropics, indicated by a variety of features. The external morphology of the genitalia of both sexes differs considerably, %% having a simple anal segment and well-developed often specialized external vomer and && have a ± distinct often specialized praeopercular organ on abdominal sternum VII and long, filiform gonapophyses in the majority of genera. The eggs furthermore differ from those of Phasmatidae s. l. by having open capitula (for a definition see: Clark-Sellick, 1988: 274). The shape of the profemora, which have the medioventral carinae lamellate and considerably displaced towards the anteroventral carinae, rather resembles the Old World Clitumninae (→ 4.4) and may either be a



21

convergent or a symplesiomorphy of [Clitumninae + Cladomorphinae], in the latter case supposing relationship between these two subfamilies. However, as discussed above Cladomorphinae can not be regarded a member of Phasmatidae s. l., raising to family level appears necessary for this subfamily. Close relationship to certain subgroups of the New World Diapheromerinae: Diapherimerini is obvious. Phasmatinae sensu Bradley & Galil, 1977 Various authors have already stated the Phasmatinae sensu Bradley & Galil, 1977 as well as the complete suborder “Anareolatae” are polyphyletic (Bradler, 2000; Tilgner, 2002; Whiting et al., 2003 & Zompro, 2004a). From the seven tribes that Bradley & Galil (1977: 192) contained in Phasmatinae, namely Phasmatini, Acanthomimini, Acanthoxylini, Pharnaciini, Baculini (= Clitumnini), Stephanacridini and Achriopterini, the latter four are obviously not closely related nor members of this subfamily. This is confirmed by striking differences concerning to the morphology of the genital exosceleton amongst the tribes included and was already pointed out by Bradler (2001), who placed Phasmatini, Acanthomimini and Acanthoxylini in his supposedly monophyletic group Lanceocercata (= Phasmatidae s. str.). Indeed, only these three tribes share several common characters which readily distinguish them from the other four tribes contained. These are the laterally compressed, foliaceous cerci, strongly tectiform anal segment of %% which forms a conspicuous clasping apparatus, lack of a praeopercular organ in && and the derived copulation position in which %% grasp the base of the && abdominal sternum VIII (= subgenital plate) instead of sternum VII (→ 4.). A specialized anal segment which consists of two movable semi-tergites is also found in %% of Pharnaciini and Clitumnini, which just like the mentioned three tribes have the external vomer strongly reduced. However, these two tribes lack the foliaceous cerci, have the semi-tergites of the anal segment structured rather differently from those of Phasmatini, Acanthomimini and Acanthoxylini, and && possess a more or less distinct often specialized praeopercular organ on sternum VII, used as an anchorage during the copulation by %% (→ 5.). Moreover, the profemora of Pharnaciini and Clitumnini differ considerably from those of the mentioned three tribes by having the medioventral carina very prominent, lamellate or ledge-like and strongly displaced towards the anteroventral carina (→ Figs. 46–47). Undoubtedly, Pharnaciini and Clitumnini are very closely related and together belong in a separate subfamily, the Clitumninae (→ 5.). The mentioned features indicate relationship between Clitumninae and Lonchodinae (→ 5.). The tribes Stephanacridini and Achriopterini were shown to be closely related to each other (Hennemann & Conle, 2004) and differ fundamentally from all other tribes contained in Phasmatinae by Bradley & Galil (1977: 192) in the morphology of the genital exosceleton (→ 4.7). %% of both tribes have a simple anal segment which consists of a single dorsal plate and have a well developed and sclerotised, hook-like external vomer. && of Stephanacridini furthermore have long, filiform gonapophyses which extend well beyond the apex of the abdomen. Furthermore, both sexes lack conspicuously foliacous cerci, have them small and cylindrical to oval in cross-section, and can hence not belong in Phasmatinae or Phasmatidae s. str. (= Lanceocercata). As discussed above only four of the six subfamilies that were included in Phasmatidae by Bradley & Galil (1977) are closely related and form the monophyletic taxon Phasmatidae s. str. (= Lanceocercata), this is Phasmatinae, Tropidoderinae, Xeroderinae and “Platycraninae”. The latter two subfamilies need further clarification since they appear to be polyphyletic (→ 4.6.2), as is Phasmatinae sensu Bradley & Galil (1977). Only the three tribes Phasmatini, Acanthomimini and Acanthoxylini form a natural group and belong in Phasmatinae. This subfamily is obviously closely related to Tropidoderinae, and based on morphological features of the genital exosceleton, together with sections of Xeroderinae and “Platycraninae” forms a monophyletic group within the family Phasmatidae sensu lato (→ 4.). The basalmost taxa of Phasmatidae s. str. (e.g. Extatosomatinae and certain Tropidoderini or Phasmatini) presumably represent some of the evolutionary oldest extant Phasmatodea, exhibiting several features plesiomorphic for the Phasmatodea, e.g. a just slightly prognathous head, ocelli, a short mesothorax, long median


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segment and abdomen, and well developed wings. In several aspects genera like Podacanthus Gray, 1833, Micropodacanthus Brock & Hasenpusch, 2007 (Tropidoderinae: Tropidoderini) or Paracyphocrania Redtenbacher, 1908 (Phasmatinae: Phasmatini) remarkably resemble fossil records of Lower Neoptera from the Tertiary supposedly belonging to Phasmatodea (e.g. Aeroplanoidea Tillyard, 1918 and Hagiphasmatidae Ren, 1997, → 3.2). However, as there appear to be differences concerning the wing venation and the organization of the cubito-median veins in particular (Nel et al., 2004) between these and Recent Phasmatodea, the phylogenetic relationship between these Tertiary orthopteroid phasmid-like insects and extant Phasmatodea still remains uncertain (Tilgner, 2001). In contrast to the hypothesis of Whiting et al. (2003) all subgroups of Phasmatidae s. str. show a clear evolution from winged to wingless forms and appear to have lost their wings during the corse of their evolution, rather than having them recovered secondarily. This is supported by various still questionable fossil records of Phasmatodea (→ 3.2), all of which are insects with well developed alae (Zompro, 2004a: 323). Evolutionary trends observed in all subgroups of Phasmatidae s. str. are a shortening of the median segment, reduction and final loss of tegmina, alae and ocelli, increasing elongation of the mesothorax, as well as a decreasing crypsis and sexual dimorphism. Distribution & biogeography Australian Region, western portion of the Pacific Islands, Wallacea, Oriental Region, southeastern Palaearctic Region as far north as Taiwan and Japan, and parts of the Indian Ocean (Seychelles, Réunion and Mauritius) (→ Fig 1). The shared presence across several of the Old World southern hemisphere landmasses suggests a common Gondwanan ancestor of Phasmatidae s. str. thus perhaps indicating an East Gondwanan origin and old evolutional age of this family. However, any broader and concrete discussion on the scenario of dispersal of Phasmatidae s. str. deserves a better understanding of the phylogeny of the Recent Phasmatodea and would be premature. Nevertheless, a short survey of some distributional patterns appear warranted. No taxa are present on the African continent and in the Neotropical Region. The taxa that retain the largest number of supposed plesiomorphies are endemic in Australia whose landmass was isolated from the rest of the world for much of the past 30 million years (Cox & Moore, 1993). The two tribes Phasmatini (subfamily Phasmatinae) and Tropidoderini (subfamily Tropidoderinae) appear to have predominantly radiated on Eucalyptus (Myrtaceae) that dominates the Australian tree flora or shrubs of the diverse genus Acacia (Fabaceae), both of which support a great variety of taxa in Australia. While the very basal Tropidoderini are endemic in Australia, several taxa of Phasmatini have dispersed northwards via New Guinea as far as to the Greater Sunda Islands (Java and Sumatra) and settled great parts of the Wallacea. The “Platycraninae” have settled great parts of the Oriental and Australian Regions extending from the Western Indian Ocean Islands Seychelles, Réunion and Mauritius as far east as Fiji and French Polynesia and appear to have radiated exclusively on palms of the families Pandanaceae (screw-palms of the genus Pandanus) and Arecaceae (e.g. the genera Chrysalidocarpus, Cocos or Trachycarpus). As for “Platycraninae” the disjunct presence of the rather basal Monandropterini (subfamily Tropidoderinae) on several of the Western Indian Ocean Islands (Réunion and Mauritius) may be explained by a fairly early settlement, perhaps during the middle or late Jurassic, via Australia or Antarctica and as well suggests a common East Gondwanan ancestor of this tribe and the Australian endemic Tropidoderini in particular. During the Pleistocene the glaciation has caused a decrease of the world-wide sea-level which developed land or island chain bridges between the Large Sunda Islands which formed one more or less united land mass, while in warmer times large areas were submerged (see Briggs, 1987; Eliot, 1992). This again may have advanced a dispersal of certain taxa in the Wallacean region as far north as the Philippines, Taiwan and even Japan. The subfamilies and tribes now contained in Phasmatidae s. str. are discussed and provisionally re-diag-



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nosed below to better characterise the groups, along with lists of the genera included in each tribe. The arrangement here proposed is meant to be a first step towards a systematization of the Old World anareolate Phasmatodea, based strictly on features of the genital exosceleton of the adult insects and egg-morphology. It is hoped that the more detailed diagnoses and new key features presented will support future studies on the higher classification of the order. Certainly, numerous questions concerning the systematic position of certain genera or true relationships between taxa are still to be answered and the subfamilies Xeroderinae, Pachymorphinae and “Platycraninae” in particular still deserve more detailed investigation. New keys to the insects and eggs of the subfamilies and tribes of Phasmatidae s. str. are provided below (→ 4.2).

Subfamilies and tribes of Phasmatidae Gray, 1835 sensu stricto (= Lanceocercata Bradler, 2001) 1. Phasmatinae Gray, 1835: 12. (→ 4.3) Tribe Phasmatini Gray, 1835: 12. Tribe Acanthomimini Günther, 1953: 555. Tribe Acanthoxylini Bradley & Galil, 1977: 192. 2. Tropidoderinae Brunner v. Wattenwyl, 1893: 97. (→ 4.4) Tribe Tropidoderini Brunner v. Wattenwyl, 1893: 97. Tribe Monandropterini Brunner v. Wattenwyl, 1893: 97. Tribe Gigantophasmatini trib. nov. 3. Extatosomatinae Clark-Sellick, 1997: 103. stat. nov. (→ 4.5) 4. Pachymorphinae Brunner v. Wattenwyl, 1893: 89 (in part). (→ 4.6.1) Tribe Pachymorphini Brunner v. Wattenwyl, 1893: 89. 5. Xeroderinae Günther, 1953: 556 (in part). (→ 4.6.2) 6. “Platycraninae” Brunner v. Wattenwyl, 1893: 97 (in part—not the type genus Platycrana Gray, 1835).

4.2 Keys to the subfamilies and tribes of Phasmatidae s. str.

&& 1. Abdomen with foliaceous lateral lobes; mesothorax strongly widened towars the posterior ......................2 - Abdomen without foliaceous lateral lobes mesothorax at best slightly widening towards posterior...........3 2. Head orthognathous; vertex flat or globose; mid and hind legs trapezoidal in cross-section, the dorsal carinae separate and not conspicuously raised ..............................................................................Xeroderinae* - Head prognathous with vertex conically elevated and spinose; mid and hind legs trigonal in cross-section with the dorsal carinae melted and together forming large, denate lobes............................ Extatosomatinae 3. Head not considerably flattened; cheeks not widened; eyes positioned laterally on head capsule; pronotum lacking distinct functional defensive glands; mesonotum not considerably flattened; cylindrical to oval in cross-section .................................................................................................................................................4 - Head ± flattened dorsoventrally; cheeks widened; eyes displaced towards dorsal surface of head capsule; pronotum with prominent defensive glands anterolaterally; mesothorax flattened and angulate laterally .... .......................................................................................................................................... ”Platycraninae”** 5. Body sub-cylindrical in cross-section, meso- and metasternum flattened; mesothorax widened towards the posterior and abdomen broadened basally or medially; profemora indistinctly triangular in cross-section, anterodorsal carina not conspicuously raised and mostly unarmed (Fig. 45).................... 6. Tropidoderinae - Body round in cross-section and of ± uniform width; profemora distinctly triangular in cross-section, anterodorsal carina strongly raised and serrate (Fig. 32); Australian Region, Wallacea & New Zealand


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(Figs. 1, 72, 73a, 75) .............................................................................................................. 8. Phasmatinae 6. Apterous........................................................................................................................................................7 - Tegmina and alae developed; Australia (Figs. 70, 71) .............................................................Tropidoderini 7. Subgenital-plate elongated, lanceolate and extending considerably over apex of abdomen; cerci very large, broadly foliaceous and much longer than anal segment; Loyalty Islands (Fig. 69a) ........................... .........................................................................................................................Gigantophasmatini trib. nov. - Subgenital plate small and scoop-shaped; cerci not conspicuously enlarged and shorter than anal segment; Indian Ocean (Figs. 74a)..................................................................................................... Monandropterini 8. Median segment longer than metanotum; Australia & Wallacea .................................................................9 - Median segment considerably shorter than metanotum, New Zealand ...................................Acanthoxylini 9. Alae longer than median segment or tegmina, usually well developed........................................Phasmatini - Alae vestigial and shorter than median segment and at best equal in length to tegmina...... Acanthomimini

%% 1. Abdomen with lateral lobes ..........................................................................................................................2 - Abdomen without lateral lobes ....................................................................................................................3 2. Head orthognathous; vertex flat or globose; mid and hind legs trapezoidal in cross-section, the dorsal carinae separate and not conspicuously raised ..............................................................................Xeroderinae* - Head prognathous with vertex conically elevated and spinose; mid and hind legs trigonal in cross-section with the dorsal carinae melted and together forming large, denate lobes............................ Extatosomatinae 3. Head not considerably flattened; cheeks not widened; eyes positioned laterally on head capsule; pronotum lacking distinct functional defensive glands; mesonotum not considerably flattened; cylindrical to oval in cross-section .................................................................................................................................................4 - Head ± flattened dorsoventrally; cheeks widened; eyes displaced towards dorsal surface of head capsule; pronotum with prominent defensive glands anterolaterally; mesothorax flattened and angulate laterally .... .......................................................................................................................................... ”Platycraninae”** 4. Medium sized to very large insects (body length 45–198 mm); antennae > 2x longer than head; scapus slender; head without a conspicuous pair of horns.......................................................................................5 - Small, rather robust insects (body length < 50 mm); antennae hardly longer than head; scapus laterally dilated; head with a pair of blunt horns or tubercles between the eyes .........................Pachymorphinae*** 5. Body sub-cylindrical in cross-section, meso- and metasternum flattened; mesothorax ± widened towards the posterior; profemora at best slightly triangular in cross-section, anterodorsal carina not conspicuously raised and unarmed (Fig. 45) ............................................................................................. 6. Tropidoderinae - Body round in cross-section and of ± uniform width; profemora distinctly triangular in cross-section, anterodorsal carina strongly raised and serrate (Fig. 44); Australian Region & Wallacea (Figs. 1, 73b) ...... ............................................................................................................................................... 8. Phasmatinae 6. Cerci foliaceous, ± oval in lateral aspect and with a ± distinct dorsal tooth; mesothorax > 1.3x longer than head and pronotum combined; anal region with ± distinct grey to blackish mottling; not Australia...........7 - Cerci lanceolate, without a dorsal tooth; mesothorax < 1.3x longer than head and pronotum combined; anal region plain transparent, pink or with pinkish mottling; Australia (Fig. 1) ......................Tropidoderini 7. Poculum roughly reaching posterior margin of abdominal tergite IX; alae projecting over abdominal segment VI; Loyalty Islands (Fig. 69b) ................................................................Gigantophasmatini trib. nov. - Poculum bulgy, almost reaching to apex of anal segment; alae at best reaching abdominal tergite V, sometimes apterous; Indian Ocean (Figs. 1, 74b) ....................................................................... Monandropterini 8. Median segment longer than metanotum; Australia & Wallacea .................................................................9 - Median segment considerably shorter than metanotum, New Zealand ...................................Acanthoxylini 9. Ocelli present; alae well developed ..............................................................................................Phasmatini



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Ocelli lacking; brachypterous ............................................................................................... Acanthomimini * Xeroderinae is here restricted to the type genus Xeroderus Gray, 1835 (→4.6.2) ** “Platycraninae” here refers to all genera contained except for the type genus Platycrana Gray, 1835, which differs considerably in genital features and consequently is not a member of the Phasmatidae (→4.1) *** Pachymorphinae here restricts to the tribe Pachymorphini (→4.6.1)

1. 2. 3. 4. -

Eggs* Micropylar plate shorter than capsule, not considerably raised above capsule surface ...............................2 Micropylar plate very elongate, reaching from opercular to posterior pole of the capsule and strongly set above capsule surface (Fig. 63) ........................................................................................... Extatosomatinae Capsule without a conspicuous dorsal ridge or keel to contain the micropylar plate; internal plate without a brown outer area ........................................................................................................................................3 Micropylar plate placed on a ± prominent longitudinal dorsal ridge or keel; internally surrounded by a broad brown area (Figs. 64–65)...............................................................................................Acanthoxylini Internal micropylar plate open......................................................................................................................4 Internal micropylar plate closed (Fig. 66b) .....................................................Gigantophasmatini trib. nov. Operculum with a distinctly stalked capitulum; micropylar plate covering > 1/3 of capsule length (Figs. 48–51) .................................................................................Phasmatini, Acanthomimini & Tropidoderini** Operculum with central region conically raised, no stalk present; micropylar plate covering < 1/3 of capsule length (Figs. 52–53) .................................................................................................... Monandropterini

*

Eggs of Xeroderus Gray, 1835 the type genus of Xeroderinae are not known. Eggs of Pachymorphinae: Pachymorphini are excluded from the key above since they include various distinct shapes and the tribe is believed to be polyphyletic (→4.6.1) ** Based on the insufficient material at hand the eggs of these three tribes appear impossible to distinguish in a key

4.3 Phasmatinae Gray, 1835 (Fig. 1) Type genus: Phasma Lichtenstein, 1796: 77. Phasmidae Gray, 1835: 12. Phasminae Karny, 1923: 240. Günther, 1953: 554. Phasmatinae Bradley & Galil, 1977: 192 (in part). [Corrected spelling of Phasminae Karny, 1923] Otte & Brock, 2005: 13. Acrophyllae Brunner v. Wattenwyl, 1893: 97. Acrophyllinae Kirby, 1904a: 379.

Description: Medium sized to very large (body length 45.0–300.0 mm), mostly stick-like, moderately robust to extremely slender and elongate Phasmatidae (Figs. 72–73, 75). Body cylindrical to subcylindrical in crosssection and mostly of more or less uniform width, abdomen of && rarely widened sub-basally or medially. Ocelli frequently present in most alate taxa, lacking in apterous taxa. Gula present. Antennae at best 3/5 the length of body, often conspicuously shortened, with less than 45 segments. Mesothorax at least as long as head and pronotum combined and much longer in apterous taxa; often granulose or spinose. Both sexes winged, having shortened tegmina and fully developed alae, brachypterous or apterous; tegmina less than half as long as alae. Wings often unequal between the sexes and better developed in %%. Abdomen at least as long as but usually distinctly longer than thorax. Median segment distinctly shorter, equal in length, or longer than


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the metanotum. Tergites V–VII often with lateral lobes or expansions (&& in particular). Sternum VII of && lacking a praeopercular organ. Gonapophyses of && elongated in certain apterous taxa, but never projecting considerably over apex of abdomen. Subgenital plate of && boat-like, distinctly and acutely keeled in basal portion. Length and shape variable, ranging from small, scoop-shaped and not reaching the apex of anal segment, to considerably elongated and lanceolate. Anal segment of %% laterally compressed, tectiform and consisting of two semi-tergites, which either have their interior surface with a longitudinal spinose bulge, or with the ventro-apical angles ± expanded, in-curving and armed with terminal teeth (Figs. 2–4, 32). External vomer of %% lacking (Fig. 38). Supraanal plate small and considerably shorter than anal segment. Cerci ± prominently enlarged, laterally compressed and ± foliaceous. Profemora distinctly triangular in cross-section with the two dorsal carinae ± distinctly approaching each other; compressed and curved basally. Posterodorsal carina reduced and anterodorsal carina ± strongly raised and serrate. Medioventral carina present, moderately distinct and about midways on ventral surface of profemur (Fig. 44). Mid- and hind legs often with prominent armature. Meso- and metafemora trapezoidal in cross-section with dorsal carinae nearing, medioventral carina present. Tibiae trapezoidal to almost triangular in cross-section, the dorsal carinae strongly nearing each another. No area apicalis on tibiae. Basitarsus distinctly longer than following tarsomere.

FIGURE 1. Geographical distribution of the subfamilies Phasmatinae, Tropidoderinae and Extatosomatinae (Phasmatidae s. str.) and Clitumninae (Phasmatidae s. l.).

Comments: The subfamily Phasmatinae was previously either attributed to Brunner v. Wattenwyl (1893), Karny (1923) or Bradley & Galil (1977). However it was Gray (1835: 12) who first introduced “Phasmidae” as a family-group name for what now is the order Phasmatodea. According to ICZN article 36.1 on the principle of coordination applied to family-group names, Gray (1835) must also be regarded the author of current REVISION OF PHARNACIINI


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Phasmatinae and the tribe Phasmatini simultaneously. Brunner v. Wattenwyl (1893: 99) used Phasmata as a term for Neotropical taxa. Doubtful genera: The Australian Echetlus Stål, 1875 (Type species: Bacillus peristhenes Westwood, 1859: 13, pl. 7: 1) was included in Platycraninae by Günther (1953: 557) and has since been retained in the subfamily by all subsequent authors. Only Brock & Hasenpusch (2007: 74) doubted Echetlus to be a member of “Platycraninae” and stated it was likely to belong in Phasmatinae. Indeed, Echetlus is obviously not at all closely related to Platycrana Gray, 1835 or any of the genera of Platycraninae. Features such as the triangular cross-section of the profemora, cylindrical body, shortened antennae and long, lanceolate cerci indicate relation to the subfamily Phasmatinae, but more material is needed for any confirmed decision on its systematic position. Two Brazilian species, E. evoneobertii Zompro & Adis 2001 and E. fulgens Zompro 2004, have recently been described in Echetlus Stål and E. evoneobertii was erroneously suggested to have been introduced from Australia (Zompro & Adis, 2001: 291). Brock & Hasenpusch (2007: 7) removed E. evoneobertii from Echetlus and transferred it to the Australian Candovia Stål, 1875 (Type species: Phasma (Bacteria) coenosa Gray, 1833: 17, 28, pl. 2: 2), but were in doubt about its Australian origin (P.D. Brock pers. comm.). More detailed examination of these two Brazilian taxa clearly proves them to be definitely not members of Echetlus or the Australasian subfamily Platycraninae but to belong in the Neotropical Diapheromeridae: Diapheromerinae: Diapheromerini. This is confirmed by the proportionally small head, very long and filiform antennae, being about 2/3 the length of the body and consisting of far more than 30 segments (conspicuously shortened and robust in Echetlus), cylindrical cerci, unarmed medioventral carina of the femora, triangular cross-section of the profemora and egg structures, such as the ovoid capsule, open internal micropylar plate and irregularly raised, rim-like structure of the operculum. Within the Diapheromerini evoneobertii and fulgens strongly resemble the Central American genus Paracalynda Zompro, 2001. Further research by the authors is in progress. Distribution (Fig. 1): Australia, Tasmania, Papuan Subregion (except Micronesia), Wallacea, Borneo, Philippines and New Zealand. Tribes included: 1. Phasmatini Gray, 1835: 12. Type genus: Phasma Lichtenstein, 1796: 77. 2. Acanthomimini Günther, 1953: 555. Type genus: Acanthomima Kirby, 1904a: 438. 3. Acanthoxylini Bradley & Galil, 1977: 192. Type genus: Acanthoxyla Uvarov, 1944: 95

4.3.1 Phasmatini Gray, 1835 (Figs. 2–4, 32, 38, 44, 48–49, 72–73, 75, 431–432, 434–435) Type genus: Phasma Lichtenstein, 1796: 77. Phasmidae Gray, 1835: 12. Phasmata Brunner v. Wattenwyl, 1893: 99. Phasmini Günther, 1953: 554. Phasmatini Bradley & Galil, 1977: 192 & 194. [Corrected spelling of Phasmini Günther, 1953] Otte & Brock, 2005: 13. Acrophyllini Redtenbacher, 1908: 436 (in part). Giglio-Tos, 1910: 43.

Description: This tribe predominantly includes very striking insects of remarkable size (body length of && 70–300 mm), most of which have well developed tegmina and alae, produced ocelli (at least %%), flattened and considerably dilated often foliaceous or lanceolate cerci, a more or less distinctly spinose or granulose


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mesothorax and serrate or otherwise armed extremities. The sexual dimorphism is distinct with %% being considerably smaller and more slender than corresponding &&. Most representatives are of a brown general colouration, only two genera being frequently green (Eurycnema and Paracyphocrania). Some taxa have the ocelli strongly reduced or lacking (e.g. Onchestus) or increasingly reduced in && (e.g. Anchiale and Paracyphocrania). In a great majority of genera the head is ovoid to almost spherical and unarmed, but may have the vertex conically raised (Peloriana) or armed with one or two pairs of spines (Onchestus and Paronchestus). The antennae of %% are longer than those of &&, but at best slightly longer than the head and complete thorax combined in %%, and usually not longer than the mesothorax in &&. The scapus is simple and rather slender. The antennomeres may either be elongate and much longer than wide or distinctly shortened towards the base of the antennae (e.g. Eurycnema, Paracyphocrania and Paractenomorpha). The mesothorax may be elongate and at laest 1.3x longer than the head and pronotum combined, often distinctly longer (e.g. Ctenomorpha and Paractenomorpha). Often the mesothorax, metapleurae and metasternum are to a variable degree set with granules, tubercles or blunt spines, those of the pleurae usually positioned in a longitudinal row. The median segment is considerably longer than the metanotum. The tegmina are usually considerably shorter than the alae and at best cover the first two abdominal segments. The alae may be as long as reaching almost to the tip of the abdomen (e.g. Phasma), or are stepwise reduced up to being just indistinctly longer than the tegmina (e.g. Ctenomorpha and Paractenomorpha). The anal region of the alae is to a various degree furnished with brown, grey or black markings, tesselated or reticulate (e.g. Acrophylla, Ctenomorphodes or Phasma) to almost entirely black, and rarely transparent (e.g. Eurycnema). There is no praeopercular organ on sternum VII of &&. The semi-tergites of the anal segment of %% are short and have the posteroventral angles swollen and the interior surfaces bearing a distinct, longitudinal keel or bulge which is armed with a row of 3–10 prominent teeth (“Dornenfeld”, Figs. 32, 38). An external vomer is lacking. The cerci are distinctly laterally flattened and carinated, more or less prominently enlarged and foliaceous or lanceolate, in some genera being as long as the combined length of abdominal tergites VIII–X. Only in a few cases the cerci are shorter than the anal segment (e.g. Cigarrophasma or && of Paracyphocrania and Onchestus). They are usually larger in %% than in corresponding &&. They are either simple, laterally flattened and ovoid in lateral aspect (e.g. Anchiale), or distinctly tri-carinate and truncate apically (e.g. Eurycnema and certain species of Phasma and Acrophylla). The subgenital plate of && is strongly keeled and boat-shaped, which in particular concerns to the basal portion, and often exceeds the apex of the abdomen. The poculum of %% is rather bulgy, strongly-convex and cup-like. All legs are more or less prominently armed with spines, teeth or lobes. The profemora are clearly triangular in cross-section with the two dorsal carinae strongly approaching each other, the anterodorsal carina considerably raised and ± distinctly serrate and the posterodorsal carina reduced; the latter carina may sometimes be slightly displaced towards the lateral surface of the femur. The medioventral carina of the profemora is very indistinct and set with a few minute spines (e.g. Onchestus) or is lacking. If present it is roughly placed midways on the ventral surface of the femur (Fig. 44). Eggs (Figs. 48–49): Eggs of Phasmatini are mostly of a very simple ovoid or spherical general shape (Figs. 48a & 49a) but some genera have the capsule covered with prominent ridges and bulges (e.g. Onchestus and Ctenomorpha), or hairy structures (e.g. Eurycnema goliath Gray, 1834). The operclum bears a distinct hat or knob-like capitulum on a short, ± conspicuous stalk. The micropylar plate is elongate, suboval or parallelsided and covers more than half of the capsule length (exception Eurycnema). Internally the plate exhibits a distinct but in most cases narrow posteromedian gap and lacks a median line (Figs. 48b, 49b). Comments: Günther (1953: 554) established Phasmini amongst six other tribes in his subfamily Phasminae. Bradley & Galil (1977: 194) corrected the spelling of Günther’s taxa to Phasmatinae and Phasmatini respectively. Both authors characterized it amongst the tribes of Phasmatinae by the following combination of characters: both sexes alate or with rudimentary wings; cerci of && strongly flattened and dilated or lanceolate and ocelli of %% distinct. Certain genera strikingly resemble representatives of the exceptionally Australian Tropidoderinae: Tropi



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FIGURES 2–19. Terminal abdominal segments of %% of Phasmatidae s. str. [scale = 5 mm] 2–4. Tribe Phasmatini: Phasma reinwardti (de Haan, 1842), 2. lateral view, 3. dorsal view, 4. ventral view 5–7. Tribe Tropidoderini: Tropidoderus childrenii Gray, 1833, 5. lateral view, 6. dorsal view, 7. ventral view 8–10. Tribe Tropidoderini: Parapodacanthus hasenpuschorum Brock, 2003, 8. lateral view, 9. dorsal view, 10. ventral view 11–13. Tribe Gigantophasmatini trib nov.: Gigantophasma bicolor Sharp, 1898, 11. lateral view, 12. dorsal view, 13. ventral view 14–16. Tribe Monandropterini: Monandroptera acanthomera (Burmeister, 1838), 14. lateral view, 15. dorsal view, 16. ventral view 17–19. Extatosomatinae: Extatosoma tiaratum tiaratum (Mac Leay, 1826), 17. lateral view, 18. dorsal view, 19. ventral view.


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doderini and indicate relation between Phasmatinae and Tropidoderinae, a fact already suggested by Stål (1877b: 65). For instance, the bright green general colouration, rather short and robust antennae with considerably shortened antennomeres, long alae with a plain anal region, medially swollen abdomen of && and rather short micropylar plate of the eggs of Eurycnema, resemble Tropidoderus Gray, 1835 (tribe Tropidoderini). In particular, %% of Paratropidoderus Brock & Hasenpusch, 2007 (Tropidoderinae: Tropidoderini) remarkably resemble those of Eurycnema by having a rather long and slender mesothorax, which dorsally bears a distinct longitudinal median carina and two rows of distinct spines, and comparatively short, ovate cerci. Another example is represented by && of Paracyphocrania which show remarkable overall similarity to Malandania Sjöstedt, 1918 (Tropidoderinae: Tropidoderini) and only differ by the key features such as the serrate profemora and convex meso- and metasternum. On the other hand, some more slender representatives resemble certain genera of the Oriental Clitumninae: Pharnaciini, at first glance appearing like winged forms of the latter. Examples include the following the generic pairs Peloriana and Tirachoidea Brunner v. Wattenwyl, 1893, and Ctenomorpha and Phobaeticus Brunner v. Wattenwyl, 1907 (→ 5.2). It both cases only key features of Tropidoderinae: Tropidoderini, or Phasmatidae respectively, distinguish these taxa. A complete revision of Phasmatini at the species level is badly needed. The taxonomic status of the Australian taxa has been clarified by Brock (1998b), Brock & Hasenpusch (2005 & 2007) and Hasenpusch & Brock (2006) and illustrated keys to the Australian genera and species will be published by Brock & Hasenpusch (in press—personal communication with P. D. Brock). Distribution: Members of Phasmatini are distributed over almost the complete Australian Region as far east as the Solomon Islands. Single species are represented as far west as the Large Sunda Islands (Java & Sumatra) and as far north as the Philippines. Genera included: 1. Acrophylla Gray, 1835: 38. Type species: Phasma titan MacLeay, 1826: 454, by subsequent designation of Kirby, 1904a: 388. = Vetilia Stål, 1875: 36. Type species: Acrophylla enceladus Gray, 1835: 39, by monotypy. [Synonymised by Brock & Hasenpusch, 2007: 72] 2. Anchiale Stål, 1875: 36 & 84. Type species: Mantis maculata Olivier, 1792: 626, by monotypy. = Ctenomorphodes Karny, 1923: 240. Type species Phasma (Diura) briareus Gray, 1834: 45, by original designation. [Synonymised by Brock & Hasenpusch, 2007: 72] 3. Cigarrophasma Brock & Hasenpusch, 2001: 4. Type species: Cigarrophasma tesselatum Brock & Hasenpusch, 2001: 6, figs. 1–7, by original designation. 4. Ctenomorpha Gray, 1833: 16 & 27. Type species Ctenomorpha marginipennis Gray, 1835: 16, by subsequent designation of Kirby, 1904a: 388. 5. Eurycnema Audinet-Serville, 1838: 235. Type species: Cyphocrana (Eurycnema) versirubra Audinet-Serville, 1838: 235, by subsequent designation of Kirby, 1904a: 391. = Clemacantha Rainbow, 1897: 34. Type species: Clemacantha regale Rainbow, 1897: 34, by monotypy. [Synonymised by Günther, 1934b: 527] 6. Onchestus Stål, 1877b: 63. Type species: Lopaphus gorgus Westwood, 1859: 102, pl. 11: 4, by subsequent designation of Kirby, 1904a: 394. 7. Paractenomorpha Hennemann & Conle, 2004: 15. Type species: Paractenomorpha baehri Hennemann & Conle, 2004: 17, figs. 1–4, by original designation. 8. Paracyphocrania Redtenbacher, 1908: 466. Type species: Paracyphocrania lativentris Redtenbacher, 1908: 466, by monotypy. 9. Paronchestus Redtenbacher, 1908: 463. Type species: Paronchestus charon Redtenbacher, 1908: 463, pl. 27: 8a & 8b, by monotypy. 10. Peloriana Uvarov, 1940: 115. Type species: Cyphocrania lobiceps MacLeay, 1884: 711, by indication.



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[Replacement name for Peloria Redtenbacher, 1908: 462] = Peloria Redtenbacher, 1908: 462, pl. 22: 7a & b. Type species: Cyphocrania lobiceps MacLeay, 1884: 711, by monotypy. [Preoccupied by Peloria Cocco, 1884 (Pisces)] 11. Phasma Lichtenstein, 1796: 77. Type species: Phasma empusa Lichtenstein, 1796: 77, by subsequent designation of Kirby, 1904a: 390 (= Phasma gigas (Linné, 1758)). = Cyphocrana St. Fargeau & Audinet-Serville, 1825: 445. Type species: Gryllus (Mantis) gigas Linné, 1758: 425, by subsequent designation of Kirby, 1904a: 390. [Synonymised by Kirby, 1904a: 390]. = Papuanoidea Werner, 1930: 181. Type species: Papuanoidea straleni Werner, 1930: 181, by monotypy [Synonymised by Günther, 1936: 339].

4.3.2 Acanthomimini Günther, 1953 (Fig. 79) Type genus: Acanthomima Kirby, 1904a: 438. Acanthomimini Günther, 1953: 555. Bradley & Galil, 1977: 193. Cliquennois & Brock, 2004b: 2. Otte & Brock, 2005: 32

Diagnosis: The following brief diagnosis is solely based on the two genera Acanthomima Kirby, 1904b and Anophelepis Westwood, 1859. Representatives of this tribe are rather small (body length of && < 12 cm), slender and stick-like, brachypterous Phasmatinae. %% in particular are very slender insects with long and slender legs. Colouration green or brown. && have great parts of their dorsal body surface granulose, but the mesothorax in particular. The body of %% is smooth. The head is > 1.3x longer than wide, broadest at the eyes and distinctly narrowing towards the posterior; often with a pair of small tubercles or humps between the eyes. Ocelli lacking. The antennae are short and consist of 25–30 segments. They are slender and roughly reach the posterior margin in %%, but have the antennomeres broad and strongly shortened with the antennae merely reaching half way along the mesonotum (&&). The scapus is dorsoventrally flattened and moderately dilated laterally. The pronotum is rectangular and of dimension similar to that of the head. The mesothorax is elongate and at least 1.7x longer than the head and pronotum combined, the mesonotum having a fine longitudinal median carina. The oval tegmina are roughly equal in length to the metanotum and just cover the bases of the alae. The alae are shorter or equal in length to the tegmina and at best reach about half way along the median segment. The anal region is steel blue to black. The median segment is considerably longer than the metanotum and the abdomen, excluding the median segment, longer than the head and thorax combined. All abdominal segments are longer than wide, the terminal three ones shortened. Tergites VIII and IX of %% are thickened and considerably broader than the previous. The anal segment of %% is typical for members of Phasmatinae, being strongly tectiform and consisting of two angulate interiorly dentate semi-tergites. An external vomer is lacking. The subgenital plate of && is very small, boat shaped and at best reaches half way along the anal segment. The cerci are laterally flattened, being small, oval and shorter than the anal segment in &&, but rather lanceolate and about as long as the anal segment in %%. All legs are long and slender (%% in particular), the two posterior pairs having the ventral carinae sparsely spinulose, or at least with 2–3 minute sub-basal teeth on the two outer carinae. A few sub-basal teeth may be present on the dorsal carinae of these femora. The profemora are distinctly triangular in cross-section, with the anterodorsal carina strongly raised and serrate. Sometimes a few small teeth are present on the posteroventral carina in &&. The medioventral carina of the profemora is acute and slightly displaced towards the anterior carina. Meso- and metafemora trapezoidal in cross-section.


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The tarsi are slender with the basitarsi longer than the following three tarsomeres combined, the two outer ventral carinae sometimes denticulate. Eggs: Rather small and ovoid with a small knob-like capitulum. The capsule surface is slightly wrinkled and the micropylar plate elongate, slender and covering about ¾ of the capsule length, with the micropylar cup placed close to the posterior end. From a superficial external examination, the micropylar plate appears to be open internally (a photo of the eggs of Anophelepis telesphorus Westwood, 1859 was kindly provided by P.D. Brock). Comments: Günther (1953: 555) established Acanthomimini for the two Australian genera Acanthomima Kirby, 1904b: 438 (= Ectus Redtenbacher, 1908: 437) and Carnacia Sjöstedt, 1918: 50 and placed it in close relation to Stephanacridini Günther, 1953. The latter genus was synonymised with Anophelepis Westwood, 1859, a member of Phasmatinae: Phasmatini, by Vickery (1983: 4). Günther characterized Acanthomimini amongst Phasminae (= Phasmatinae) by the following combination of features: cerci of && short and slender, %% without ocelli; tegmina and alae of && strongly reduced; head flat; middle and hind femora at most with a few small spines at the apex. Cliquennois & Brock (2004: 2) transferred Anophelepis from Platycraninae to Acanthomimini and Brock & Hasenpusch (2007: 74) again supposed it was misplaced in Platycraninae and likely to belong in Phasmatinae. Indeed, Anophelepis Westwood is not a member of Platycraninae which is obviously seen in a number of features such as the triangular cross-section of the profemora, slender cheeks, laterally positioned eyes, or lack of functional prothoracic defensive glands. The type genus of Acanthomimini Acanthomima is unfortunately as yet only known from the &&. Features such as the distinctly triangular cross-section of the profemora, which have the anterodorsal carina raised decidedly serrate, cylindrical body, short antennae, laterally compressed cerci and keeled subgenital plate obviously place Acanthomima in the subfamily Phasmatinae and support relationship between Acanthomimini and Phasmatini. Amongst this tribe Acanthomima and Anophelepis show affinity to Paractenomorpha Hennemann & Conle, 2003 with which they share the short subgenital plate, shortened alae and short, robust antennae of &&. This relationship is furthermore indicated by structures of the eggs, such as the ovoid and rather plain capsule, knob-like capitulum and elongate micropylar plate which is supposedly open internally. In general they resemble those of certain Phasmatini, e.g. some species of Anchiale Stål, 1875 or Phasma Lichtenstein, 1796. New diagnoses and illustrations of the species of these two genera will shortly be available in a Field Guide of the Australian Phasmatodea by Brock & Hasenpusch (in press—pers. comm. with P. D. Brock). Doubtful genera: Cliquennois & Brock (2004b) described Mauritiophasma, a new genus from Mauritius, which they suggested to belong in Acanthomimini. The egg of Mauritiophasma is remarkable for being glued to a support by the operculum, a feature so far only known from another not closely related Mauritian genus, Monoiognosis Cliquennois & Brock, 2004b. Unlike Acanthomima, it has a closed internal micropylar plate (personal communication with N. Cliquennois). %% of Mauritiophasma have the vomer rudimentary and functionally replaced by spiny forceps on the interior surfaces of the semi-tergites of the tectiform anal segment. However, in contrast to Acanthomimini the spines of the semi-tergites are positioned at their posterodorsal angles. Whether Mauritiophasma is correctly placed in Acanthomimini needs further research and requires knowledge of the % and egg of Acanthomima, the type genus of Acanthomimini. However, since several features, such as the anal segment of %% and remarkable eggs with a closed internal micropylar plate, violate the diagnostic features of Acanthomimini, Mauritiophasma is rather unlikely to belong here. The presence of a possible representative of Acanthomimini on the volcanic island Mauritius could however be explained by an east-west directed stream in the Indian Ocean, which passes near coastal Western Australia and arrives close to Mauritius. As the eggs are glued to branches, eggs of the Australian ancestor might have reached Mauritius thanks to a floating raft (Cliquennois & Brock, 2004b: 2). The following two Australian genera are apparently closely related to each other. Within the Phasmatinae Arphax Stål, 1875 keys close to Acanthoxylini due to having a median segment that is shorter than the metan-



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otum. Vasilissa Kirby, 1896 on the other hand has a median segment which is approximately as long as in Acanthomimini but %% are alate and rather resemble certain taxa of Phasmatini. They are here provisionally attributed to Acanthomimini but with doubt. Certainly, more material is necessary to decide about the systematic position of these two genera, but research is believed to be on-going. Arphax was previously included in Acanthoxylini Bradley & Galil, 1977, a tribe endemic to New Zealand (→ 5.3.3). However, the distribution in Australia and features of the insects and eggs violate the diagnostic features of Acanthoxylini, which is believed to be endemic in New Zealand. %% possess the typical tectiform anal segment found throughout Phasmatinae or Tropidoderinae, consisting of two separate semi-tergites which have the posteroventral angle swollen and interiorly armed with a row of conspicuous spines (= “Dornenfeld”). Both sexes are slender, cylindrical and apterous, stick-like insects with short and rather robust antennae. The median segment is shorter than the metanotum. The head is cylindrical and distinctly longer than wide. Ocelli are lacking. && of certain species have the subgenital plate very long and lanceolate, others short and barely reaching the apex of the abdomen. Both sexes have rather lanceolate, laterally compressed cerci. The eggs are rather typical for Phasmatinae, being ovoid to elliptical and bearing a conspicuous capitulum. The micropylar plate is elongate, slender and covers almost the entire dorsal surface of the capsule. Internally the plate appears to be open (based on illustration provided by Brock & Lowe, 1998: 76, fig. 6c). Vasilissa was previously placed in Tropidoderinae: Tropidoderini, but the type species V. walkeri Kirby, 1896 from Queen’s Islet (NW-Australia) is obviously closely related to Arphax. It was therefore transferred to Phasmatinae by Brock & Hasenpusch (2007: 74) who commented it may only warrant subgenus status. It is merely distinguished from representatives of Arphax by having the median segment slightly longer than the metanotum, && possessing scale-like tegmina, and %% having rather well developed alae, which reach about half way along abdominal segment V. The anal region of the alae of %% is transparent and tesselated with dark brown, resembling e. g. Anchiale Stål, 1875 (tribe Phasmatini). The Philippine Vasilissa tecticollis Redtenbacher, 1908 was shown to belong in the genus Paracyphocrania Redtenbacher (Hennemann & Conle, 2006a: 97). Distribution: Australia, believed to be endemic. Genera included: 1. Acanthomima Kirby, 1904b: 438. Type species: Anophelepis ripheus Westwood, 1859: 70, pl. 8: 10, 10a, 10b, by original designation. = Ectus Redtenbacher, 1908: 437. Type-secies: Ectus solitarius Redtenbacher, 1908: 437, pl. 21: 1, by monotypy. [Synonymised by Vickery, 1983: 3] 2. Anophelepis Westwood, 1859: 68. Type species: Anophelepis telesphorus Westwood, 1859: 69, pl. 8: 3, 7, 7a, , by subsequent designation of Kirby, 1904a: 382. [Here removed from Platycraninae] = Carnacia Sjöstedt, 1918: 50. Type species: Carnacia obscura Sjöstedt, 1918: 51, pl. 6: 4a–c, by monotypy. [Synonymised by Vickery, 1983: 4] ? Arphax Stål, 1875: 37. Type species: Bacillus australis Charpentier, 1845: plate 57, by monotypy. [Here removed from Acanthoxylini] = Austroclonistria Redtenbacher, 1908: 438. Type species: Austroclonistria serrulata Redtenbacher, 1908: 438, by monotypy. [Synonymised by Brock & Hasenpusch, 2007: 72] ? Vasilissa Kirby, 1896: 468. Type species: Vasilissa walkeri Kirby, 1896: 469, by monotypy. [Here removed from Phasmatinae: Acanthoxylini] ? Mauritiophasma Cliquennois & Brock, 2004b: 2. Type species: Mauritiophasma motalai Cliquennois & Brock, 2004: 3, figs. 1–8, 33, by original designation.


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4.3.3 Acanthoxylini Bradley & Galil, 1977 (Figs. 64–65) Type genus: Acanthoxyla Uvarov, 1944: 95. Macracanthini Günther, 1953: 555. Acanthoxylini Bradley & Galil, 1977: 192. Jewell & Brock, 2003: 192. Otte & Brock, 2005: 32.

Description: This tribe containes exceptionally apterous, medium-sized (body length 50–150 mm), slender to moderately broad insects. The dorsal surface of the head and body of both sexes is to a various degree covered with spines, rarely unarmed. The head is longer than wide, broadest at the eyes and ± distinctly narrowing towards the posterior; vertex flat or very gently rounded. Ocelli are lacking. The antennae are ± as long as the head, pro and mesonotum combined and consist of less than 30 segments. The scapus is strongly dorsoventrally flattened and roundly elevated laterally. A small gula is present. The mesothorax is of moderate length and 1.3–2.8x longer than the head and pronotum combined. The median segment is very short, being less than 1/3 the length of the metanotum. Abdominal tergite VII of && usually bears a ± distinct, rounded posterolateral lobe; sometimes similar but smaller lobes are also present on V and VI. && lack a praeopercular organ on sternum VII and have the subgenital plate is strongly keeled, boat shaped but never extending considerably over the apex of the abdomen. Certain && are typical for having the subgenital plate with a prominent, backcurving spine (Acanthoxyla) or tubercle (Pseudoclitarchus) near the base. %% have a tectiform and split anal segment, which consists of two separate semi-tergites and forms a conspicuous clasping apparatus. The semitergites are however of a rather typical shape in each of the three genera contained (%% of Acanthoxyla are not known). In Argosarchus they have the posteroventral angles elevated into a finger-like, distinctly in-curving process which bears 4–5 small, terminal teeth. In Clitarchus they are rounded and moderately elongated posteroventrally and have 4–5 distinct interior teeth along the posterior margin. %% of Pseudoclitarchus have the semi-tergites rather broad and rounded with a single, prominent interior tooth. An external vomer is missing. Both sexes possess conspicuously enlarged, laterally compressed cerci, which are ± oval in lateral aspect. The profemora are distinctly triangular in cross-section with the posterodorsal carina reduced and the anterodorsal carina ± distinctly raised. The medioventral carina is prominent, ledge-like and slightly displaced towards the anteroventral carina. The remainder femora are trapezoidal in cross-section, the tibiae triangular with the dorsal carinae strongly approaching each other. All carinae of legs to a various degree armed with teeth, serrations or lobules, single carinae may be unarmed. Basitarsi at least as long as following two tarsomeres combined and slender (exception && of Argosarchus). Eggs (Figs. 64–65): Eggs are of various shapes, ranging from roughly ovoid (Clitarchus, Fig. 64a) to laterally compressed and alveolar with a ± prominent dorsomedian bulge (Acanthoxyla, Fig. 65a). The capsule is distinctly longer than high and the surface ± strongly sculptured, being to a variable degree punctured or irregularly set with acute, tooth-like ridges. Opercular collar roughly tuberculose. Polar-area flattened or impressed. The operculum is circular and ± strongly conically raised. In Clitarchus it is hollow conical operculum, whilst in Acanthoxyla and Argosarchus it is a spongy cone (Clark-Sellick, 1988: 277). The micropylar plate is rather small, rhomboidal or lozenge-shaped and ± distinctly displaced towards the polar end of the capsule. The micropylar cup is positioned in the posterior half of the plate. Internally, the plate is distinctly pointed at both ends and appears to be closed (Clark-Sellick, 1998: 225). It is however remarkable for having the inner portion white, as usual in Phasmatodea, but surrounded by first a cream-coloured and then a dark brown outer area. The micropylar stalk is placed well within the borders of the white inner portion of the plate (Argosarchus), but may either have a long and slender extension which almost reaches the polar end of the white portion (Acanthoxyla, Fig. 65b), or is connected with the outer brown area by a raised, longitudinal



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ridge, which resembles a median line (Clitarchus, Fig. 64b). If the white inner portion alone is interpreted as the micropylar plate in the latter genus, this would give rise to the impression of an open plate with a median line. Similar internal micropylar plates with a dark outer area are otherwise only known from the Neotropical genus Agathemera Stål. Comments: Günther (1953: 55) originally placed Macracanthini (= Acanthoxylini Bradley & Galil, 1977) in Phasminae (= Phasmatinae Bradley & Galil) and included the following four genera in the tribe: Acanthoxyla Uvarov, 1944 (= Macracantha Kirby, 1904), Argosarchus Hutton, 1898 (= Mimarchus Carl, 1913), Clitarchus Stål, 1875 and Arphax Stål, 1875. The Australian Arphax Stål is obviously not a member of Acanthoxylini and here provisionally attributed to Acanthomimini which however requires further evaluation (→ 4.3.2). Already Jewell & Brock (2003: 189) have pointed out that the New Zealand fauna are not closely related to Australian Phasmatodea. Acanthoxylini is endemic to New Zealand, which was separated from Australia by the Tasman Sea for about 80 million years (Austin et al., 2004: 227), and moreover features of the eggs, such as the conical operculum, dorsomedian bulge and small lozenge-shaped micropylar plate, as well the genitalia of the insects, which include laterally compressed cerci, lack of a praeopercular organ in && and specialised clasping apparatus of the %% anal segment, indicate close relationship to the other New Zealandean genera currently placed in the subfamily Pachymorphinae, namely Asteliaphasma Jewell & brock, 2003, Tectarchus Salmon, 1954, Spinotectarchus Salmon, 1991 (tribe Hemipachymorphini) and Micrarchus Carl, 1913 and Niveaphasma Jewell & Brock, 2003 (tribe Pachymorphini). With the exception of Micrarchus, which may despite genital differences be close to the Australian Pachymorpha Gray, 1835 (Jewell & Brock, 2003: 189), all of these genera are apparently difficult to distinguish from Acanthoxylini on a tribal basis and likely to belong in this tribe. This hypothesis however requires further evaluation and availability of sufficient material for detailed investigation. Studies discussing the relationships of these genera and their position within the family Phasmatidae are believed to be currently on-going. Due to the laterally compressed and ± foliaceous cerci and anal segment of %% forming a clasping apparatus Bradler (2001: 183) included Acanthoxylini in his Lanceocercata (= Phasmatidae). Acanthoxylini exhibits the typical feature, which already Günther (1953) and Bradley & Galil (1977) used to characterize Phasmatinae, the triangular cross-section of the profemora, and it is mainly due to this feature that the tribe is here retained in the Phasmatinae. There are several characters which readily distinguish members of Acanthoxylini from other Phasmatinae, e.g.: the spiny body; distinct and acute medioventral carina of the profemora; strongly sculptured, ± alveolar egg-capsule; lack of a capitulum but conically raised operculum instead, closed internal micropylar plate (→ above), and geographical distribution. Other features such as the basal spine or tubercle of the subgenital plate of && of Acanthoxyla and Pseudoclitarchus, shape and structure of the semi-tergites of the anal segment of %% as well as the egg-morphology show affinity to certain members of the subfamily “Platycraninae”, e.g. Megacrania Kaup, 1871. Eggs of Acanthoxylini (Acanthoxyla & Argosarchus in particular) strongly resemble those of certain “Platycraninae” (e.g. Megacrania Kaup, 1871, Ophicrania Stål, 1875 or Graeffea Brunner v. Wattenwyl, 1868), the capsule being alveolar, laterally compressed and with a more or less distinct longitudinal dorsal bulge, the capsule surface strongly sculptured, the micropylar plate small, rhomboidal or lozenge-shaped and displaced towards the posterior pole of the egg, and the operculum ± strongly conically raised. A hyopothetical relationship to “Platycraninae” (not type genus Platycrana Gray, 1835) however needs further evaluation. Salmon (1991: 36) stated Clitarchus, Pseudoclitarchus and Acanthoxyla to be closely related to one another and that Pseudoclitarchus formed an evolutionary link between Clitarchus and Acanthoxyla. ClarkSellick (1998: 225) studied the internal micropylar plates of the four genera currently included and suggested the tribe to be polyphyletic, which is however not supported by the exosceleton of the insects. Certainly, further consideration of the position of Acanthoxylini in the classification of Phasmatodea requires detailed examination and extensive comparison of all taxa concerned and careful comparison with supposedly closely related groups.


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Distribution: New Zealand, endemic. Genera included: 1. Acanthoxyla Uvarov, 1944: 95. Type species: Acanthoderus prasinus Westwood, 1859: 49, pl. 3: 2, by indication. [Replacement name for preoccupied Macracantha Kirby, 1904] = Macracantha Kirby, 1904a: 340. Type species: Acanthoderus prasinus Westwood, 1859: 49, pl. 3: 2, by original designation. [New name for Acanthoderus Hutton, 1899 (not Gray, 1835). Preoccupied by Macracantha Simon, 1864 (Araneae)] 2. Argosarchus Hutton, 1898: 165. Type species: Phasma (Acanthoderus) horridus White, 1846: 24, pl. 5: 3, by subsequent designation of Kirby, 1904a: 340. = Mimarchus Carl, 1913: 22. Type species: Mimarchus tarsatus Carl, 1913. 23, by monotypy. [Synonymised by Jewell & Brock, 2003: 194] 3. Clitarchus Stål, 1875: 34, 82. Type species: Clitarchus laeviusculus Stål, 1875: 82, by subsequent designation of Kirby, 1904a: 339. 4. Pseudoclitarchus Salmon, 1991: 35, 77. Type species: Acanthoxyla senta Salmon, 1948: 301, pl. 56: 1–4, pl. 57: 1–3, by original designation.

4.4 Tropidoderinae Brunner v. Wattenwyl, 1893 (Fig. 1) Type genus: Tropidoderus Gray, 1835: 31. Tropidoderi Brunner v. Wattenwyl, 1893: 97. Tropidoderinae Bradley & Galil, 1977: 194. Otte & Brock, 2005: 13. Podacanthinae Günther, 1953: 553.

Description: Medium sized to large (body length 43.0–187.0 mm), moderately broad and robust robust Phasmatidae, && in particular. Body slightly compressed dorsoventrally and ± oval in cross-section (&& in particular). Abdomen ± widened sub-basally or medially in &&, parallel-sided in %%. Head orthognathous, vertex either flattened or gently convex nad unarmed. Ocelli present in alate and lacking in apterous taxa. Gula present. Antennae at best 2/3 the length of body and with less than 45 segments, often conspicuously shortened in &&. Mesothorax short, often hardly longer than pronotum in alate taxa, but increasingly elongated and at best 1.8x longer than head and pronotum combined in certain apterous taxa; ± distinctly widened towards the posterior. Mesonotum, meso- and metapleurae often granulose or spinose. Meso- and metasternum ± flattened. Either both sexes winged, having shortened tegmina and long, fully developed alae which in most cases cover the complete abdomen, both sexes or only the && apterous. Tegmina less than half as long as alae. Abdomen longer than head and complete thorax combined. Median segment may be distinctly shorter (apterous insects), equal in length, or longer than the metanotum (alate insects). Lateral margins of abdominal tergites II–VII of && often roundly or uniformly expanded. Sternum VII of && lacking a praeopercular organ. Gonapophyses of && may be slightly elongated but never project considerably over apex of abdomen. Subgenital plate of && boat-like, distinctly and acutely keeled in basal portion. Length and shape variable, ranging from small, scoop-shaped to distinctly boat-shaped but usually not exceeding apex of anal segment, rarely elongated, lanceolate and projecting over abdomen. Anal segment of %% laterally compressed, tectiform and consisting two semi-tergites, which either have the interior surface with a dentate longitudinal bulge, or have the ventro-apical angle elevated, in-curving and armed with several terminal teeth. External vomer of %% lacking. Supraanal plate very small. Cerci ± prominently enlarged, laterally compressed, ± dilated and lan-



37

ceolate or foliaceous. Profemora indistinctly triangular in cross-section with the two dorsal carinae ± approaching each other, compressed and curved basally; mostly unarmed. Anterodorsal carina slightly raised and usually unarmed, rarely with minute serrations. Medioventral carina present, moderately distinct and about midways on ventral surface of profemur. Mid- and hind legs often with prominent armature, the dorsal carinae mostly smooth. Meso- and metafemora trapezoidal in cross-section with anterior carinae ± decidedly nearing, medioventral carina present, often spinose. Tibiae trapezoidal to almost triangular in cross-section, with the dorsal carinae strongly nearing each another. No area apicalis on tibiae. Basitarsus distinctly longer than following tarsomere. Doubtful genera: The systematic position of Didymuria Kirby, 1904 (Type species: Phasma violescens Leach, 1814: 26, by original designation; Fig. 68) is doubtful. It was previously placed in Tropidoderinae: Tropidoderini by all former authors (e.g. Günther, 1953 and Bradley & Galil, 1977: 195). But although the insects are remarkably similar in appearance and habitus to e.g. Tropidoderus the closed internal micropylar plate of the eggs clearly separates Didymuria from Tropidoderini, since the type of internal micropylar plate is uniform for an individual tribe (→ 3.3 and Clark-Sellick, 1998). The fact that both genera occur in the same habitats in southeast Australia supposes the similar habitus and colouration may be the result of convergent adaption to similar ecological pressure. Egg morphology and certain features of the insects, such as the conspicuously broadened metafemora of %% which bear several prominent spines on their medioventral carina, suppose relationship to the tribe Gigantophasmatini trib. nov.. Consequently, Didymuria is here retained in Tropidoderini but with doubt. Any broader discussion on the exact systematic position of the genus requires the availability of sufficient material, careful investigation and comparison of the exosceleton and may find support by genetic analyses. A DNA-barcoding project of predominantly Australian Phasmatodea is in progess by Brock & Mantovani (personal communication with P. D. Brock). The rarely known genus Lysicles Stål, 1877 (Type species: Lysicles hippolytus Stål, 1877b: 65, by monotypy) was originally described as being close to Tropidoderus and since attributed to Tropidoderini. Photos of the % and & of an unidentified species of Lysicles from Australia were kindly provided for examination by P.D. Brock (Brockenhurst, England). Several features strongly violate the diagnostic features of Tropidoderini, e.g. the very slender and cylindrical body, elongate mesothorax which is roughly 2x longer than the head and pronotum combined, strongly shortened tegmina and alae of &&, plain transparent anal region of the alae of %%, very short subgenital plate of && which hardly projects over the posterior margin of abdominal tergite IX, and dorsally dentate meso- and metafemora of &&. On the other hand, the shortened tegmina and alae of && and broadened metafemora of %%, which bear two or three prominent long spines on their medioventral carina, resemble the Australian Didymuria and the tribe Gigantophasmatini trib. nov.. Unfortunately, the eggs of Lysicles are as yet unknown which makes any confirmed decision on its systematic position impossible. Apparently more material and eggs are required for any broader discussion. A re-description of the genus and illustrations of L. hippolytus Stål will be provided in a field guide of the Australian phasmids by Brock & Hasenpusch (personal communication with P.D. Brock). Distribution (Fig. 1): Australia, New Guinea, New Caledonia and Loyalty Islands as well as Islands of the Indian Ocean (Réunion, Mauritius, Madagascar and Comoros). Tribes included: 1. Tropidoderini Brunner v. Wattenwyl, 1893: 97. Type genus: Tropidoderus Gray, 1833: 13, 31. 2. Monandropterini Brunner v. Wattenwyl, 1893: 97. Type genus: Monandroptera Audinet-Serville, 1838: 242. 3. Gigantophasmatini trib. nov. Type genus: Gigantophasma Sharp, 1898: 87.


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FIGURES 20–31. Terminal abdominal segments of %% of Clitumninae and Lonchodinae [scale = 5 mm] 20–22. Tribe Pharnaciini: Tirachoidea jianfenglingensis (Bi, 1994), 20. lateral view, 21. dorsal view, 22. ventral view 23–25. Tribe Clitumnini: Cuniculina cuniculus (Westwood, 1859), 23. lateral view, 24. dorsal view, 25. ventral view 26–28. Tribe Medaurini trib. nov.: Medaura jobrensis (Brock & Cliquennois, 2000), 26. lateral view, 27. dorsal view, 28. ventral view 29–31. Subfamily Lonchodinae: Lonchodes brevipes Gray, 1835), 29. lateral view, 30. dorsal view, 31. ventral view

4.4.1 Tropidoderini Brunner v. Wattenwyl, 1893 (Figs. 5–10, 33, 39, 50–51, 70–71) Type genus: Tropidoderus Gray, 1835: 31. Tropidoderi Brunner v. Wattenwyl, 1893: 97. Tropidoderini Bradley & Galil, 1977: 195. Otte & Brock, 2005: 13. Podacanthini Günther, 1953: 553. Phibalosomini (III. Sectio: Diurae) Redtenbacher, 1908: 379 (in part).

Description: The six genera currently attributed to Tropidoderini share several common features and form a rather uniform group of moderately sized (body lengths %% 43–130 mm, && 53–163 mm), bright green often



39

very colourful Phasmatinae with large tegmina and well developed alae covering most of the abdomen (Figs. 70 & 71). All representatives are rather robust for the subfamily with the body sub-cylindrical in cross-section and slightly compressed dorsoventrally and the meso- and metasternum conspicuously flattened. %% are slightly smaller but much more slender than &&, the abdomen except the three basal segments being of uniform width. In && the abdomen is broadened basally or medially and may have the lateral margins of tergites II–VII expanded to a various degree (Tropidoderus). The head is oval to slightly globose and armed. Ocelli are either well developed (Podacanthus, Parapodacanthus and Micropodacanthus), conspicuously reduced (Paratropidoderus) or lacking (Tropidoderus and Malandania). The antennae are at best a little longer than the head and thorax combined but often conspicuously shorter in &&. They are slender and filiform with the antennomeres distinctly longer than wide (Parapodacanthus and Micropodacanthus) to rather robust with the antennomeres considerably shortened (e.g. Tropidoderus and Paratropidoderus). The mesothorax is conspicuously widened towards the posterior (&& in particular) and often remarkably short being at best 1.3x longer than the head and pronotum combined (Tropidoderus and Paratropidoderus) but sometimes just hardly longer than the pronotum (e.g. Podacanthus). Often the mesonotum bears a number of tubercles or ± distinct spines. The meso- and metapleurae (&& in particular) are angulate and have the lower outer margin gently elevated and set with a longitudinal row of spiniform tubercles or distinct spines. The meso- and metasternum are strongly flattened and minutely granulose but destitute of any enlarged tubercles or spines. The median segment is considerably longer than the metanotum and the abdomen at least 1.5x longer than the head and complete thorax combined. The alae are well developed and at least reach to abdominal segment VII, the anal region being either plain transparent (Tropidoderus and Paratropidoderus), plain pink (e.g. Podacanthus) or tesselated with pink (Micropodacanthus). Abdominal sternum VII of && lacks a praeopercular organ. The semi-tergites of the anal segment of %% are rather broad with the apex broadly rounded to truncate (e.g. Tropidoderus and Paratropidoderus, Figs. 5–7), or have the lower posterior angles elevated into a conspicuous slender, strongly in-curving, finger-like process (Podacanthus, Micropodacanthus and Parapodacanthus, Figs. 8–10). Their interior surfaces bear a distinct, longitudinal keel or bulge which is armed with a row of prominent teeth (“Dornenfeld”, Figs. 33 & 39), or the finger-like projections each bear several terminal teeth. An external vomer in the %% is lacking (Fig. 39). The cerci are strongly laterally compressed, carinate dorsally and ventrally and ± distinctly lanceolate; with the exception of Paratropidoderus they considerably exceed the anal segment in length. The subgenital plate of && is strongly longitudinally keeled, boat-shaped and at best slightly projects over the apex of the abdomen (Tropidoderus). The legs are relatively short and rather robust, with the ventral carinae of the mid and hind legs dentate or spinose to a various degree; dorsal carinae unarmed (at least of tibiae). Tropidoderus has the anteroventral carina of the meso- and metafemora roundly elevated in &&. The profemora are unarmed and indistinctly triangular in cross-section with the anterodorsal carina just gently raised. The posteroventral carina either smooth (e.g. Parapodacanthus) or sparsely set with a few very minute spines (Tropidoderus and Paratropidoderus). The medioventral carina of the profemora is very indistinct, unarmed and roughly midways on the ventral surface of the femur. Eggs (Figs. 50–51): In most aspects eggs are very similar to those of Phasmatinae: Phasmatini. The capsule is longer than wide, roughly ovoid and usually strongly sculptured (Figs. 50a & 51a), certain taxa having the capsule densely covered with hair-like structures (e.g. Parapodacanthus and certain species of Podacanthus). The operculum bears a conspicuous knob-like capitulum, either on a very short and indistinct stalk or strongly appressed to the operculum. The micropylar plate is elongate, oval to lanceolate and less than half as long as the capsule. Internally it is open with a very narrow posteromedian excavation or slit and lacks a median line (Figs. 50b & 51b). Comments: Within his Acrophyllidae Brunner v. Wattenwyl (1893: 97) first united the genera Tropidoderus, Podacanthus, Diura Gray, 1833 (= Didymuria Kirby, 1904) and Extatosoma Gray, 1833 in a tribe termed Tropidoderi. A separate tribe, the Extatosomatini, was established by Clark-Sellick (1997: 103) for the latter genus (→ 4.4.3), which differs considerably from all other Tropidoderini by e.g. the conical head, spi-


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nose and lobate body, lobate legs and features of the egg-morphology. Podacanthini Günther, 1953 is a junior synonym of Brunner v. Wattenwyl’s Tropidoderini (see Bradley & Galil, 1977: 194). Based on the morphology of the insects two generic groups can be distinguished amongst present Tropidoderini. The first group, which contains Tropidoderus, Paratropidoderus and Malandania, is characteristic for the rather elongate and flat head, lack of ocelli, mesothorax which is longer than the head and pronotum combined, small poculum and slender, apically rounded anal semi-tergites of the %%. The second group contains all remaining genera and is distinctive for the globose head, distinct ocelli, strongly shortened mesothorax, which is shorter than the head and pronotum combined, more bulgy poculum and in-curving, finger-like ventro-apical process of the anal semi-tergites of %%. Several taxa of Tropidoderini remarkably resemble certain genera of Phasmatinae: Phasmatini, which indicates relation between these two groups. For instance, in view of ther general habitus, body length relations, wing length and colouration && of Malandania strongly resemble those of Paracyphocrania Redtenbacher, 1908 (tribe Phadmatini) from Sulawesi. A similar case is presented by Tropidoderus or Paratropidoderus and the well-known Australasian genus Eurycnema Audinet-Serville, 1838 (tribe Phasmatini). These genera resemble each other by e.g. the short antennae and shortened antennomeres, similar relative length of the mesothorax, plain transparent anal region of the alae and green general colouration. In both cases differences between these genera generally restrict to the key features of Tropidoderini such as the unarmed profemora, short mesothorax, unarmed dorsal carinae of the meso- and metatibiae and flattened, unarmed meso- and metasternum. The elongated, finger-like ventro-apical projections of the anal semi-tergites of %% of Podacanthus and Parapodacanthus (Figs. 8–10) moreover resemble the elongated semi-tergites seen in most Clitumninae (tribes Clitumnini and Pharnaciini). The genus Vasilissa Kirby, 1896 was removed from Tropidoderini and transferred to the subfamily Phasmatinae by Brock & Hasenpusch (2007: 74). Distribution: Australia, endemic. Genera included: 1. Malandania Sjöstedt, 1918: 48. Type species: Malandania pulchra Sjöstedt, 1918: 49, pl. 7: 1, by original designation. 2. Micropodacanthus Brock & Hasenpusch, 2007: 54. Type species: Micropodacanthus sztrakai Brock & Hasenpusch, 2007: 58, figs. 103–113, by original designation. 3. Parapodacanthus Brock, 2003: 31. Type species: Parapodacanthus hasenpuschorum Brock, 2003: 32, figs. 46–53 & 60–62. 4. Paratropidoderus Brock & Hasenpusch, 2007: 60. Type species: Paratropidoderus spinosus Brock & Hasenpusch, 2007: 61, figs. 114–117, by original designation. 5. Podacanthus Gray, 1833: 28. Type species: Podacanthus typhon Gray, 1833: 17, pl. 2: 1, by monotypy. 6. Tropidoderus Gray, 1835: 31. Type species: Trigonoderus childrenii Gray, 1833: 18, pl. 3: 1, by indication. [Replacement name for Trigonoderus Gray, 1833] = Athertonia Sjöstedt, 1918: 45. Type species: Athertonia prasina Sjöstedt, 1918: 46, pl. 7: 7a–b, by monotypy. [Synonymised by Balderson, Rentz & Roach, 1998: 375] = Kimberleyana Sjöstedt, 1918: 46. Type species: Kimberleyana gracilifemur Sjöstedt, 1918: 46, pl. 7: 8, by monotypy. [Synonymised by Balderson, Rentz & Roach, 1998: 375] = Trigonoderus Gray, 1833: 18. Type species: Trigonoderus childrenii Gray, 1833: 18, pl. 3: 1, by monotypy. [Preoccupied by Trigonoderus Westwood, 1832 (Hymenoptera: Cleonymidae)]



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FIGURES 32–39. Anal segments of %% [scale = 0.5 mm] TIX = tergite IX; TX = tergite X (anal segment); Ce = cercus; Pa = paraproct; Ep = epiproct (supraanal plate); Vo = vomer; DoF = “Dornenfeld”. 32. Tribe Phasmatini: Acrophylla wuelfingi (Redtenbacher, 1908), caudal view 33. Tribe Tropidoderini: Tropidoderus childrenii Gray, 1833, caudal view 34. Tribe Monandropterini: Monandroptera acanthomera (Burmeister, 1838), caudal view 35. Tribe Pharnaciini: Tirachoidea jianfenglingensis (Bi, 1994), caudal view 36. Tribe Clitumnini: Cuniculina cuniculus (Westwood, 1859), caudal view 37. Tribe Stephanacridini: Hermarchus pythonius (Westwood, 1859), caudal view 38. Tribe Phasmatini: Acrophylla wuelfingi (Redtenbacher, 1908), ventral view 39. Tribe Tropidoderini: Tropidoderus childrenii Gray, 1833, ventral view


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4.4.2 Monandropterini Brunner v. Wattenwyl, 1893 (Figs. 1, 14–16, 34, 42, 45, 52–53, 74) Type genus: Monandroptera Audinet-Serville, 1838: 245. Monandropterae Brunner v. Wattenwyl, 1893: 97. Monandropterini, Bradley & Galil, 1977: 195. Cliquennois & Brock, 2004a: 42. Otte & Brock, 2005: 13. Rhaphiderini Günther, 1953: 553.

Description: Small to rather large representatives of the Tropidoderinae (body lengths %% 45.5–138.0 mm, && 58.0–168.5 mm). Body sub-cylindrical and of almost uniform width in %%, oval in cross-section, ± broadened with the abdomen narrowing towards the posterior in && (Figs. 74a, 74b). The head is globose to almost spherical and unarmed. Ocelli are lacking. The antennae are at best slightly longer than the head, pro- and mesonotum combined, the scapus being simple and slender. The mesothorax is at least 1.3x longer than the head and pronotum combined, but never considerably elongated; in && distinctly widening towards the posterior. The meso- and metapleurae (&& in particular) have the lower outer margin gently elevated and set with a longitudinal row of spiniform tubercles or distinct spines. Often a cluster of tubercles or spines is present in the anterior portion of the mesonotum. The median segment is a little shorter than the metanotum. && are apterous or have very small, vestigial tegmina. %% of Heterophasma and Rhaphiderus are apterous, only the type genus Monandroptera having alate %%, which have the anal region transparent and tesselate with darker colour in a similar manner to certain Phasmatini; the alae at best extending to abdominal segment VI. && lack a praeopercular organ on sternum VII and have the abdomen ± prominently widened towards the base; occasionally lateral margins of abdominal tergites are roundly elevated (Monandroptera). The anal segment of %% is very similar in structure to those seen in Tropidoderini and Phasmatini, the semi-tergites being short and broad with the posteroventral angles roundly swollen. The interior surfaces are apically rather irregularly set with several small teeth (= “Dornenfeld”, Figs. 34 & 42). %% possess a very prominent, strongly elongated and gently curved, papillate phallus, which reaches as far back as to the posterior margin of the poculum. The cerci are strongly laterally compressed and carinate but usually shorter than the anal segment (conspicuously enlarged, foliaceous and with a spine-like posterior process in %% of Monandroptera). An external vomer is lacking (Fig. 42). The subgenital plate of && is very small, scoop-shaped, just slightly keeled in the basal portion and at best reaches to the posterior margin of tergite IX. The gonapophyses are well developed, exceed the apex of the subgenital plate and are used to hold the egg by the opercular extension before it is flicked away. The profemora are indistinctly triangular in cross-section with the two dorsal carinae moderately approaching each other. The anterodorsal carina is not conspicuously raised nor serrate and the posterodorsal carina not displaced towards the lateral surface of the femur. The medioventral carina of the profemora is represented by an indistinct longitudinal keel midways on the ventral surface of the femur (Fig. 45). Dorsal carinae of mid and hind legs unarmed. Eggs (Figs. 52–53): Eggs are similar in general appearance to those of Tropidoderini. The capsule is ovoid, longer than wide, slightly laterally compressed and narrowed towards the anterior. The micropylar plate is small, suboval, pointed anteriorly, covers less than half of the capsule length and is slightly displaced towards the polar end of the capsule (Figs. 52a, 52b). Internally it is open and lacks a median line (Figs. 52b, 53b). In contrast however, the operculum is conically elevated in the centre but lacks a stalked capitulum. Comments: Günther (1953: 553) originally established Rhaphiderini as a tribe of Podacanthinae Bradley & Galil (1977: 195) replaced Günther’s tribal name to the senior available family-group name Monandropterini and characterized it by the following characters: & apterous or with rudimentary tegmina, % apterous or alate; in latter case inner carina of ventral surface of posterior tibiae of % spinose. Bradler (2000) included Monandropterini in “Lanceocercata” and thus was the first author who recognized the close relation to e.g. the REVISION OF PHARNACIINI


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subfamily Phasmatinae. A detailed revision of Monandropterini at the species level was published by Cliquennois & Brock (2004a). Relationship between Monandropterini and Tropidoderini is obvious and was already suggested by most former authors (e.g. Brunner v. Wattenwyl, 1893; Redtenbacher, 1908; Günther, 1953 or Bradley & Galil, 1977). This is emphasized by the sub-cylindrical to oval cross-section of the body, short mesothorax which is widened towards the posterior (&& in particular), spinose meso- and metapleurae, basally broadened abdomen of &&, occasionally laterally dilated abdominal tergites, just indistinctly triangular cros-section of the profemora, and unarmed dorsal carinae of all femora and tibiae. However, Monandropterini frequently differs from Tropidoderini by the convex meso- and metasternum (conspicuously flattened in Tropidoderini), distinct phallus of %% and very short, scoop-like subgenital plate of &&. The eggs furthermore differ by lacking a stalked capitulum. Furthermore, close relationship obviously exists with the Gigantophasmatini trib. nov., a tribe only known from a single genus endemic to the Loyalty Islands (near New Caledonia). With this tribe the type genus Monandroptera shares the lack of wings in &&, dorsal tooth of the cerci and distinct ventral spines on the medioventral carina of the metafemora of %%. Monandropterini however differs from this peculiar tribe by e.g. the small cerci of && and open internal micropylar plate of the eggs. The presence of representatives of Tropidoderinae in Madagascar, the Comoros, Réunion and Mauritius may be explained by a fairly early settlement, perhaps during the middle or late Jurassic, via Australia or Antarctica. The evolution of several features shows convergence with the corresponding features in other nonrelated taxa distributed in the same region, which supposes a strong selection pressure. The convergent evolution of characters in three non-related taxa from Mauritius has already been discussed by Cliquennois & Brock (2004b). Distribution (Fig. 1): Monandropterini is restricted to the islands of the Western Indian Ocean and endemic to Réunion and Mauritius. The literary records of Monandropterini from Madagascar and the Comoros are doubtful and believed to be erroneous (Cliquennois, 2007: 12). No representatives have so far been recorded from the Seychelles or the African continent. Genera included: 1. Heterophasma Redtenbacher, 1908: 386. Type species: Heterophasma gaudichaudi Redtenbacher, 1908: 386, pl. 19: 2, 2a, 2b, by monotypy. 2. Monandroptera Audinet-Serville, 1838: 242. Type species: Monandroptera inuncans Audinet-Serville, 1838: 244, by monotypy. = Craspedonia Westwood, 1841: 25. Type species: Diapherodes (Cranidium) undulata Westwood, 1843: 49, by monotypy. [Synonymised by Westwood, 1859: 80] 3. Rhaphiderus Audinet-Serville, 1838: 245. Type species: Bacteria scabrosa Percheron, 1829, pl. 53: 4 (in: Guérin-Méneville, F. E.), by monotypy.

4.4.3 Gigantophasmatini trib. nov. (Figs. 11–13, 66, 69) Type genus: Gigantophasma Sharp, 1898: 87, by present designation.

Description: Large to very large (body length %% 90–100 mm, && 163–187 mm), moderately slender Tropidoderinae with strong sexual dimorphism. && apterous (Fig. 69a), %% with scale-like tegmina and well developed alae (Fig. 69b). Body surface of both sexes slightly glabrous and smooth except for a few small tubercles on the mesonotum, meso- and metasternum and a marginal row of spine-like tubercles along the meso- and


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metapleurae. && with very distinctive colouration, being brown with the greatest portions of the dorsal surfaces of the meso-, metanotum, and abdominal tergites I–VII creamish white or amber. Head ovoid, about 1.3x longer than wide and slightly narrowing behind the eyes. Vertex gently rounded and unarmed. Ocelli lacking. Antennae slightly shorter (&&) or a little longer (%%) than the head, pro- and mesonotum combined; consisting of about 30 antennomeres. Pronotum rectangular and slightly narrower than the head. Mesothorax about 1.8x longer than head and pronotum combined, constricted anteriorly and gently broadened towards the posterior in &&. Metanotum rectangular and about 2.2x longer than wide in &&. Tegmina of %% about as long as metanotum, the alae projecting to abdominal tergite VI. Anal region of alae transparent. Abdomen excluding median segment longer than head and complete thorax combined. Median segment less than half the length of metanotum in && and considerably longer than the metanotum in %%. Abdominal tergites II–VII of && slightly increasing in length with the three basal segments rougly as long as wide, each tergite swollen and with the lateral margins roundly expanded. Tergites II–VII of %% slender, parallel-sided, of uniform width and considerably longer than wide. Sternites II–VII unarmed, a praeopercular organ in && is missing. Tergites VIII–X much narrower than previous and roughly parallel-sided in && and broader than previous in %%. Anal segment of %% strongly tectiform and forming two semi-tergites, which have the ventro-apical angle elevated, in-curving and interiorly armed with several terminal teeth (Fig. 12). External vomer of %% lacking (Fig. 13). Cerci of both sexes laterally flattened, strongly enlarged and considerably longer than the anal segment; in %% with a distinct triangular tooth on the dorsal carina. Poculum of %% deep, cup-like and roughly reaching the posterior margin of tergite IX (Fig. 11). Subgenital plate of && with the basal portion very strongly convex and keeled and the apical portion elongated, lanceolate and extending over the apex of the abdomen by more than the combined length of tergites IX and X. Legs moderately long and slender with most carinae dentate. Profemora longer than mesothorax and slightly triangular in cross-section with the anterodorsal carina very slightly raised. The anterodorsal and posteroventral carinae minutely and sparsely denate in &&, unarmed in %%. The medioventral carina moderately distinct and midways on the ventral surface of the profemur. Mid and hind legs trapezoidal in cross-section with dorsal carinae nearing. Metafemora conspicuously broadened and on the medioventral carina armed with four prominent and long, pointed spines; a further prominent subapical spine is present on the posteroventral carina. Basitarsi longer than following two tarsomeres combined, the dorsal carina very gently rounded in &&. Eggs (Fig. 66): Small (overall length 3.7–3.8 mm), ovoid and roughly circular in cross-section (Fig. 66a). Capsule surface very minutely rugose. Micropylar plate elongate, about ¾ as long as the capsule with both ends pointed and slightly constricted below the micropylar cup. Internally the plate is closed with the micropylar stalk placed well away from the posterior end of the plate (Fig. 66b). Operculum flat with a large, knoblike but strongly appressed central capitulum. Comments: Sharp (1898: 87) originally placed Gigantophasma from the Loyalty Islands in close relation to the genus Monandroptera Audinet-Serville, 1838, now in the tribe Monandropterini. Although relationship to Monandropterini and the subfamily Tropidoderinae is obvious, neither subsequent author followed Sharp’s original placement of the genus. Redtenbacher (1908) listed it amongst his Acrophyllini and in close relation to Hermarchus Stål, 1875 (here transferred to the tribe Stephanacridini, → 4.6.2). Günther (1953: 555) placed Gigantophasma in the tribe Pharnaciini (subfamily Phasmatinae), which was overtaken by all subsequent authors (e.g. Bradley & Galil, 1977 and Otte & Brock, 2005). Gigantophasma is however definitely not a member of Pharnaciini (here transferred to Clitumninae, → 5.) and here excluded from this tribe, due to strongly violating the diagnostic features of Pharnaciini. The foliaceos cerci and charcteristic shape of the anal semi-tergites of the %%, which have the ventro-apical angles protuded and armed with several terminal teeth, obviously place Gigantophasma in the family Phasmatidae s. str. (= Lanceocercata). Although the insects show remarkable similarity to the genus Monandroptera (tribe Monandropterini) in several aspects, the large foliaceous cerci and lanceolate subgenital plate of &&, features of the eggs such as the closed internal micropylar plate, and the far off, rather isolated distribution in the Loyalty Islands clearly



45

distinguish Gigantophasma from this tribe. Features shared with Monandropterini, and the type genus Monandroptera in particular, are the lack of wings, broadened body and laterally expanded abdominal tergites of &&, and thickened metafemora of %% which have the medioventral carina armed with several prominent spines. Consequently, since the type of internal micropylar plate (open or closed) is uniform within a unique tribe, and the fact that Gigantophasma clearly differs from its closest relative, the Monandropterini, by numerous obvious morphological features of the insects, the new tribe Gigantophasmatini is here proposed to include this peculiar genus. The broadened and ventrally armed metafemora and features of the egg morphology such as the closed internal micropylar plate indicate relationship to the Australian genera Didymuria Kirby, 1904 and Lysicles Stål, 1875, both of which are here left in an unknown position within the Tropidoderinae (→ 4.4). Distribution (Fig. 1): Loyalty Islands, endemic. Genus included: 1. Gigantophasma Sharp, 1898: 87. Type species: Gigantophasma bicolor Sharp, 1898: 87, pl. 7: 6, by subsequent designation of Kirby, 1904a: 386.

4.5 Extatosomatinae Clark-Sellick, 1997 stat. nov. (Figs. 1, 17–19, 63, 433) Extatosomatini Clark-Sellick, 1997: 103. Otte & Brock, 2005: 33.

Description: Large (body length %% 80–115 mm, && 100–165 mm), the body broad and very massive in &&, moderately slender in %%. && with tegmina and vestigial alae, %% with fully developed alae roughly reaching apex of abdomen. Head prognathous with the vertex strongly conically raised and spinose. Ocelli lacking in &&, but prominent in %%. Antennae at best as long as the head, pro- and mesonotum combined in &&, much longer than the head and entire thorax and strongly pubescent in %%; with less than 35 segments. Pronotum narrower than the head and rectangular to slightly trapezoidal, soinose in &. Mesothorax about as long (%%) or shorter than the head and pronotum combined (&&); strongly widened towards the posterior in && with the posterior portion roughly 3x wider than anterior margin. Mesonotum slightly trapezoidal and in && with a posteromedian pair of spines or spinose lamellae. Mesosternum moderately convex and at best with few slender spines. Metasternum flat and smooth (%%) or set with a few scattered spines (&&). In && tegmina scalelike, with the anterior margin angulate and rougly reaching to posterior margin of metanotum; alae vestigial, shorter than tegmina and completely or in great parts covered by tegmina. %% with tegmina reaching about half-way along median segment, and alae well developed, reaching to about apex of abdomen. Anal region transparent with slight brownish mottling. Median segment slightly shorter than the metanotum in &&, considerably longer in %%. Abdomen excluding median segment longer than head and complete thorax combined. Abdomen slender in %% with tergites II–VII > 1.5x longer than wide and unarmed; V–VII with large, roundly angulate lateral lobes. Abdomen of && very massive with the basal tergites transverse and V–VII bearing very large and spinose, foliaceous lateral lobes. Tergites II–VII each with one or two pairs of spinose lamellae and tergites VIII–X considerabyl narrower than previous. Sternites II–VII strongly rugulose in %% and all over set with long, slender spines in &&. Praeopercular organ lacking in &&. Aanal segment of %% strongly tectiform, consisting of two semi-tergites which have the ventro-apical angles expanded into a short and in-curving finger-like process which bears 5–6 distinct terminal teeth (Figs. 17–19). External vomer in %% lacking. Cerci rather small, laterally compressed, carinate and slightly lanceolate, about as long (%%) or considerably shorter (&&) than the anal segment. Poculum of %% very bulgy, deep and cup-like. Subgenital plate of && boat-shaped, acutely longitudinally keeled and roughyl reaching apex of the abdomen; basal por-


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tion of the subganital plate with the median keel strongly raised and dentate. Gonapophyses of && elongate, filamentous, up-curving apically and slightly extending over apex of subgenital plate. Legs all rather short, robust and furnished with large, dentate ± foliaceous appendages and spines. Profemora trapezoidal in crosssection with dorsal carinae strongly nearing and medioventral carina lacking; all carinae with ± distinct dentate lobes. The remainder femora and all tibiae triangular in cross-section with the anterior carinae melted and forming one or two large, dentate foliaceous lobes. Anteroventral carinae of mid and hind legs roundly elevated and dentate, the posteroventral carinae spinose, that of the metatibiae in particular armed with prominent hooked spines. The dorsal and anteroventral extensions of the mid and hind legs together form a large, almost plain lateral surface of the corresponding femur or tibia. Medioventral carina of meso- and metafemora very faint but set with a few small spines. Probasitarsus about as long as following three tarsomeres combined and with a ± apical dorsal lobe in &&. Meso- and metabasitarsi slightly shorter. Eggs (Fig. 63): Moderately sized (overall length 4.5–6.5 mm), capsule ovoid with the polar-area slightly narrowed. Capsule surface very gently rugose or punctured and ± glabrous. Colouration variable, usually pale cream or grey and irregularly marbled with brown or dull green. Micropylar plate very elongate, parallelsided, extending from the opercular rim to the posterior pole of the capsule and set conspicuously above the capsule surface; pale colour strongly contrasting with the capsule. Various degrees of expansion are seen in the region of the micropylar cup. Internal micropylar plate closed with the micropylar stalk well within the borders of the plate (Fig. 63b) and connected to the surface with an inwardly turning lip (Clark-Sellick, 1997: 103). Operculum convex and with a prominently raised, irregulary cone-shaped or funnel-like capitulum. Comments: Both, Günther (1953: 553) and Bradley & Galil (1977: 195) omitted to mention Extatosoma in their taxonomic arrangement of the order and since the genus had been treated as a member of Tropidoderinae: Tropidoderini. Based on the striking features of the insects, such as the strongly conically elevated head, foliaceous appendages of the abdominal tergites and large leaf-like extensions of the femora and tibiae, as well as egg-structures Clark-Sellick (1997: 103) removed Extatosoma from Tropidoderini and established the tribe Extatosomatini to contain it. Due to the laterally compressed cerci, distinct clasping apparatus of the anal segment of %% and derived copulation position (Fig. 433) in which %% grasp the base of the && subgenital plate, Brader (2001: 181) contained Extatosomatini in Lanceocercata. Indeed, the mentioned features show close relationship to taxa of this group, hence Extatosomatini must be regarded a member of Phasmatidae s. str.. Indeed, the following striking features at once distinguish the highly cryptic Extatosomatini from entire Tropidoderinae and well characterise it amongst the family Phasmatidae s. str.: head prognathous with vertex strongly conically raised and spinose; abdominal tergites of && with paired spinose lamellae dorsally; tergites V–VII with greatly expanded, leaf-like lateral lobes (spinose in &&); abdominal sternites II–VII of && spinose; subgenital plate of && with the longitudinal keel dentate basally; meso- and metafemora and all tibiae distinctly triangular in cross-section with dorsal carinae melted and forming very large, dentate lobes. Furthermore, && exhibit the remarkable ability to curl over their abdomen with live insects usually found in a very characteristic position, and eggs have the very long micropylar plate conspicuously set above the capsule surface. Consequently, as these features, most of which may represent apomorphies, cleary distinguish Extatosomatini from Tropidoderinae and any other subfamily of Phasmatidae s. str., Extatosomatini is here raised to subfamily level. Since Extatosomatinae is monotypic the tribe Extatosomatini is technically not needed anymore. The rather basal ground pattern of Extatosomatinae, like the remarkably prognathous head, presence of distinct ocelli in the %% and short thorax, suggest this group must have splitted off rather early from the remaing Phasmatidae. Mainly in behavioural aspects Extatosomatinae however shows to be highly specialized, which concerns to the ability of adult && to curl their strongly procryptic abdomen forward over the ir back and symbiosis of the Australian E. tiaratum Mac Leay, 1826 with ants of the genus Leptomyrmex Mayr, 1821 (Formicidae: Dolichoderinae) in its natural habitats (Key, 1991 and Hughes & Westoby, 1992). The ability to



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curl the abdomen is observed in nymphs of many Phasmatodea, but none retain this ability in such degree in the adult insects. The && of several species (e.g. of the genus Eurycnema Audinet-Serville, 1838) slightly curl up their five terminal segments, but in Extatosomatini the complete abdomen is curled forward from the 2nd segment onwards, with the apex of the abdomen almost touching the dorsal surface of the body near the wings or median segment. This peculiar behaviour is supported by extensive membranes between the abdominal segments and requires a strong and specialized dorsal and ventral musculature internally. In nymphs the curled up abdomen is perhaps a specialisation due to the symbiosis with ants that are known to be used as a host to hatch the eggs in their formicary. The ants are attracted by the nutritious capitulum as a food source with the rest of the egg discarded in their nest. This benefits the phasmids as it helps dispersal and a decreases the rate of parasitism. The aposematic colouration of black, with an orange head and white collar on the mesothorax, in the newly hatched nymphs remarkably mimics the Leptomyrmex ants in colour, posture and movements, and thus insures the nymphs to escape from the formicary safely (Key, 1991: 402). This symbiosis has also been reported from other Australian species whose eggs bear a capitulum (Hughes & Westoby, 1992) but in neither species the nymphs represent “ant mimics”. A revision of the genus Extatosoma at the species level was provided by Brock (2001). Distribution (Fig. 1): East Coast of Australia and New Guinea. Genus included: 1. Extatosoma Gray, 1833: 23. Type species: Phasma tiaratum Mac Leay, 1826: 455, by subsequent designation of Kirby, 1904a: 380. = Ectatosoma Gray, 1835: 29. [Incorrect subsequent spelling]

4.6 Problematic taxa The following two subfamilies are obviously polyphyletic and deserve more detailed investigation to make any confirmed decision on their position within the Phasmatidae s. str.. Certainly the type-genera of both subfamilies belong in Phasmatidae (= Lanceocercata), but other genera contained in either subfamiliy strongly violate the diagnostic features, e.g. by %% having a well developed and sclerotised vomer, or lacking the clasping apparatus of the anal segment. Below brief critical discussions of these subfamilies, namely Pachymorphinae and Xeroderinae, are provided and emphasize on highlighting the differences between the taxa contained. 4.6.1 Subfamily Pachymorphinae Brunner v. Wattenwyl, 1893 The subfamily Pachymorphinae as interpreted by Karny (1923: 235) and subsequent authors (Günther, 1953; Bradley & Galil, 1977 and Otte & Brock, 2005) is obviously polyphyletic which was already suggested by various authors (e.g. Tilgner, 2002; Whiting et al., 2003 and Cliquennois, 2004). Although Karny (1923) and Günther (1953) placed Pachymorphinae in the family Phasmatidae or Phasmidae respectively, Bradley & Galil (1977) contained it in Heteronemiidae (= Diapheromeridae Kirby, 1904). The subfamily was traditionally characterised amongst Heteronemiidae by having short antennae and being apterous in both sexes. Three tribes were attributed to Pachymorphinae by Bradley & Galil (1977: 184): 1) Pachymorphini Günther, 1953 includes the type genus Pachymorpha Gray, 1835 and Acanthoderus Gray, 1835 both from Australia, two New Zealandean genera, as well as a doubtful genus from Sri Lanka, 2) Hemipachymorphini Günther, 1953 contains the type genus Hemipachymorpha Kirby, 1904a from South Africa, one genus from New Guinea and two from New Zealand, and 3) Gratidiini Bragg, 1995 (= Ramulini Günther, 1953) comprises small to moderately sized, mostly very slender and plain stick-insects, with slender and unarmed legs. Sections of the latter


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tribe were suggested to be closely related to Clitumnini (→ here transferred to the subfamily Clitumninae) by Tilgner (2002) and Whiting et al. (2003). This however requires further evaluation. The laterally flattened cerci and genital structures of %% appear to place the tribe Pachymorphini in Lanceocercata (see Bradler, 2001: 183). The exact systematic position and true relationships of the type genus, the Australian Pachymorpha Gray, 1835 (Type species: Pachymorpha squalida Gray, 1835: 28) however remain as yet questionable. Brock & Hasenpusch (2005: 48) stated that the eight other species from Africa and Asia were unlikely to belong in the genus. The egg of P. squalida was illustrated by Key (1991: 397, fig. 25.4: K) and shows strong resemblance to eggs of certain taxa of Phasmatinae, being ovoid with a small, knob-like capitulum and having an open internal micropylar plate (obviously seen on Key’s figure). Moreover, similarity concerning to the morphology of the insects is in particular seen in the shape and structure of the antennae and genitalia. This suggests relation to Phasmatinae and may place Pachymorphini as a tribe of that subfamily, but deserves further research and examination of sufficient material. The Sri Lankan Miniphasma Zompro, 2007 is unlikely to be closely related to Pachymorpha Gray, and relationship of Pachymorpha to the two New Zealandian genera Micrarchus Carl, 1913 and Niveaphasma Jewell & Brock (2003) and the Australian Acanthoderus Gray, 1835 deserve further evaluation. At least the latter genus shows affinity rather to Acanthoxylini (Phasmatinae) than Pachymorphinae (→ 4.3.3). In contrast to Pachymorpha, the eggs of Micrarchus hystriculeus (Westwood, 1859), the type species of Micrarchus Carl, 1913, are laterally compressed, strongly sculptured, a conical operculum and appear to have a closed internal micropylar plate (see Clark-Sellick, 1998: 219, figs. 28d & e), another indication for the polyphyly of Pachymorphini. The intergeneric relationships and systematic position of Hemipachymorphini are very obscure and barely deserve more detailed research. As presently treated the tribe is certainly polyphyletic. Studies clarifying the relationships and systematic positions of these two tribes are believed to be currently on-going. Tribe Gratidiini Bragg, 1995 The tribe Gratidiini Bragg, 1995c (= Ramulini sensu Günther, 1953) was critically reviewed and re-diagnosed by Cliquennois (2004) who has shown it to be highly polyphyletic in its present recognition. Tilgner (2002) and Whiting et al. (2003) supposed parts of present Gratidiini were likely to be related to Clitumnini. Indeed three Oriental genera (Parapachymorpha Brunner v. Wattenwy, 1893, Gongylopus Brunner v. Wattenwyl, 1907 and Cnipsomorpha Hennemann, Conle, Zhang & Liu, 2007) are here removed from Gratidiini and transferred to Clitumninae: Clitumnini (→ 5.2). Based on sperm ultrastructure, Baccetti (1987) concluded the genera Clonaria Stål, 1875 (Pachymorphinae : Gratidiini), Ramulus Saussure, 1862 (Clitumninae : Clitumnini) and Carausius Stål, 1875 (Lonchodinae) were closely related, which supports the assumption of Tilgner (2002) and Whiting et al. (2003). Indeed, the very slender and stick-like body, very short median segment, unarmed legs, short antennae with a laterally dilated sapus and egg-structures, such as the increasingly elongated capsule and peripheral outer rim of the operculum in certain genera or the open internal micropylar plate with a median line (Figs. 57a & b) support close relationship between Gratidiini and Clitumnini. Moreover, the profemora are of a rather similar structure, being distinctly triangular with the two dorsal carinae nearing and the anterior carina raised, and the medioventral carina distinct but at best slightly displaced towards the anteroventral carina. The %% genital morphology of Gratidiini however differs considerably from members of Clitumnini. All representatives have a distinct, well developed and sclerotised external vomer and a simple anal segment which consists of a single dorsal plate. Often the anal segment has the posterior margin excavated or with one or two straight finger-like appendices. The cerci are more or less strongly elongated and often in-curving and specialized, being broadened or forked at the apex. In correation to the external vomer in %%, && usually have a ± distinct praeopercular organ on abdominal sternum VII. In contrast to Clitumnini several members of Gratidiini glue their grass-seed like eggs to a surface. Nevertheless, relationship with Clitumnini is likely but any confirmed decision on the systematic position of Gratidiini s. str. certainly deserves more comprehensive and



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detailed investigation of taxa and comparison with members of Clitumnini. The centre of distribution of Gratidiini sensu stricto is Africa, with the highest diversity of taxa in eastern Africa. Representatives have dispersed as far north as the Mediterranean and as far east as the northern portions of the Oriental Region (Cliquennois, 2004: 13). Genera of Gratidiini (sensu stricto): 1. Adelungella Brunner v. Wattenwyl, 1907: 229. Type species: Adelungella insignis Brunner v. Wattenwyl, 1907: 22, by monotypy. 2. Burria Brunner v. Wattenwyl, 1900: 37. Type species: Burria longixipha Brunner v. Wattenwyl, 1907: 38, by subsequent designation of Kirby, 1904a: 336. 3. Camax Karsch, 1898: 365, 380. Type species: Camax acutus Karsch, 1898: 380, by monotypy. [= Leptynia] 4. Clonaria Stål, 1875: 5. Type species: Bacillus natalis Westwood, 1859: 6, by subsequent designation of Kirby, 1904a: 33. = Gratidia Stål, 1875: 14, 17. Type species: Gratidia sansibara Stål, 1875: 70, by original designation. [Synonymised by Rehn, 1933: 61] 5. Gharianus Werner, 1908: 714. Type species: Gharianus klaptoczi Werner, 1908: 714, by monotypy. [Transferred from Clitumnini by Cliquennois, 2004: 24] 5. Gratidiinilobus Brock, 2005: 31. Type species: Gratidiinilobus capensis Brock, 2005: 31, figs. 10–17, by original designation. 6. Ladakhomorpha Hennemann & Conle, 1999: 13. Type species: Ladakhomorpha longipes Hennemann & Conle, 1999: 14, by original designation. 7. Leptynia Pantel, 1890: 385, 398. Type species: Leptynia attenuata Pantel, 1890: 402, by subsequent designation of Kirby, 1904a: 335. = Leptyniella Bolívar, 1926: 101. Type species: Leptynia attenuata Pantel, 1890: 402, by original designation. [Objective synonym] 8. Linocerus Gray, 1835: 13, 19. Type species: Linocerus gracilis Gray, 1835: 20, by monotypy. [Removed from erroneous synonymy with Bacillus St. Fargeau & Audinet-Serville, 1827 by Hennemann, Conle & Zhang, 2008: 12] 9. Macellina Uvarov, 1940: 112. Type species: Bacillus souchongia Westwood, 1859: 12, by indication. [Replacement name for preoccupied Macella Stål, 1875] = Macella Stål, 1875: 13. Type species: Bacillus souchongia Westwood, 1859: 12, by subsequent designation of Rehn, 1904: 80. [Preoccupied by Macella Walker, 1858 (Lepidoptera)] 10. Maransis Karsch, 1898: 365, 381. Type species: Bacillus mozambicus Westwood, 1859: 179, by monotypy. 11. Paraclonaria Brunner v. Wattenwyl, 1893: 89. Type species: Paraclonaria hamuligera Schulthess, 1898: 83, by subsequent designation of Rehn, 1904: 83. 12. Paraleptinia Carl, 1913: 20. Type species: Paraleptinia schulthessi Carl, 1913: 21, by monotypy. 13. Phtoa Karsch, 1898: 365, 368. Type species: Phtoa prolixa Karsch, 1898: 368, by original designation. 14. Sceptrophasma Brock & Seow-Choen, 2000: 3. Type species: Bacillus hispidulus Wood-Mason, 1873: 47, by original designation. 15. Zehntneria Brunner v. Wattenwyl, 1907: 233. Type species: Zehntneria mystica Brunner v. Wattenwyl, 1907: 233, by subsequent designation of Otte & Brock, 2005: 344.


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FIGURES 40–43. Anal segments of %%, ventral view [scale = 1 mm] TIX = tergite IX; TX = tergite X (anal segment); Ce = cercus; Pa = paraproct; Ep = epiproct (supraanal plate); Vo = vomer; DoF = “Dornenfeld”. 40. Tribe Pharnaciini: Tirachoidea jianfenglingensis (Bi, 1994) [Semi–tergites spread to show the vomer (Vo)] 41. Tribe Clitumnini: Cuniculina cuniculus (Westwood, 1859) 42. Tribe Monandropterini: Monandroptera acanthomera (Burmeister, 1838) 43. Tribe Stephanacridini: Hermarchus pythonius (Westwood, 1859)



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FIGURES 44–47. Cross-section of profemur of && of Phasmatinae and Stephanacridini [scale = 0.5 mm] 44. Tribe Phasmatini: Acrophylla wuelfingi (Redtenbacher, 1908) 45. Tribe Monandropterini: Monandroptera acanthomera (Burmeister, 1838) 46. Tribe Pharnaciini: Tirachoidea jianfenglingensis (Bi, 1994) 47. Tribe Clitumnini: Cuniculina cuniculus (Westwood, 1859)

4.6.2 Subfamily Xeroderinae Günther, 1953 Bradler (2001: 183) stated the entire subfamily Xeroderinae Günther, 1953 to belong in Lanceocercata. Currently nine distinct genera are attributed to Xeroderinae (Otte & Brock, 2005: 33, → see list below), which are distributed throughout the Australian Region and Pacific Ocean as far east as Fiji and have one genus each in Mauritius (Epicharmus Stål, 1875) and continental Southeast Asia (Xenophasmina Uvarov, 1940). Detailed examination of all genera with particular emphasis on the external morphology of the genitalia and egg features has revealed reasonable evidence that the subfamily as presently treated is polyphyletic. Indeed, certain taxa match with the characteristic features given for Lanceocercata by Bradler but others apparently do not. Although all genera included have laterally flattened cerci, these are only considerably enlarged, foliaceous or lanceolate in the type genus Xeroderus Gray, 1835, Xenophasmina Uvarov, 1940, and certain species of Nisyrus Stål, 1875 and Cotylosoma Wood-Mason, 1878. All other genera have them very small and shorter than the anal segment. The external morphology of the %% genitalia reveals at least two distinct generic units and apparently supposes a polyphyly of present Xeroderinae. The typically tectiform and longitudinally split anal segment which consists of two interiorly dentate semi-tergites is only true for the type genus Xeroderus and the Mauritian Epicharmus, the structure of the forceps being very similar to that of other Lanceocercata (for Epicharmus see: Cliquennois & Brock, 2004: 12, fig. 36). These two genera and Dimorphodes Westwood, 1859 all lack an external vomer, but Dimorphodes has the anal segment shaped rather unlike that of typical Lanceocercata. It is just very slightly tectiform and the posterior margin bears a few small teeth ventrally, but the posterolateral angles are not expanded, nor do they form any kind of forcep-like structure. Therefore, close relation to these two genera and a placement of Dimorphodes in Xeroderinae is questionable. In contrast to these three genera %% of Nisyrus, Cotylosoma and Xenophasmina (confirmed upon material at hand from the author’s collections) have a simple anal segment and a well developed and sclerotised, hook-like external vomer. Although adult insects are very similar to Xeroderus in most aspects, a close relation with Xeroderus and placement in Xeroderinae deserves further evaluation. Furthermore, two fundamentally distinct types of internal plates in the eggs of present Xeroderinae (closed in Cotylosoma and Nisyrus vs. open in Dimorphodes) indicate this subfamily is either polyphyletic or deserves splitting into two


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separate tribes (confirmed upon eggs at hand from the author’s collections). The %% of Caledoniophasma are not known, so the systematic position of this New Caledonian genus remains as yet questionable. Otte & Brock (2005: 76) have privisionally transferred it to Xeroderinae. Unfortunately, the unique & holotype of the type species (the only representative known) is rather incomplete and lacks most legs and both antennae, but the very bulgy and cup-like subgenital plate, leg armature and shape of the head indicate relationship to Dimorphodes. Zompro (2001b: 19) removed the small Caledonian genus Cnipsus Redtenbacher, 1908 from Eurycanthinae and provisionally transferred it to Xeroderinae. Its exact position however remains questionable and warrants detailed examination of the genitalia of both sexes, the antennae and unknown eggs in particular. The disjunct distribution of Xenophasmina, the only representative of Xeroderinae known from Vietnam, Myanmar and Thailand, is remarkable and warrants clarification. The presence of a member of Xeroderinae on Mauritius, namely Epicharmus, was discussed by Cliquennois & Brock (2004). Certainly more detailed investigation of all taxa currently attributed to Xeroderinae is required to decide about the position of the genera marked with an asterix (*) in the list below. Genera included: 1. Caledoniophasma Zompro, 2001: 29. Type species: Caledoniophasma marshallae Zompro, 2001: 30, figs. 1–2, by original designation.* 2. Cnipsus Redtenbacher, 1908: 350. Type species: Acanthoderus rachis Saussure, 1868: 64, by subsequent designation of Zompro, 2001b: 19.* 3. Cotylosoma Wood-Mason, 1878: 102. Type species: Cotylosoma dipneusticum Wood-Mason, 1878: 102, by monotypy.* 4. Dimorphodes Westwood, 1859: 80. Type species: Dimorphodes prostasis Westwood, 1859: 81, pl. 34: 4–5, by monotypy.* 5. Epicharmus Stål, 1875: 60. Type species: Prisopus marchali Audinet-Serville, 1838: 284, by subsequent designation of Kirby, 1904a : 404. 6. Leosthenes Stål, 1875 : 60, 102. Type species: Leosthenes aquatilis Stål, 1875: 102, by monotypy. = Cacomorpha Sharp, 1898: 91. Type species: Cacomorpha aberrans Sharp, 1898: 91, pl. 8: 12 (= Leosthenes aquatilis Stål, 1875), by monotypy. [Synonymised by Redtenbacher, 1908: 438]* 7. Nisyrus Stål, 1877: 66. Type species: Nisyrus spinulosus Stål, 1877: 66, by subsequent designation of Kirby, 1904a: 407.* 8. Xenophasmina Uvarov, 1940: 379. Type species: Xenophasma fimbriatum Redtenbacher, 1908: 442, pl. 21: 3, by indication. [Replacement name for preoccupied Xenophasma Redtenbacher, 1908]* = Xenophasma Redtenbacher, 1908 : 442. Type species: Xenophasma fimbriatum Redtenbacher, 1908: 442, pl. 21: 3, by Zompro, 2004: 323. [Preoccupied by Xenophasma Dognin, 1905 (Lepidoptera: Pyralidae)] 9. Xeroderus Gray, 1835: 32. Type species: Xeroderus kirbii Gray, 1835: 32, by monotypy. = Cooktownia Sjöstedt, 1918: 43. Type species: Cooktownia plana Sjöstedt, 1918: 44, pl. 7: 2–3 (= Xeroderus kirbii Gray, 1835), by monotypy. [Synonymised by Brock & Hasenpusch, 2007: 75]

4.7 Tribes excluded from the family Phasmatidae s. str. The tribes Achriopterini Günther, 1953 and Stephanacridini Günther, 1953 from the Papuan Subregion were both placed in Phasminae (= Phasmatinae Bradley & Galil, 1977) by Günther (1953: 555). They however differ fundamentally from Phasmatinae and the entire Phasmatidae s. str. (Phasmatinae, Tropidoderinae, Extatosomatinae, Xeroderinae, Pachymorphinae and “Platycraniinae”) in genital morphology and are here removed



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from that family. In both, Achriopterini and Stephanacridini, both sexes have small, cylindrical to sub-cylindrical cerci, %% have a simple anal segment and a well produced, sclerotised external vomer, and && possess a more or less distinct praeopercular organ. Consequently, as these features strongly violate the diagnostic characters of Phasmatidae s. str., neither tribe belongs in the concerned family and may provisionally be interpreted as subordinate taxa of Phasmatidae sensu lato. In addition to the homologous genital morphology, several features of the insects and eggs indicate a close relationship between Achriopterini and Stephanacridini (Hennemann & Conle, 2004: 48). Both tribes are characteristic for having the abdomen longer than the head and thorax combined, the subgenital plate of && considerably elongated often lanceolate or spatulate, and all tibiae triangular in cross-section with the dorsal carinae strongly approaching each other. The eggs of both tribes have closed internal micropylar plates with the micropylar cup placed more or less in the centre of the plate. Externally the plate is rather small, at best covers half of the capsule length, and lenticular or rhomboidal in shape. Moreover, certain representatives of Stephanacridini (e.g. members of the type genus Stephanacris Redtenbacher, 1908) are remarkably similar in general appearance to Achriopterini, having rather short, prominently armed legs, head and mesothorax and half-sized (&&) or fully developed alae (%%). The presence of a close relative of Stephanacridini in Madagascar and the Comoros may be explained by a fairly early settlement, perhaps during the middle or late Jurassic, via Australia or Antarctica (Hennemann & Conle, 2004: 52). In several respects the fauna of Madagascar exhibits close affinity with the Austro-Malayan, Papuan and Polynesian faunas.

4.7.1 Achriopterini Günther, 1953 A detailed revision of Achriopterini on the species level and a new diagnosis of the tribe were presented by Hennemann & Conle (2004). Achriopterini differs from the closely related Stephanacridini by: the lack of a medioventral carina on the femora; fused dorsal carinae of the meso- and metatibiae; short gonapophyses and distinct praeopercular organ of &&; and two terminal points of the external vomer of %%. The eggs clearly differ from those of Stephanacridini by the prominent extension of the polar-area and peripheral extension on the outer margin of the operculum (Hennemann & Conle, 2004: 50). Distribution: Madagascar and Republic of Comoros, endemic. Genera included: 1. Achrioptera Coquerel, 1861: 495. Type species: Achrioptera fallax Coquerel, 1861: 495, pl. 9: 1, by monotypy. = Enetia Kirby, 1891: 151. Type species: Enetia spinosissima Kirby, 1891: 151, by monotypy. [Synonymised by Redtenbacher, 1908: 439] = Hovaspectrum Rehn, 1940: 1. Type species: Hovaspectrum lobipes Rehn, 1940: 3, by original designation. [Synonymised by Hennemann & Conle, 2004: 8] 2. Glawiana Hennemann & Conle, 2004: 41. Type species: Glawiana glawi Hennemann & Conle, 2004: 42, figs. 16, 34–35, 71, 73 & 92, by original designation.

4.7.2 Stephanacridini Günther, 1953 (Figs. 37, 43, 60–61, 80–82) Type genus: Stephanacris Redtenbacher, 1908: 441.


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Stephanacridini Günther, 1953: 555. Bradley & Galil, 1977: 194. Hennemann & Conle, 2004: 48. Otte & Brock, 2005: 33.

The tribes Pharnaciini Günther, 1953 and Stephanacridini Günther, 1953 were both included in the subfamily Phasminae (= Phasmatinae Bradley & Galil, 1977) by Günther (1953: 555). Striking differences concerning to the genitalia and eggs however prove that these two tribes are not closely related and that Stephanacridini is not a member of Phasmatidae s. str. (→ 4.1) nor the subfamily Clitumninae (→ 5.). Several genera included in Pharnaciini by Günther (1953) and Bradley & Galil (1977) fundamentally violate the diagnostic features of the tribe, e. g. by the rather indistinct medioventral carina of the profemora which is positioned midways on the ventral surface of the profemur, simple anal segment and well produced, sclerotised external vomer of %% (Figs. 37, 43), more or less distinct praeopercular organ and elongate, filiform gonapophyses of && which considerably exceed the apex of the abdomen, and closed internal micropylar plate of the eggs (Figs. 60–61). Pharnaciini in contrast, which include the type genus Pharnacia Stål, 1877, have the medioventral carina of the profemora prominently elevated, ledge-like and strongly displaced towards the anteroventral carina (Fig. 46), the anal segment strongly tectiform, more or less deeply split and consisting of two movable, interiorly dentate semi-tergites, which function as a clasping apparatus to grasp the & sternum VII during copulation (Figs. 35, 40), the external vomer of %% lacking and the eggs with an open internal micropylar plate and a median line (Fig. 54, → 6.). This obviously shows Pharnaciini as presently treated is polyphyletic and further research has proven the taxa with a well developed external vomer of %% and closed internal micropylar plate of the eggs formerly placed in Pharnaciini to be closely related to Stephanacris, hence to belong in Stephanacridini. Consequently, Hennemann & Conle (2006) have transferred Hermarchus from Pharnaciini to Stephanacridini and described a new genus, Macrophasma, from New Guinea. Four further genera, namely Diagoras, Eucarcharus, Phasmotaenia and Sadyattes, are here removed from Pharnaciini and transferred to Stephanacridini (→ see list of genera below). Due to similar reasons the genus Nesiophasma Günther, 1934 is removed from Clitumnini (= Baculini sensu Bradley & Galil, 1977) and also included in Stephanacridini. The peculiar genus Gigantophasma Sharp, 1898 differs considerably from other Phasmatinae in several aspects and is here transferred to the tribe Gigantophasmatini trib. nov. of Tropidoderinae (→ 4.4.3). A more detailed comparison and distinction of the tribes Pharnaciini and Stephanacridini is provided in table 1 below. Clark-Sellick (1998) studied the internal micropylar plate of the eggs of Phasmatodea and its relevance for the classification of the order. Having examined the eggs of different taxa included in Pharnaciini by Günther (1953) and subsequent authors, Clark-Sellick (1998: 226) stated that the plate and capsule morphology within this tribe show so much variation that it can be considered polyphyletic. The author observed two distinct types of eggs which are clearly distinguished by the structure of the internal micropylar plate. This is either open (Clark-Sellick mentioned the genera Baculolonga, Phryganistria and Tirachoidea), or closed (Hermarchus, Phasmotaenia and Gigantophasma). Clark-Sellick (1998: 226) commented: “The eggs of this tribe fall into two groups. These appear to be associated with two quite distinct types of plates. Group (I) with open internal plates […]. Group (II) with closed internal plates […].”. As stated above comparison of all genera attributed to Pharnaciini by Günther (1953) and subsequent authors has shown the type of internal micropylar plate is in accordance to the shape and structure of the %% anal segment and thus clearly confirms Clark-Sellick’s (1998) remarks on the polyphyletic nature of Pharnaciini. Taxa with a tectiform anal segment, that consists of two separate interiorly dentate semi-tergites always have open plates, whereas it is closed in taxa with a simple anal segment and a well developed vomer. Clark-Sellick (1998) did not have eggs of Stephanacris Redtenbacher, the type genus of Stephanacridini, at hand for examination and hence was not able to detect the close relationship between his “group II” (genera Hermarchus, Phasmotaenia and Gigantophasma) and this tribe.



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Günther (1953: 555) established Stephanacridini solely for the New Guinean genus Stephanacris Redtenbacher, 1908. Based on Günther’s key features, Bradley & Galil, (1977: 192) characterized Stephanacridini as follows: “Operculum [= subgenital plate] of female extended broadly over apex of abdomen” and “Head swollen; middle and hind femora with tooth-like, serrate inferior carinae. New Guinea”. Apparently, since Stephanacridini now inludes other genera than the type genus Stephanacris, these diagnostic features of former authors do no longer hold to characterize the tribe. A complete revision of Stephanacridini on the species level and a new diagnosis of the tribe by Hennemann & Conle is in preparation. Structures of the genitalia, such as the well developed vomer, cylindrical cerci and simple anal segment of %%, lanceolate subgenital plate and long, filiform gonapophyses of &&, and closed internal micropylar plate of the eggs suppose relation to the genus Platycrana Gray, 1835 (subfamily “Platycraninae”). This is in particular emphasized by && of Macrophasma, which in addition to the features mentioned has the head, antennae, mesothorax, abdominal tergites and legs very similar in structure to Platycrana, with some species appearing like apterous Platycrana. A supposed relationship between Platycrana and the tribe Stephanacridini however deserves further evaluation and will be part of a revision of Stephanacridini by the authors (in preperation). Distribution (Fig. 82): Philippines, Taiwan, West Malaysia, Indonesia, New Guinea, Salomon Islands, NE-Australia, Micronesia and Melanesia (Fiji, Tonga, Samoa, Vanuatu, New Caledonia & Western French Polynesia). Genera included: 1. Diagoras Stål, 1877b: 66. Type species: Diagoras ephialtes Stål, 1877b: 66, by monotypy. = Eustygera Brunner v. Wattenwyl, 1907: 186. Type species Eustygera godeffroyi Brunner v. Wattenwyl, 1907: 186, pl. 7: 2, by monotypy. [Synonymised by Günther, 1932b: 756] 2. Eucarcharus Brunner v. Wattenwyl, 1907: 185. Type species: Lonchodes feruloides Westwood, 1859: 45, pl. 6: 5a–b, by subsequent designation of Günther, 1935b: 138. 3. Hermarchus Stål, 1875: 45. Type species: Phibalosoma pythonius Westwood, 1859: 73, pl. 12: 1 & 35: 3, by subsequent designation of Kirby, 1904: 361. 4. Macrophasma Hennemann & Conle, 2006b: 3. Type species: Hermarchus biroi Redtenbacher, 1908: 445, by original designation. 5. Nesiophasma Günther, 1934c: 5. Type species: Nesiophasma eremothocus Günther, 1934c: 6, fig. 1, by original designation. = Mylothrus Günther, 1935c: 18. Type species: Mylothrus oligarches Günther, 1935c: 18, pl. 2: 12, by original designation. [Synonymised by Günther, 1935c: 29] 6. Phasmotaenia Návas, 1907: 10. Type species: Taeniosoma sanchezi Bolivar, 1897: 31, fig. 1, by indication. [replacement name for the preoccupied Taenionema Bolivar, 1906] FIGURES 48–59. Eggs and internal micropylar plates, dorsal view (open plates) [scale = 1 mm] 48. Tribe Phasmatini: Phasma gigas (Linné, 1758), a. egg, b. internal micropylar plate 49. Tribe Phasmatini: Acrophylla wuelfingi (Redtenbacher, 1908), a. egg, b. internal micropylar plate 50. Tribe Tropidoderini: Tropidoderus childrenii Gray, 1833, a. egg, b. internal micropylar plate 51. Tribe Tropidoderini: Podacanthus viridiroseus Stål, 1875, a. egg, b. internal micropylar plate 52. Tribe Monandropterini: Rhaphiderus scabrosus (Percheron, 1829), a. egg, b. internal micropylar plate 53. Tribe Monandropterini: Monandroptera acanthomera (Burmeister, 1838), a. egg, b. internal micropylar plate 54. Tribe Pharnaciini: Tirachoidea jianfenglingensis (Bi, 1994), a. egg, b. internal micropylar plate 55. Tribe Clitumnini: Cuniculina cuniculus (Westwood, 1859), a. egg, b. internal micropylar plate 56. Tribe Medaurini trib. nov.: Medauroidea extradentata (Brunner v. Wattenwyl, 1907), a. egg, b. internal micropylar plate 57. Tribe Gratidiini: Sceptrophasma hispidulum (Wood-Mason, 1873), a. egg, b. internal micropylar plate 58. Lonchodinae: Lonchodini: Lonchodes brevipes Gray, 1835, a. egg, b. internal micropylar plate 59. Lonchodinae: Neohiraseini trib. nov.: Neohirasea maerens (Brunner v. Wattenwyl, 1907), a. egg,



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b. internal micropylar plate

FIGURES 60–67. Eggs and internal micropylar plates, dorsal view (closed plates) [scale = 1 mm] 60. Tribe Stephanacridini: Nesiophasma plateni (Dohrn, 1910), a. egg, b. internal micropylar plate 61. Tribe Stephanacridini: Hermarchus pythonius (Westwood, 1859), a. egg, b. internal micropylar plate 62. Tribe Tropidoderini?: Didymuria violascens (Leach, 1814), a. egg, b. internal micropylar plate 63. Extatosomatinae: Extatosoma tiaratum tiaratum (Mac Leay, 1826), a. egg, b. internal micropylar plate 64. Tribe Acanthoxylini: Clitarchus hookeri (White, 1846), a. egg, b. internal micropylar plate 65. Tribe Acanthoxylini: Acanthoxyla geisovii (Kaup, 1871), a. egg, b. internal micropylar plate 66. Tribe Gigantophasmatini trib nov.: Gigantophasma bicolor Sharp, 1898, a. egg, b. internal micropylar plate 67. Subfamily “Platycraninae”: Megacrania tsudai Shiraki, 1933, a. egg, b. internal micropylar plate


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FIGURES 68–69. 68a. Tribe Tropidoderini?: Didymuria violescens (Leach, 1814), % [after Key, 1991, fig. 25.11A] 68b. Tribe Tropidoderini?: Didymuria violescens (Leach, 1814), & [after Key, 1991, fig. 25.11B] 69a. Tribe Gigantophasmatini trib nov.: Gigantophasma bicolor Sharp, 1898, & [after Sharp, 1898, pl. 7: 6] 69b. Tribe Gigantophasmatini trib nov.: Gigantophasma bicolor Sharp, 1898, % [after Sharp, 1898, pl. 7: 5]



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= Taeniosoma Bolivar, 1897: 31. Type species: Taeniosoma sanchezi Bolivar, 1897: 31, fig. 1, by monotypy. [Preoccupied by Taeniosoma Stimpson, 1857 (Nematoda)] = Taenionema Bolivar, 1906: 393. Type species: Taeniosoma sanchezi Bolivar, 1897: 31, fig. 1, by indication. [Replacement name for preoccupied Taeniosoma Bolivar, 1897—preoccupied by Taenionema Banks, 1905 (Plecoptera)] = Taeniophasma Uvarov, 1940: 379. Type species: Taeniosoma sanchezi Bolivar, 1897: 31, fig. 1, by indication. [Homonym of Phasmotaenia Návas, 1907 & unnecessary replacement name for Taenionema Bolivar, 1906] = Phasmotaenionema, Günther, 1933: 155. [Misspelling of Phasmotaenia Návas, 1907] = Phasmatotaenionema, Bradley & Galil, 1977: 193. [Misspelling of Phasmotaenia Návas, 1907] 7. Sadyattes Stål, 1875: 44, 88. Type species: Sadyattes borrii Stål, 1875: 88, by monotypy. 8. Stephanacris Redtenbacher, 1908: 441. Type species: Stephanacris brevipes Redtenbacher, 1908: 441, pl. 21: 4a–b, by subsequent designation of Bradley & Galil, 1977: 194. TABLE 1. Comparison and differentiation of Pharnaciini and Stephanacridini Pharnaciini

Stephanacridini

Body cross-section (&&)

Cylindrical

Sub-cylindrical to oval

Mesothorax (&&)

± cylindrical; never considerably elevated; Mesonotum and/or mesopleurae often ± eleunarmed (exception Phobaeticus foliatus vated and lamellate; often spinose Bragg)

Wings (&&)

Apterous

Apterous or brachypterous; if alae present then often colourful

Medioventral carina of Distinctly elevated, ledge- or lamella-like Indistinct and roughly midways on ventral profemora and strongly displaced towards anteroventral surface of femur; may be spinose carina; unarmed Gonapophyses (&&)

Not considerably projecting over apex of ± elongated and filiform; projecting anal segment over apex of anal segment by as much as combined length of tergites IX and X

Praeopercular organ (&&) ± distinct; formed by a scale-like hump, pair No external structure present; a fold or of ridges, tubercles, spines or lobes rounded impression Anal segment (%%)

Strongly tectiform and split; consisting of A single tergite; not distinctly tectiform two separate semi-tergites with a distinct (Figs. 37 & 43) “Dornenfeld” intero-apically (= clasping apparatus, Figs. 35 & 40)

External vomer (%%)

Rudimentary (Fig. 40)

Well developed and sclerotised; a triangular plate or hook-like organ (Fig. 43)

Internal micropylar plate Open with a median line of eggs (Fig. 54b)

Closed (Figs. 60b & 61b)

Capitulum of eggs

Not stalked; ± strongly appressed to operculum

Stalked

Geographical distribution Oriental Region (Fig. 82)

Australia, New Guinea, Melanesia, Micronesia, western Polynesia, Wallacea and southern portions of the Oriental Region (Fig. 82)


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FIGURES 70–72. 70. Tribe Tropidoderini: Tropidoderus childrenii Gray, 1833, & [after Gray, 1833, pl. 3: 1] 71. Tribe Tropidoderini: Podacanthus typhon Gray, 1833, & [after Gray 1833, pl. 2: 1] 72. Tribe Phasmatini: Peloriana lobiceps (Mac Leay, 1884), & [after Redtenbacher, 1908, pl. 22: 7b] REVISION OF PHARNACIINI


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5 Subfamily Clitumninae Brunner V. Wattenwyl, 1893 Clitumninae Brunner v. Wattenwyl, 1893 Type genus: Clitumnus Stål, 1875: 9. Clitumnidae Brunner v. Wattenwyl, 1893: 87. Bruner, 1915: 37. Ramulinae Kevan, 1982: 382.

Description: Medium sized to very large (body length 28.0–357.0 mm), stick-like, slender to extremely slender and often very elongate Phasmatodea (Figs. 76–78). Body cylindrical in cross-section and of ± uniform width. && apterous, %% may be brachypterous or alate. Ocelli lacking. Gula present. Antennae at best 3/5 the length of body, but mostly conspicuously shortened and often just a little longer than head and pronotum combined; with less than 35 segments. Mesothorax mostly very elongate; in a very few cases just a little longer than head and pronotum combined; rarely granulose or spinose. Median segment distinctly shorter (majority of taxa), equal in length, or slightly longer than the metanotum (a few alate %%). Abdominal tergite VII often with a lateral lobe or expansion (&& in particular). Sternum VII of && with a ± distinct praeopercular organ; formed by a posteromedian scale-like structure, hump, or pair of spines or lobes. Gonapophyses of && elongated in certain taxa, but never projecting considerably over apex of abdomen. Subgenital plate of && boatlike, distinctly longitudinally keeled and variable in length; ranging from small, scoop-shaped and not reaching the apex of anal segment, to strongly elongated and lanceolate. Anal segment of %% laterally compressed, slightly tectiform and consisting of two semi-tergites, which have the apex ± elongated and finger-like, slightly in-curving and the interior surface armed with numerous small teeth (Figs. 20–25, 29–31, 35–36, 40– 41). External vomer of %% lacking in a majority of taxa; a rather slender, papillate or filiform external vomer may however be present ina few more derived genera. Supraanal plate usually small and considerably shorter than anal segment; rarely elongated and lanceolate. Cerci small and usually shorter than anal segment, cylindrical to sub-cylindrical in cross-section; rarely laterally compressed or slightly lanceolate. Profemora distinctly triangular in cross-section with the two dorsal carinae ± distinctly approaching each other, compressed and curved basally. Posterodorsal carina considerably reduced, anterodorsal carina ± strongly raised and serrate. Medioventral carina very prominent, acutely ledge or lamella-like and strongly displaced towards anteroventral margin (Figs. 46–47). Mid- and hind legs often with prominent armature (increasingly reduced im more derived forms). Meso- and metafemora trapezoidal in cross-section with dorsal carinae nearing, medioventral carina present. Tibiae trapezoidal to almost triangular in cross-section, the dorsal carinae strongly nearing each another. No area apicalis on tibiae. Basitarsus distinctly longer than following tarsomere. Eggs: These may be of various shapes and structures, ranging from simply ovoid with a central capitulum to considerably compressed laterally, angulate and bark-like with variable structures on the outer margin of the operculum. A common feature is the open internal micropylar plate with a ± distinct median line. Comments: Since the tribes Clitumnini Brunner v. Wattenwyl, 1893 and Pharnaciini Günther, 1953 obviously do not belong in the Phasmatidae s. str. (→ 4.) and subfamily Phasmatinae respectively, they together form a separate subfamily of Phasmatidae sensu lato, the Clitumninae. Clitumnidae was first used for a tribe by Brunner v. Wattenwyl (1893: 87) for taxa now in Clitumnini and Pachymorphini (subfamily Pachymorphinae, → 4.6.1). Subsequently, Brunner v. Wattenwyl (1907: 181) re-named the tribe Clitumnini, and as this is an available name and the monophyletic group here proposed (Clitumnini + Pharnaciini) includes the type genus Clitumnus Stål, 1875, Clitumninae is the valid name for the subfamily that contains these two tribes. Representatives of Clitumninae were predominantly included in the subfamily Phasmatinae by former authors (e.g. Bradley & Galil, 1977). There are however several features of the genital exosceleton and mor-


HENNEMANN & CONLE

phology of the profemora, which readily distinguish the tribes Clitumnini and Pharnaciini from the subfamily Phasmatinae. These are the small and cylindrical to sub-cylindrical cerci, presence of a praeopercular organ on sternum VII of && and profemora, which have the medioventral carina very distinct, ledge-like or lamellate and conspicuously displaced towards the anteroventral carina. The anal semi-tergites of %% of Clitumninae are of a rather distinct shape and structure if compared to those of Phasmatinae and Phasmatidae s. str. respectively, having the apex ± strongly elongated, finger-like and in-curving with numerous small teeth on their interior surface (= “Dornenfeld”). Moreover, %% of certain derived taxa possess a well sclerotised, papillate, hook-like or filiform external vomer, never present in Phasmatidae s. str. and Phasmatinae. Differences are also observed in the copulation position of Clitumninae which is less derived than that seen in representatives of Phasmatidae s. str. (= Lanceocercata). %% either grasp the && abdomen on the corresponding anchorage on sternum VII, the praeopercular organ, or in the case an external vomer is present this is inserted into a slit or hole of that organ to hold fast to the & during copulation. The shape and length of the %%’s semi-tergites in many cases shows correlation to the shape of the appendages of the praeopercular organ of &&. In taxa whose && have a prominent praeopercular organ that is formed by a pair of lobes or appendages close to the posterior margin of sternum VII, %% have usually have the semi-tergites conspicuously elongated and finger-like. During the copulation these appendages of the && praeopercular organ are grasped and used as an anchorage by the %%, and mostly the shape of the semi-tergites fits very well with the shape of the appendages of the praeopercular organ. %% of both tribes contained have the external vomer more or less strongly reduced or lacking, mostly due to being functionally replaced by the movable semi-tergites of the anal segment as an anchorage to hold fast to the & abdomen during copulation. However, in certain taxa of Clitumnini the external vomer is developed and seen to be a sclerotised tube-like, papillate or slender, hook-like organ. This appears to have been re-evolved secondarily and functions as an additional anchorage, with the corresponding %% seen to grasp the praeopercular organ of && and insering their vomer into a posterior slit or hole of the same. In certain taxa of Pharnaciini which at first glance appear to lack an external vomer, it can seen to be an elongate and straight tube-like or papillate organ if the semi-tergites are spread and the paraprocts pushed aside (Fig. 40). Phasmatini, Acanthomimini, Acanthoxylini (all subfamily Phasmatinae, → 4.3), Pharnaciini and Clitumnini all have profemora that are distinctly triangular in cross-section, having the dorsal carinae strongly approching each other, the posterodorsal carina reduced and the anterodorsal carina more or less strongly raised and ± distinctly serrate. In Tropidoderini and Monandropterini, they are just indistinctly triangular in cross-section, with the dorsal carinae only moderately approaching each other and unarmed. Triangular profemora are however also found elsewhere throughout the Phasmatodea and thus this character cannot be used at a higher systematic rank than at subfamily level. This has led to much confusion since Redtenbacher (1908: 436) mentioned this as the main character to distinguished his Acrophyllini from other “Anareolatae”, which however is clearly an artificial group (→ 3.1). No subsequent author has since recognized the most obvious and characteristic feature of the profemora which serves for distinguishing the subfamilies Phasmatinae and Clitumninae, the structure and position of the medioventral carina. This is conspicuously raised, ledge-like or lamellate, ± in-curving and strongly displaced towards the anteroventral carina in Pharnaciini and Clitumnini (subfamily Clitumninae, Figs. 46–47), but merely a rather indistinct longitudinal keel midways on the ventral surface of the profemur in Phasmatinae (family Phasmatidae s. str., Fig. 44). As stated above the eggs of Clitumninae are very polymorphic but characteristic for having an open internal micropylar plate and a ± distinct median line (Figs. 54–56). This means, the micropylar cup is placed outside the borders of the plate, usually in a more or less distinct posterior incision or gap. The external plate is usually longer than wide, but may be transverse and does not reach the anterior and posterior poles of the egg capsule. It shows a more or less prominent posteromedian incision or gap similar in shape to that of the internal plate. The external plate is either oval, lanceolate, strongly elongate and parallel-sided, bilobed with an anteromedian indention, or even shaped like an inverted “Y”. The latter shape is atttained by elongation of the



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posterolateral regions on both sides of the median gap. While the micropylar plate usually covers more than ? of the capsule length in Pharnaciini it may be as short as just ¼ of the capsule length in certain Clitumnini. The operculum always bears more or less prominent opercular structures, such as a ± distinctly stalked capitulum (Pharnaciini, Fig. 54), or a raised crown-like outer margin, hairy structures or pseudocapitulum (Clitumnini, Figs. 55–56). The capsule is mostly ovoid and longer than wide, more rarely spherical in Pharnaciini and certain Clitumnini. In many Clitumnini however it is very elongate and strongly laterally flattened with the capsule surface unevenly sculptured, giving the egg a bark or grass-seed like appearance. Some genera of Pharnaciini have laterally flattened, lens-shaped eggs, which exhibit a distinct, more or less elevated dorsoventral keel. In general, egg features are observed to be very uniform and rather typical within each individual genus and therefore very well serve for the distinction of genera amongst Clitumninae. Close relationship between Clitumnini and Pharnaciini not only becomes obvious by the morphological features of the external genitalia and profemora here mentioned but is also emphasized by habitual similarity between taxa of both tribes. The more derived apterous representatives of Pharnaciini in particular strongly resemble certain genera of Clitumnini, the most remarkable example being represented by Baculonistria gen. nov. (Pharnaciini) and Entoria Stål, 1875 (Clitumnini). These two genera are merely distinguished by the key features which distinguish Pharnaciini from Clitumnini, namely the shape of the scapus, leg armature and opercular structures of the eggs. In Baculonistria gen. nov. the scapus is slender, all ventral carinae of the mid and hind legs distinctly serrate and the ovoid eggs bear a prominent, stalked central capitulum, whereas in Entoria the scapus is considerably dilated laterally, the mid and hind legs at best armed with single teeth and lobes or a with few minute basal serrations on the tibiae, and the eggs are laterally compressed, considerably longer than wide and lack a stalked central capitulum. Another example is represented by the very slender && of certain species of Phobaeticus Brunner v. Wattenwyl, 1907 with a short subgenital plate, which remarkably resemble && of the genera Ramulus Saussure, 1862 or Cuniculina Brunner v. Wattenwyl, 1907. Pharnaciini certainly inherit the more basal taxa amongst Clitumninae which is in particular seen in the relatively longer median segment of most genera and presence of tegmina and ± well developed alae in the %% of certain species. Interpreting Pharnaciini as the more basal forms of Clitumninae obviously reveals a loss of wings during the course of evolution of this particular subfamily, and suggests a secondary recovery of the external vomer, since this is only present in certain Clitumnini and the supposedly most derived genus of Pharnaciini (Baculonistria gen. nov.), but lacking in the more basal Pharnaciini. Evolutional trends recognized within Clitumninae are e.g. a decreasing body size, leg armature and sexual dimorphism, but an increasing elongation of the body segments and legs. Distribution & Biogeography (Fig. 1): Complete Oriental Region, southern portion of the Palearctic Region (China, Korea, Far East Russia & Japan), complete Wallacea and as far east as New Guinea (→ Map 1). The wide dispersal of Clitumninae in the Indo-Malayan Archipelago appears to have been mainly advanced by the glaciation during the Pleistocene which has caused a decrease of the world-wide sea-level and developed land or island chain bridges between the Asian continent and the Large Sunda Islands. These quite certainly allowed an interchange of the faunas of the Asian continent and Sundaland (Cox & Moore, 1993). Subsequent diversification and specialization, especially on the Asian continent, becomes obvious in the very speciose and derived Clitumnini. Tribes included: 1. Pharnaciini Günther, 1953: 555. Type genus: Pharnacia Stål, 1877a: 40. 2. Clitumnini Brunner v. Wattenwyl, 1907: 181. Type genus: Clitumnus Stål, 1875: 9 (= Ramulus Saussure, 1862: 471). 3. Medaurini trib. nov. Type genus: Medaura Stål, 1875 : 69.


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FIGURES 73–74. 73a. Tribe Phasmatini: Acrophylla enceladus (Gray, 1833), & [after Westwood, 1859, plate 39: 2] 73b. Tribe Phasmatini: Acrophylla enceladus (Gray, 1833), % [after Westwood, 1859, plate 39: 1] 74a. Tribe Monandropterini: Monandroptera acanthomera (Burmeister, 1838), & [after Redtenbacher, 1908, plate 18: 3b] 74b. Tribe Monandropterini: Monandroptera acanthomera (Burmeister, 1838), % [after Redtenbacher, 1908, plate 18: 3a]



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5.1 Pharnaciini Günther, 1953 (Figs. 20–22, 35, 40 46, 54, 76–77, 82) Type genus: Pharnacia Stål, 1877: 40. Pharnaciini Günther, 1953: 555 (in part). → for a complete list see chapter 6.

Pharnaciini are distributed over great parts of the Oriental region, including complete Sundaland and as far west as western India with the most northern dispersal being Bhutan and northern Central China (→ Fig. 1). The tribe includes some of the world’s longest Phasmatodea and exhibits less primitive characters than the more derived Clitumnini. The mesothorax is usually smooth and destitute of spines (exception Phobaeticus foliatus (Bragg)). The insects exhibit strong sexual dimorphism with %% much smaller and more slender than &&. && are apterous, %% either fully winged, brachypterous or apterous as well. The median segment is of variable length; it is equal in length or even slightly longer than the metanotum in taxa with winged %%, and shorter but at least ¼ the length of the metanotum in taxa with apterous %%. Ocelli are absent. The seventh abdominal tergite is more or less distinctly dilated posterolaterally (&& in particular). && of most taxa exhibit a prominent praeopercular organ on sternum VII, often formed by a pair of spine-like appendages or lobes. The semi-tergites of the %% anal segment are either short and roughly triangular in lateral aspect, or more or less prominently elevated apically, being ± elongate, slender and finger-like. The interior surfaces are in the apical portion either densely armed with minute teeth or bear a longitudinal, dentate bulge (= “ Dornenfeld”, Figs. 35 & 40). The cerci are small, usually cylindrical or oval in cross-section but my be laterally compressed and sickle-shaped in certain species. The external vomer of %% is strongly reduced and hidden underneath sternum XI (= paraproct). Sometimes it is seen to be an elongate, straight, tube-like or papillate organ (Fig. 40), or in Baculonistria gen. nov. is represented as a conspicuous “Y”-shaped sclerotised structure. The antennae of %% are slightly longer than those of the corresponding &&, but at best slightly longer than the head and complete thorax combined; with > 25 segments. The scapus is simple and slender. All antennomeres are elongate and distinctly longer than wide (exception Baculonistria gen. nov.). Most carinae of the legs are more or less prominently spinose, serrate or armed with foliaceous lobes. The medioventral carina of the meso- and metafemora is spinose. Eggs of Pharnaciini are similar to those of Phasmatinae: Phasmatini or Tropidoderinae: Tropidoderini (Phasmatidae s. str.), mostly being of an ovoid or spherical general shape (Fig. 54a). In some cases the capsule is laterally compressed, lens-shaped and surrounded by a more or less prominent dorsoventral keel which may be considerably elevated. The micropylar plate is of a variable shape, either being transverse and kidneyshaped (Phobaeticus), lanceolate and pointed towards the anterior (Phryganistria), or shaped like an inverted “Y” (Pharnacia and Tirachoidea). Internally the plate is open and has a median line (Fig. 54b). The operculum always bears a distinct hat or knob-like, stalked capitulum. The broader taxa of Pharnaciini with pterous %% show remarkable habitual similarities to certain genera of taxa of Phasmatinae: Phasmatini (Phasmatidae s. str.), and some && may at first glance appear like wingless forms of Phasmatini. Examples include Tirachoidea and Peloriana Uvarov, 1940 or Phobaeticus and Ctenomorpha Gray, 1833. A complete revision of Pharnaciini at the species level and a detailed new diagnosis is provided below (→ 6.). Genera included: 1. Baculonistria gen. nov. 2. Pharnacia Stål, 1877a: 40. 3. Phobaeticus Brunner v. Wattenwyl, 1907: 183.


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= Baculolonga Hennemann & Conle, 1997a: 347. = Lobophasma Günther, 1935: 138. syn. nov. = Nearchus Redtenbacher, 1908: 448. syn. nov. 4. Phryganistria Stål, 1875: 6 & 63. 5. Tirachoidea Brunner v. Wattenwyl, 1893: 83. stat. rev.

5.2 Clitumnini Brunner v. Wattenwyl, 1893 (Figs. 11–13, 23–25, 36, 41, 47, 55, 78, 82) Type genus: Clitumnus Stål, 1875: 9. Clitumnini Brunner v. Wattenwyl, 1907: 181. Cliquennois, 2004: 14. Baculini Günther, 1953: 555. Bradley & Galil, 1977: 192. Ramulini, Otte & Brock, 2005: 16. [Not: Baculini sensu Zompro, 2005: 134. → An exceptionally neotropical tribe of the subfamily Cladomorphinae]

Description: Small to large (body length 45.0–211.0 mm), mostly very slender and stick-like insects with very long and slender legs (%% in particular, Figs. 78a & b). The body is elongate and of ± uniform width, the abdominal segments II–VII at least 2x longer than wide. The sexual dimorphism is distinct, %% being considerably more slender and shorter than corresponding &&. The antennae are often strongly shortened and in some cases (&& in particular) just a little longer than the head and pronotum combined; reaching to the posterior of the mesonotum in certain %%. They consist of < 28 segments. The scapus is ± prominently enlarged and laterally dilated (&& in particular). The head is smooth (most %%) or armed with two conspicuous spines, horns or lobes between the eyes. The body is mostly unarmed and smooth but in a few cases finely granulose (&& in particular). The mesothorax is ± elongate and at least 2.5x longer than the head and pronotum combined. The median segment is strongly shortened and at best ¼ the length of the metanotum. The abdominal segments II–IV are increasing in length, the longest segments (V–VII) at least 2.5x longer than wide. Abdominal tergite VII slender. && have a more or less distinct praeopercular organ on sternum VII, usually formed by one or two spines or lobes. %% have the semi-tergites of the anal segment more or less prominently elevated apically, usually being slender and ± finger-like (Figs. 23–25). Interiorly they are set with minute teeth either all over or along the outer margins (“Dornenfeld”, Figs. 36 & 41). In both sexes the cerci are very small to slightly lanceolate, at best equal in length to the anal segment, and cylindrical or oval in cross-section. An external vomer lacking and the lower margins of the anal segment strongly approach each other basally and almost form a closed tube (Fig. 41). && of most taxa have a small and scoop-like to strongly convex and boatshaped subgenital plate which at best slightly extends over the anal segment. In certain genera however it is slightly elongated and pointed apically (e.g. Prosentoria), and others may in addition have an elongated supraanal plate to form a bird beak-like ovipositor (e.g. Entoria and Rhamphophasma). The armature of the extremities is increasingly reduced and in %% restricted to minute sub-apical spines of the femora and minute teeth in the apical portion of the tibiae. Conspicuous armature may be present on the mid and hind legs of && and represented by single enlarged teeth or lobes, but neither carina is densely serrate as in Pharnaciini. The medioventral carina of the meso- and metafemora is indistinct and unarmed. The profemora are very similar in structure to those of Pharnaciini having the two dorsal carinae conspicuously approaching each other, the anterodorsal carina considerably raised and the posterior carina reduced, usually being ± decidedly displaced towards the lateral surface of the femur. The medioventral carina is ± prominently elevated, ledge or lamellalike, curved inwards, and conspicuously displaced towards the anteroventral carina (Fig. 47). The anterodorsal carina is serrate in certain &&. REVISION OF PHARNACIINI


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FIGURES 75–78. 75. Tribe Phasmatini: Ctenomorpha marginipennis (Gray, 1833), & [after Redtenbacher, 1908, pl. 22: 6] 76. Tribe Pharnaciini: Phobaeticus serratipes (Gray, 1835), & [after Beier, 1957, fig. 11c] 77. Tribe Pharnaciini: Phobaeticus hypharpax (Westwood, 1859), % [after Westwood, 1859, pl. 13: 6] 78a. Tribe Clitumnini: Cuniculina cuniculus (Westwood, 1859), & [after Westwood, 1859, pl. 6: 1] 78b. Tribe Clitumnini: Ramulus stilpnus (Westwood, 1859), % [after Westwood, 1859, pl. 5: 7]


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FIGURES 79–81. 79. Tribe Acanthomimini: Acanthomima rhipheus (Westwood, 1859), & [after Westwood, 1895, pl. 8: 10] 80a. Tribe Stephanacridini: Hermarchus pythonius (Westwood, 1859), & [after Westwood, 1895, pl. 12: 1] 80b. Tribe Stephanacridini: Hermarchus pythonius (Westwood, 1859), % [after Westwood, 1895, pl. 35: 3] 81. Tribe Stephanacridini: Stephanacris brevipes Redtenbacher, 1908, & [after Redtenbacher, 1908, pl. 21: 4b]



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FIGURE 82. Geographical distribution of the subfamily Clitumnini, Medaurini trib. nov. and Pharnaciini and Stephanacridini (Phasmatidae s. l.)

Eggs (Fig. 55): Eggs are very diverse and exhibit a wide variety of different shapes, most of which can be regarded specializations due to evolutionary pressure (overall-length 2.5–11.0 mm). Most are of a bark or grass-seed like appearance (Fig. 55a). The capsule is increasingly elongated and up to 4x longer than wide. It is ± strongly laterally flattened with the cross-section ranging from oval to almost rectangular. In some cases the lateral surfaces of the capsule are flat and more or less parallel-sided. The capsule surface is either minutely granulose or strongly wrinkled or rugulose. The operculum is flat and has a ± strongly raised, setose, dentate or crown-like rim, or a prominent peripheral extension on the outer margin. The micropylar plate is small and covers less than half of the capsule length. It is either elongate and pointed towards the anterior end, suboval or transverse and bilobed. Internally the plate exhibits a wide posteromedian notch and a distinct median line (Fig. 55b). Comments: The tribe Baculini was established by Günther (1953: 555) and based on the generic name Baculum Saussure, 1861. Günther distinguished Baculini (= Clitumnini Brunner v. Wattenwyl) from the other tribes of Phasminae (= Phasmatinae sensu Bradley & Galil) by the following two features: both sexes completely lacking wings and median segment much shorter than metanotum. Saussure (1861: 127) however intended to use Baculum as a subgenus of Bacillus Latreille, 1825 and for a species from Brazil Bacillus (Baculum) ramosum, which is the type species by monotypy. Kirby (1904a: 327) introduced an erroneous usage of Baculum by synonymizing it with Clitumnus Stål, 1875, an Oriental genus. Furthermore, Kirby unnecessarily selected Bacillus cuniculus Westwood, 1859 as the type species although this species had not been included in


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Baculum by Saussure. All subsequent authors accepted Kirby’s error until clarification by Bragg (1995c), Hennemann (2002) and Zompro (2005). Bragg (1995c) discussed the usage of the generic names Baculum Saussure, 1861, Ramulus Saussure, 1862 and Gratidia Stål, 1875 and changed the tribal name Ramulini to Gratidiini. Hennemann (2002) re-established Cuniculina Brunner v. Wattenwyl, 1907 (Type species: Bacillus cuniculus Westwood, 1859: 9, pl. 6: 1, 2), discussed the status of Ramulus Saussure, 1861 (Type species: Bacillus (Ramulus) humberti Saussure, 1862: 472, fig. 2), provided new descriptions for both genera and stated Baculum Saussure, 1862 to be a Neotropical genus. Zompro (2005) discussed the status of Baculum Saussure and re-described the type species B. ramosum Saussure from Brazil. As Baculum Saussure is no longer a member of the group which Günther termed Baculini, and Baculum has been transferred to the neotropical subfamily Cladomorphinae, the tribal name Baculini is no longer available for the group Günther intended to define. So, the senior tribal name Clitumnini introduced by Brunner v. Wattenwyl (1907: 181) is the valid name for the Oriental taxa which Günther included in Baculini (Cliquennois, 2004). Three genera formerly attributed to Clitumnini, namely Interphasma Chen & He, 2007, Medaura Stål, 1875 and Medauroidea Zompro, 2000, differ considerably from all other representatives of the tribe by features of the %% genitalia and eggs and are here transferred to Medaurini trib. nov. (→ 5.3). The genus Gongylopus Brunner v. Wattenwyl, 1907 was obviously misplaced in Pachymorphinae: Gratidiini (Cliquennois, 2004: 19) and is here transferred to Clitumnini. Gharianus Werner, 1908 was removed from Clitumnini and transferred to Pachymorphinae: Gratidiini (→ 4.6.1) by Cliquennois (2004: 24). Distribution (Fig. 82): Members of Clitumnini are found throughout the entire Oriental Region and Wallacea as far east as New Guinea. The most northern dispersals are Northern China, Korea and Japan in the Palearctic Region. Genera included: 1. Cuniculina Brunner v. Wattenwyl, 1907: 196. Type species: Bacillus cuniculus Westwood, 1859: 9, pl. 6: 2, by subsequent designation of Hennemann, 2002: 63. 2. Ectentoria Brunner v. Wattenwyl, 1907: 211. Type species: Entoria bilobata Brunner v. Wattenwyl, 1893: 90, pl. 3: 30a, 30 b, by monotypy. 3. Entoria Stål, 1875: 15. Type species: Entoria denticornis Stål, 1875: 72, by subsequent designation of Kirby, 1904a: 327. 4. Erringtonia Brunner v. Wattenwyl, 1907: 212. Type species: Erringtonia malaccensis Brunner v. Wattenwyl, 1907: 212, pl. 9: 2, by monotypy. 5. Gongylopus Brunner v. Wattenwyl, 1907: 233. Type species: Gongylopus adiposus Brunner v. Wattenwyl, 1907: 233, pl. 9: 7, by monotypy. [Here transferred from Pachymorphinae: Gratidiini] 6. Mesentoria Chen & He, 2007: 323. Type-species: Mesentoria acuticaudata Chen & He, 323, figs. 292 a-e, by original designation. 7. Metentoria Brunner v. Wattenwyl, 1907: 211. Type species: Metentoria regina Brunner v. Wattenwyl, 1907: 211, by subsequent designation of Zompro, 2004a: 314. 8. Parabaculum Brock, 1999: 127. Type species: Parabaculum pendleburyi Brock, 1999: 128, figs. 86a–h, by original designation. 9. Paraentoria Chen & He, 1997: 113. Type species: Paraentoria sichuanensis Chen & He, 1997: 114, figs. 1a–c, by original designation. 10. Prosentoria Brunner v. Wattenwyl, 1907: 207. Type species: Prosentoria arrogans Brunner v. Wattenwyl, 1907: 207, pl.11: 1a–c, by subsequent designation of Bragg, 2001: 371. 11. Paraleiophasma Chen & He, 2007: 321. Type species: Leptynia xinganense Chen & He, 1993: 36, by original designation. = Leiophasma Chen & He, 2007: 320. Type-species: Leiophasma yunnanense Chen & He, 2007: 321, by original designation. [Preoccupied by Leiophasma Uvarov, 1940: 114 (Phasmatodea: Anisacanthidae) --> For explanation see addendum] syn. nov.



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12. Ramulus Saussure, 1862: 471. Type species: Bacillus (Ramulus) humberti Saussure, 1862: 472, fig. 2, by monotypy. [Originally established as a subgenus of Bacillus Latreille, 1825] = Clitumnus Stål, 1875: 9. Type species: Phasma (Bacteria) nematodes de Haan, 1842: 132, pl. 11: 6, by subsequent designation of Kirby, 1904a: 327. [Synonymised with the Neotropical Baculum Saussure, 1861 by Kirby, 1904a: 327] = Dagys Günther, 1935c: 3. Type species: Dagys balia Günther, 1935c: 3, pl. 1: 1, by original designation. [Synonymised by Hennemann, 1998: 124] = Dubreuilia Brunner v. Wattenwyl, 1907: 208. Type species: Paraclitumnus lineatus Brunner v. Wattenwyl, 1893: 91, pl. 4: 31a–c, by indication. [Unnecessary replacement name for Paraclitumnus Brunner v. Wattenwyl, 1893, → see Hennemann, 2002: 68] = Paraclitumnus Brunner v. Wattenwyl, 1893: 91. Type species: Paraclitumnus lineatus Brunner v. Wattenwyl, 1893: 91, pl. 4: 31a–c, by subsequent designation of Kirby, 1904a: 330. [Synonymised by Hennemann, 2002: 68] 13. Rhamphophasma Brunner v. Wattenwyl, 1893: 92. Type species: Rhamphophasma modestum Brunner v. Wattenwyl, 1893: 93, pl. 4: 32a–d, by monotypy. 14. Woodmasonia Brunner v. Wattenwyl, 1907: 211. Type species: Bacillus oxytenes Wood-Mason, 1873: 48, pl. 5: 5, by monotypy.

5.3 Medaurini trib. nov. (Figs. 26–28, 56) Type genus: Medaura Stål, 1875: 69.

Description: Small to medium-sized (body length 28.4–110.0 mm), robust to moderately slender Clitumninae with the legs very long in relation to the body. Sexual dimorphism moderate with %% more slender but just little shorter than &&. Body surface smooth to granulose in %% or granulose, tuberculose or even spinose in && (thorax in particular). Vertex set with a variable number of granules, tubercles or spines and usually with a pair of ± distinct spines or scale-like tubercles between the eyes. Antennae consisting of < 28 segments and at best reaching to posterior margin of mesonotum. Scapus ± enlarged and dilated laterally. Mesothorax at best 2.5x (&&) or 3x (%%) longer than head and pronotum combined, in && of certain taxa gently widening towards the posterior or swollen medially. Mesopleurae of && granulose or tuberculose. Median segment wider than long, at best 1/3 as long as metanotum. Abdomen about as long as head and thorax combined; sometimes swollen medially. Abdominal segments II–IV distincly increasing in length, the longest segments (V and VI) at best 3x longer than wide (some %%). In Cnipsomorpha all abdominal tergites bear a triangular posterolateral tooth or lobe. Abdominal tergite IX of && with a ± distinct posteromedian tubercle or hump. Abdominal sternum VII of && with a ± distinct praeopercular organ formed by a posterior pair of tubercles, spines or lobes. Anal segment of %% longitudinally split and consisting of two movable semi-tergites. These ± triangular in lateral aspect with the apex acute and the interior surface set with minute teeth (= “Dornenfeld”); lower margins cleft ventro-basally. Certain species of Parapachymorpha have an additional finger-like ventro-apical appendix on the lower margin. External vomer produced but rather small and slender, sometimes filiform or papillate (Medauroidea and Parapachymorpha). Cerci of both sexes small, distinctly shorter than the anal segment and oval in cross-section; the apex may be laterally compressed. Subgenital plate of && rather small and scoop-shaped, longitudinally keeled and not reaching to apex of anal segment. Legs very long and slender. Profemora ± triangular in cross-section with the anterodorsal carina ± conspicuously raised. Medioventral carina distinct and acute, slightly lamellate and displaced towards anteroventral carina. Anterodorsal and posteroventral carinae may be serrate (Medauroidea). Mid and hind-legs mostly unarmed but


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sometimes with a few dorsal lobes (Medaura and Medaurouidea) and in one case armed with several enlarged teeth (Cnipsomorpha). Medioventral carina of meso- and metafemora indistinct and unarmed. Tibiae often with dentations in apical portion. Tarsi rather short and simple. Eggs (Fig. 56): Small (overall-length < 4.0 mm) ovoid to almost spherical with the capsule minutely punctured or covered with a raised net-like structure. The operculum is either flat and slightly impressed in the centre or elevated and evaginated into a pseudocapitulum (Fig. 56a). The micropylar plate is small, oval or rhomboidal and covers less than half of the capsule length. Internally the plate exhibits a wide posteromedian notch and distinct median line (Fig. 56b). Comments: Morphology of the %% genitalia and eggs has revealed two distinct generic groups within Clitumnini, which indicates the tribe as presently treated to be polyphyletic. The two genera Medaura and Medauroidea differ from all other representatives of Clitumnini by the produced and sclerotised external vomer of %% and ovoid to spherical eggs. Cliquennois (2004) critically discussed the tribe Gratidiini Bragg, 1995 and showed this taxon to be polyphyletic (→ 4.5.1). Cliquennois (2004) and other authors (e.g. Tilgner, 2002 & Whiting et al., 2003) suggested parts of Pachymorphinae: Gratidiini to be related to Clitumnini. Indeed, Gongylopus Brunner v. Wattenwyl, 1907 was misplaced in Gratidiini and is here transferred to Clitumnini. Three further genera, namely Parapachymorpha, Interphasma and Cnipsomorpha, are apparently closely related to Medaura and Medauroidea (formerly in Clitumnini) and hence also misplaced in Pachymorphinae: Gratidiini or Clitumninae: Clitumnini. These five genera together share several common features (→ see above) which distinguish them from all other Clitumnini and consequently are transferred to another tribe, the Medaurini trib. nov.. Distribution (Fig. 82): Continental SE-Asia (Thailand, Vietnam, Laos, Kampuchea, China, Myanmar, Bangladesh and E-India) Genera included: 1. Cnipsomorpha Hennemann, Conle, Zhang & Liu, 2008: 54. Type species: Cnipsomorpha erinacea Hennemann, Conle, Zhang & Liu, 2008: 58, by original designation. [Here transferred from Pachymorphinae: Gratidiini] 2. Interphasma Chen & He, 2007: 328. Type-species: Interphasma lushanense Chen & He, 2007: 329, figs. 297 a-d, by original designation. [Here transferred from Clitumnini] 3. Medaura Stål, 1875: 69. Type species: Medaura brunneri Stål, 1875: 69, by subsequent designation of Kirby, 1904: 341. [Here transferred from Clitumnini] = Menaka Wood-Mason, 1877: 342. Type species: Bacillus scabriusculus Wood-Mason, 1873: 55, by monotypy. [Synonymised by Kirby, 1904a: 341] 4. Medauroidea Zompro, 2000: 68. Type species: Clitumnus extradentatus Brunner v. Wattenwyl, 1907: 193, by original designation. [Here transferred from Clitumnini] 5. Parapachymorpha Brunner v. Wattenwyl, 1893: 95. Type species: Parapachymorpha nigra 1893: 96, pl. 4: 35a, 35b, by subsequent designation of Kirby, 1904a: 342. [Here transferred from Pachymorphinae: Gratidiini]

5.4 Subfamily Lonchodinae Brunner v. Wattenwyl, 1893 Type genus: Lonchodes Gray, 1835: 19. Lonchodidae Brunner v. Wattenwyl, 1893: 80. Lonchodinae Rehn, 1904: 38. Kirby, 1904a: 317. Günther, 1953: 548.



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Bradley & Galil, 1977: 180. Bragg, 2001: 431. Otte & Brock, 2005: 30. Prisomerinae Karny, 1923: 236.

Lonchodinae Brunner v. Wattenwyl, 1893 is a very large subgroup of the anareolate Phasmatodea which presently contains 37 genera (→ see list below) and more than 300 described species. It shows a wide geographical range and extends from the Seychelles and India as far east as to New Guinea, the Solomon Islands and northern Australia. The most northern disperseals are Japan and central China. Günther (1953: 560) characterized Lonchodinae by the following characters: anal segment of %% split and bilobed, or at least with two finger-shaped posterior processes; median segment considerably shorter than the metanotum; antennae long and filiform, usually as long as, or longer than body; ventral carinae of meso- and metafemora not distinctly serrate; both sexes apterous. Bradley & Galil (1977) placed Lonchodinae in Heteronemiidae and currently, Lonchodinae is treated as a subfamily of the Diapheromeridae Kirby, 1904 (Zompro, 2001a: 192). Several features of the genital exosceleton of Lonchodinae however clearly prove this subfamily is not a member of Diapheromeridae, but is closely related to the Old World subfamily Clitumninae of (Phasmatidae s. l.). These are in particular the longitudinally split anal segment of %% which consists of two interiorly dentate (= “Dornenfeld”) and reduced or lacking external vomer, both of which correspond to Clitumninae (→ 5.). The profemora of Diapheromeridae are triangular in cross-section with the medioventral carina distinct, ledge or lamella-like and ± conspicuously displaced towards the anteroventral carina, && have a praeopercular organ on sternum VII and the eggs have an open internal micropylar plate and conspicuous opercular structures, but the genital structures are strikingly distinct. %% of Diapheromeridae have the anal segment consisting of a single plate and never tectiform or split, with the posterior margin at best notched medially. The external vomer is either well developed and seen to be a sclerotised, roughly triangular or hooklike organ, or it is functionally replaced by strongly elongated often highly specialized cerci which enable %% to grasp the & abdomen during copulation. Furthermore, as at least Diapheromerinae are restricted to the New World close relationship to Lonchodinae is rather unlikely. As a result, the striking genital exosceleton of the %% obviously places Lonchodinae near Clitumninae, hence it must be regarded a sub-ordinate taxon of Phasmatidae sensu lato rather than Diapheromeridae. Although several features confirm a close relationship between Lonchodinae and Clitumninae (→ see below), Lonchodinae is well distinguished by the longer, filiform antennae which consist of more than 50 segments (exception Pericentrus Redtenbacher, 1908) and exceed half of the body length, as well as the lack of dentations or serrations of the legs. Günther (1953: 560) divided Lonchodinae into two tribes, and in discussing the tribe Lonchodini, noted “Es liegt für diese Tribus nahe, an eine engere Verwandtschaft zwischen ihr und den Necrosciinae zu denken, zu denen sie in ähnlicher Beziehung stehen könnten, wie die Baculini [= Clitumnini] zu den Pharnaciini in der subfam. Phasminae [= Phasmatinae]” [For this tribe it seems appropriate to consider a closer relationship to Necrosciinae, to which they might stand in a similar relation as do Baculini to Pharnaciini in the subfamily Phasminae]. For the second tribe of Lonchodinae, Neopromachini, Günther (1953: 561) supposed a possible relationship to the subfamily Eurycanthinae (family Phasmatidae). Bradley & Galil (1977: 182) misinterpreted Günther’s Neopromachini and erroneously changed the name to Menexenini based on Menexi Brunner v. Wattenwyl, 1893. However, Günther had definitely placed the Menexenus Stål, 1875 in the tribe Lonchodini. Zompro (2001b) transferred three of the five genera which Günther placed in Neopromachini to Eurycanthinae (these are: Neopromachus Giglio-Tos, 1912, Brachyrtacus Sharp, 1898 and Eupromachus Brunner v. Wattenwyl, 1907). As these include the type genus of Neopromachini, Neopromachus, Zompro transferred Neopromachini to Eurycanthinae but did not accept it as a separate tribe. Up to date Lonchodinae was not subdivided into tribes, but based on differences of genital structures Hennemann (2007: 24) united several genera in a generic group provisionally termed the “Neohirasea-complex”.


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Lonchodinae is quite heterogeneous both in egg morphology and several features of the insects, and so far no author has attempted to subdivide it into tribes, an exercise long overdue. The morphology of the eggs and genitalia of the insects indicate it will become necessary to subdivide Lonchodinae into at least three distinct tribes but this should be subject to a complete revision of Lonchodinae on the generic level. However, as the “Neohirasea-complex” differs from all remaining genera of Lonchodinae, this group certainly represents a distinct tribal unit within the subfamily. Due to some genera (e.g. Mortites Günther, 1935) are probably misplaced in Lonchodinae and belong to other subfamilies, and the status of the “Neohirasea-complex” is questionable (→ Neohiraseini trib. nov., see below), Lonchodinae as presently treated appears to be nonmonophyletic. Moreover, two predominantly New Guinean genera currently contained in the subfamily Necrosciinae (Leprocaulinus and Phenacocephalus) obviously belong in Lonchodinae: Lonchodini and are here transferred. This is emphasized by numerous features like the longitudinally split anal segment of %% which consists of two movable semi-tergites, lack of an external vomer, laterally compressed meso- and metafemora, dorsally lobed two basal tarsomeres, lamellate dorsal carina of the protibiae, and ovoid eggs with a stalked capitulum and an open micropylar plate. All of these features indicate relation to e.g. Hermagoras, Austrocarausius or sections of Carausius. Affinity of Leprocaulinus to the subfamily Lonchodinae was already suggested by Clark-Sellick (1997a: 106 & 1998: 216) who included Leprocaulinus in a “Carausiuscomplex” of Lonchdinae, but did not formally transfer the genus. Examination of Cladomimus from the Loyalty Islands has shown this genus to be misplaced in Pharnaciini and to belong in Lonchodinae: Lonchodini. The elongated and apically pointed anal segment suggests it is close to the Australian Hyrtacus. The laterally compressed meso- and metafemora, dorsally lobed two basal tarsomeres and lamellate dorsal carina of the protibiae show affinity to e.g. Leprocaulinus, Phenacocephalus or Hermagoras and clearly place Cladomimus in the Lonchodinae. Carl (1915: 192) stated the antennae of the holotype of Cladomimus griseus Carl, 1915 (the only specimen known of the entire genus) to be shorter than the fore legs and to consist of only 25 segments, but examination of the specimen has proven them to be broken. They must therefore be assumed to have been considerably longer when complete.

5.4.1 Lonchodini Brunner v. Wattenwyl, 1893 (Figs. 29–31, 58) Type genus: Lonchodes Gray, 1835: 19. Lonchodini Brunner v. Wattenwyl, 1907: 239. Günther, 1953: 560. Bradley & Galil, 1977: 181.

The vast majority of genera contained in Lonchodinae, which include the type genus Lonchodes Gray, 1835, share numerous common morphological features. As stated above some of these strikingly resemble the corresponding structures in Clitumninae and are most certainly homologous, thus indicating much closer relationship to this subfamily rather than to the New World Diapheromeridae. Since five genera, that strikingly differ from the all other Lonchodinae in several aspects, are here transferred to Neohiraseini trib. nov., a tribal name becomes necessary for the remainder genera. Lonchodini was introduced by Brunner v. Wattenwyl (1907: 239) but technically not needed, since Lonchodinae was not sub-divided into tribes. Diagnosis: %% of Lonchodini have the anal segment distinctly tectiform, longitudinally split and consisting of two ± elongate often finger-like semi-tergites, which are dentate interiorly (= “Dornenfeld”). They are very similar in structure to those seen in Clitumninae and form a conspicuous clasping apparatus used to grasp the & abdomen during copulation. The external vomer is strongly reduced or lacking and its morphological function during the copulation replaced by the dentate, movable semi-tergites of the anal segment. && exhibit



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a ± distinct praeopercular organ on abdominal sternum VII, which serves as an anchorage for the anal semitergites of the %%. The subgenital plate of && is strongly keeled, boat-shaped but never considerably elongated or lanceolate and extending considerably over the apex of the abdomen. In some genera && may have the supraanal plate considerably elongated (e.g. certain species of Lonchodes Gray, 1835, Hermagoras Stål, 1875 or Mnesilochus Stål, 1877). As an exception, && of the Philippine Manduria Stål, 1877 have the supraanal plate and subgenital plate elongated to form a bird-beak like ovipositor. Ocelli and wings are lacking in both sexes, although scale-like, rudimentary alae may be present in %% of certain genera (e.g. Chondrosthetus Kirby, 1896). The body is mostly very elongate, stick-like and cylindrical (%% in particular), the mesothorax being distinctly longer than the head and pronotum combined. The median segment is shorter than the metanotum. The sexual dimorphism is rather obvious with %% being much more slender than &&. The body surface is in most genera densely granulose to extremely verrucose or may even be armed with spines (body much less armed and structured in %%). The antennae are long, filiform and 2/3 as long as the body (often much longer), and consist of at least 50 segments (exception Pericentrus). The apical 15–25 antennomeres are strongly shortened. The legs are never serrate or dentate as in Clitumninae and merely bear one or two pairs of sub-apical spines on the two outer ventral carinae of the meso- and metafemora. Occasionally however, prominent tooth or leaf-like lobes may be present, and/or the fore legs have several carinae considerably elevated and lamellate (&& in particular). The meso- and metafemora are often considerably laterally compressed and lobate dorsally, the medioventral carina being unarmed. In several genera the dorsal carinae of the tibiae as well as the meso- and metafemora are melted to form a single dorsal keel (e.g. Lonchodes, Hermagoras or Mnesilochus). The profemora are curved and compressed basally and ± distinctly triangular in cross-section with the dorsal carinae conspicuously nearing and the anterodorsal carina ± decidedly raised. As in Clitumninae the medioventral carina is distinct, often elevated and ledge- or lamella-like and conspicuously displaced towards the anteroventral carina. Several taxa are real masters of camouflage being strongly procryptic and adapted to their natural environments. && in particular exhibit a wide range of conspicuous specialisations like e.g. wart-like swellings or lobes on the body which is accompanied by remarkable intraspecific variation (e.g. Hermagoras or Phenacephorus). The eggs as well indicate close relationship between Lonchodinae and Clitumninae, having an open internal micropylar plate with the micropylar stalk placed in a posteromedian incision or gap of the plate (Fig. 58b). The capitulum bears a more or less distinct hat or knob-like, often stalked capitulum, similar in structure to those seen in Clitumninae: Pharnaciini (Fig. 58a). Genera included: 1. Austrocarausius Brock, 2000: 51. Type species: Lonchodes nigropunctatus Kirby, 1896: 453, by original designation. 2. Breviphetes Zompro, 1998: 163. Type species: Periphetes rammei Günther, 1929: 661, pl. 2: 1 & 2, by original designation. 3. Carausius Stål, 1875: 8. Type species: Carausius strumosus Stål, 1875: 64, by subsequent designation of Rehn, 1904: 42. = Dixippus Stål, 1875: 9. Type species: Phasma (Bacteria) crawangense de Haan, 1842: 132, pl. 13: 1, by subsequent designation of Rehn, 1904: 41. [Synonymised by Bragg, 1992: 129] 4. Chondrostethus Kirby, 1896: 455. Type species: Chondrostethus woodfordi Kirby, 1896: 455, by monotypy. 5. Cladomimus Carl, 1915: 192. Type species: Cladomimus griseus Carl, 1915: 192, by monotypy. [Here transferred from Clitumninae: Pharnaciini] 6. Denhama Werner, 1912: 53. Type species: Denhama aussa Werner, 1912: 53, by monotypy. 7. Echinoclonia Carl, 1913: 43, pl. 1: 4 & 5. Type species: Echinoclonia borneensis Carl, 1913: 43 (= Apora laetior Brunner v. Wattenwyl, 1907: 242), by monotypy. = Apora Brunner v. Wattenwyl, 1907: 241. Type species: Apora laetior Brunner v. Wattenwyl, 1907: 242, by subsequent designation of Bragg, 1993: 34. [Synonymised by Bragg, 1993: 34 – preoccupied by Apora


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Gunnerus, 1768 (Echinodermata)] 8. Echinothorax Günther, 1932b: 757. Type species: Menexenus gazellae Brunner v. Wattenwyl, 1907: 246, pl. 11: 4, by original designation. 9. Greenia Kirby, 1896: 456. Type species: Lonchodes furcatus Bates, 1865: 335, pl. 44: 5, by monotypy. 10. Hermagoras Stål, 1875: 8. Type species: Lonchodes personatus Bates, 1865: 336, pl. 44: 7, by subsequent designation of Kirby, 1904a: 322. = Hermogenes Kirby, 1896: 456. [Misspelling of Hermagoras Stål, 1875] 11. Hyrtacus Stål, 1875: 10. Type species: Hyrtacus tuberculatus Stål, 1875: 67, by subsequent designation of Kirby, 1904a: 331. = Marcenia Sjöstedt, 1918: 37. Type species: Marcenia cunctatrix Sjöstedt, 1918: 37, fig. 1, by subsequent designation of Vickery, 1983: 8. [Synonymised by Brock & Hasenpusch, 2007: 70] 12. Leprocaulinus Uvarov, 1940: 176. Type species: Leprocaulus alte-cornutus Redtenbacher, 1908: 473, by indication. [Replacement name for preoccupied Leprocaulus Redtenbacher, 1908. Here transferred from the subfamily Necrosciinae] = Leprocaulus Redtenbacher, 1908: 473. Type species: Leprocaulus alte-cornutus Redtenbacher, 1908: 476, by subsequent designation of Hennemann, 1998: 121. [Preoccupied by Leprocaulus Fairmaire, 1896 (Coleoptera)] 13. Lonchodes Gray, 1835: 19. Type species: Lonchodes brevipes Gray, 1835: 19, by subsequent designation of Kirby, 1904a: 321. = Phasgania Kirby, 1896: 461. Type species: Phasgania everetti Kirby, 1896: 461, pl. 40: 2, 2a, by original designation. [Synonymised by Bragg, 2001: 435] = Staelonchodes Kirby, 1904: 372. Type species: Lonchodes geniculatus Gray, 1835: 19, by original designation. [Synonymised by Brock, 1999: 169] 14. Lonchodiodes Hennemann & Conle, 2007: 28. Type species: Lonchodiodes samarensis Hennemann & Conle, 2007: 44, figs. 68–76 & 137, by original designation. 15. Manduria Stål, 1877a: 40. Type species: Lonchodes systropedon Westwood, 1859: 44, by original designation. 16. Matutumetes Hennemann & Conle, 2007: 60. Type species: Matutumetes amoenus Hennemann & Conle, 2007: 61, figs. 97–102, by original designation. 17. Menexenus Stål, 1875: 18. Type species: Phasma (Acanthoderus) lacertinum Westwood, 1848: 80, pl. 39: 6, by monotypy. 18. Mithrenes Stål, 1877a: 39. Type species: Mithrenes asperulus Stål, 1877a: 39, by monotypy. 19. Mnesilochus Stål, 1877a: 39. Type species: Mnesilochus capreolus Stål, 1877a: 39, by subsequent designation of Rehn, 1904: 320. 20. Mortites Günther, 1935c: 13. Type species: Mortites enarges Günther, 1935c: 13, pl. 2: 9, by original designation. 21. Myronides Stål, 1875: 8. Type species: Lonchodes pfeifferae Westwood, 1859: 44, pl. 5: 6 & 6a, by subsequent designation of Rehn, 1904: 38. 22. Paraprisomera Hennemann, 2002: 46. Type species: Clitumnus coronatus Brunner v. Wattenwyl, 1907: 251, by original designation. 23. Pericentropsis Günther, 1936: 336. Type species: Pericentropsis aculeata Günther, 1936: 336, fig. 13, by original designation. 24. Pericentrus Redtenbacher, 1908: 351. Type species: Pericentrus moewisi Redtenbacher, 1908: 351 (= Phasma (Acanthodera) bicoronata Westwood, 1848: 15, pl. 7: 2, by subsequent designation of Hebard, 1923: 359. 25. Periphetes Stål, 1877a: 40. Type species: Phasma graniferum Westwood, 1859: 35, pl. 2: 4 & 4a, by original designation.



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26. Phenacephorus Brunner v. Wattenwyl, 1907: 291. Type species: Phenacephorus cornucervi Brunner v. Wattenwyl, 1907: 291, by subsequent designation of Bragg, 1994: 232. 27. Phenacocephalus Werner, 1930: 179. Type species: Phenacocephalus coronatus Werner, 1930: 179. [Here transferred from the subfamily Necrosciinae] 28. Phraortes Stål, 1875: 8. Type species: Phasma elongata Thunberg, 1815: 299, by monotypy. 29. Prisomera Gray, 1835: 15. Type species: Prisomera spinicollis Gray, 1835: 15, by subsequent designation of Kirby, 1904a: 323. 30. Pseudostheneboea Carl, 1913: 25. Type species: Pseudostheneboea segregata Carl, 1913: 26, pl. 1: 2, by subsequent designation of Hennemann, 1998: 102. 31. Spinophetes Zompro & Eusebio, 2000: 131, figs. 1–6. Type species: Spinophetes spinotergum Zompro & Eusebio, 2000: 132, by original designation. 32. Stheneboea Stål, 1875: 10. Type species: Stheneboea malaya Stål, 1875: 68, by subsequent designation of Kirby, 1904a: 324. = Sthenobaea Kirby, 1904a : 323. [Misspelling of Stheneboea Stål, 1875]

5.4.2 Neohiraseini trib. nov. (Fig. 59) Type genus: Neohirasea Rehn, 1904: 84.

Diagnosis: The five genera placed in the “Neohirasea-complex” by Hennemann (2007: 24) differ considerably from all other members of Lonchodinae and violate the key features of this subfamily in several aspects. %% have the anal segment not split but only with a more or less conspicuous posteromedian indention. It consists of single sclerite which shows a fine longitudinal median fissure along the dorsal surface and has the posterolateral angles more or less prominently swollen or with a small, finger-like process, which are ventrally armed with a variable number of small teeth (= “Dornenfeld”). The vomer is well developed and represented as a prominent, sclerotised and roughly triangular or hook-like organ, sometimes being highly specialized with a characteristic number and shape of the terminal hooks (e.g. Spinohirasea Zompro). As in other Lonchodinae && usually exhibit a distinct praeopercular organ on abdominal sternum VII, but this is rather differently structured, consisting of two small spiniform projections (e.g. Neohirasea) or represented by a prominent characteristically shaped and specialized, setose hump (e.g. Spinohirasea). The subgenital plate of && is small, scoop-shaped and never extends beyond the apex of the abdomen. It is longitudinally keeled but never convex and boat-shaped as in other Lonchodinae. The profemora are rectangular to trapezoidal in crosssection with the anterodorsal carina just indistinctly raised and the medioventral carina distinct but never elevated and roughly midways on the ventral surface of the femur. The meso- and metafemora lack any conspicuous armature or lobes, except for having a few sub-apical spines on the two outer ventral carinae, and the tibiae are entirely unarmed and slender. The tarsi are slender and lack any conspicuous specializations. All representatives are small to medium-sized, rather robust insects with the mesothorax less than 2x the length of the head and pronotum combined and ± conspicuously broadened or swollen, and the dorsal body surface ± tuberculose or spinose. The eggs of these genera are mostly spherical, with a small, shield-shaped to almost circular micropylar plate and lack a conspicuous central capitulum (Fig. 59a). Internally they correspond with other Lonchodinae, having an open internal plate with a distinct median line (Fig. 59b). Comments: The genetic analysis and phylogeny presented by Whiting et al. (2003) placed the genus Neohirasea as a basal taxon of a clade which contains genera currently in the subfamily Necrosciinae (e.g. Lopaphus Westwood, 1859 and Sipyloidea Brunner v. Wattenwyl, 1893). As this study does not reveal any close relationship to members of Lonchodinae, this would suggest the genera of the “Neohirasea-complex” to


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belong in Necrosciinae, but this certainly needs further evaluation. Nevertheless, the result of Whiting et al. (2003) emphasizes the striking differences between the “Neohirasea-complex” and all remaining Lonchodinae and the features summarized above well serve to distinguish the concerned genera from all other Lonchodinae on a tribal level. Therefore, the tribe Neohiraseini trib. nov. is here introduced to contain the five genera previously referred to as the “Neohirasea-complex” (→ see list of genera below, 5.4.1). As relationship with Lonchodini or the subfamily Necrosciinae, as suggested by Whiting et al. (2003), clearly need further clarification Neohiraseini trib. nov. is here retained in Lonchodinae. A key to the genera was provided by Hennemann (2007: 24). Genera included: 1. Andropromachus Carl, 1913: 48. Type species: Andropromachus scutatus Carl, 1913: 49, pl. 1: 1 & 3, by subsequent designation of Zompro, 2002b: 68. = Spiniphasma Chen & He, 2000: 32. Type species: Spiniphasma guangxiense Chen & He, 2000: 32, fig. 1, by original designation. [Synonymised by Hennemann, 2007: 16] 2. Neohirasea Rehn, 1904: 84. Type species: Phasma (Acanthoderus) japonicum de Haan, 1842: 135, pl. 12: 4, by original designation. = Paracentema Redtenbacher, 1908: 477. Type species: Paracentema stephanus Redtenbacher, 1908: 477, by monotypy. [Synonymised by Zompro, 2002b: 68] 3. Pseudocentema Chen, He & Li, 2002 : 106. Type species: Pseudocentema bispinatum Chen & He (in: Chen, He & Li), 2002: 107, figs. 8a & b, by original designation. 4. Qiongphasma Chen, He & Li, 2002: 106. Type species: Qiongphasma jianfengense Chen & He (in: Chen, He & Li), 2002: 106, figs7a & b, by original designation. 5. Spinohirasea Zompro, 2002b: 67. Type species: Spinohirasea crassithorax Zompro, 2002b : 68, figs. 1–7 (= Menexenus bengalensis Brunner v. Wattenwyl, 1907), by original designation.

5.5 Keys to the tribes of Clitumninae and Lonchodinae As discussed above Clitumninae and Lonchodinae are obviously closely related to each other, and therefore the keys to the insects and eggs below cover the tribes of both subfamilies. It may well be Clitumninae and Lonchodinae together form a family of their own separate from Phasmatidae sensu stricto (→ Fig. 367). Lonchodidae was established by Brunner v. Wattenwyl (1893: 80), would be an available family-name and has page priority over Clitumnidae (Brunner v. Wattenwyl, 1893: 87).

&& 1. Antennae with < 35 segments; apical antennomeres not conspicuously shortened; at least apical portion of meso- and metatibiae with dentations or serrations.................................................................2.Clitumninae - Antennae with > 50 segments; antennomeres strongly shortened in apical portion of antennae; legs not dentate or serrate but single carinae often lobate or lamellate................................................4.Lonchodinae 2. Antennae distinctly longer than head and pronotum combined; scapus slender; legs ± densely serrate / dentate; medioventral carina of meso- and metafemora spinose (Fig. 76).................................. Pharnaciini - Antennae at best slightly longer than head and pronotum combined; scapus ± dilated laterally; mid and hindlegs at best with single teeth or serrations; medioventral carina of meso- and metafemora unarmed ..3 3. Vertex and body unarmed; abdominal tergite IX flat (Fig. 78a)................................................... Clitumnini - Vertex and body tuberculose or spinose; abdominal tergite IX with a ± distinct posteromedian hump ........ ...................................................................................................................................... Medaurini trib. nov. 4. Elongate, stick-like insects; body cylindrical and of uniform width; profemora distinctly triangular in



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-

cross-section ................................................................................................................................ Lonchodini Robust insects; mesothorax and abdomen ± swollen and broadened; profemora trapezoidal in cross-section .............................................................................................................................Neohiraseini trib. nov.

%% 1. Antennae with < 35 segments; apical antennomeres not conspicuously shortened; at least apical portion of meso- and metatibiae with dentations or serrations.................................................................2.Clitumninae - Antennae with > 50 segments; antennomeres strongly shorten in apical portion of antennae; legs not dentate or serrate but single carinae often lobate or lamellate .....................................................4.Lonchodinae 2. Antennae at least reaching ¾ along mesonotum; scapus slender; legs ± densely serrate / dentate; medioventral carina of meso- and metafemora spinose (Fig. 77) ........................................................ .Pharnaciini - Antennae at best reaching half way along mesonotum; scapus ± dilated laterally; mid and hindlegs at best with single teeth or apical serrations on tibiae; medioventral carina of meso- and metafemora unarmed...3 3. Anal segment forming a closed tube basally; external vomer lacking (Fig. 41) .......................... Clitumnini - Ventral margins of anal segment cleft basally; external vomer produced (Fig. 28) ..... Medaurini trib. nov. 4. anal segment longitudinally split and consisting of two movable semi-tergites; external Vomer reduced or lacking.......................................................................................................................................... Lonchodini - anal segment not split and consisting of a single plate; external vomer produced....Neohiraseini trib. nov.

1. 2. 3. 4. -

Eggs Operculum with a ± distinctly stalked, hat or knob-like, closed central capitulum .....................................2 Operculum with other structures...................................................................................................................3 Capsule not punctured (Figs. 145–193)....................................................................................... Pharnaciini Capsule ± distinctly punctured (often with a polar mound, Fig. 58) ........................................... Lonchodini Operculum lacking conspicuous raised structures (Fig. 59)......................................Neohiraseini trib. nov. Operculum with distinct structures ...............................................................................................................4 Capsule laterally flattened and ± elongated and of grass-seed or bark-like appearance; operculum with various structures on outer margin (Fig. 55) ................................................................................ Clitumnini Capsule ovoid; operculum with a raised central pseudocapitulum (Fig. 56)................ Medaurini trib. nov.

6 Revision of the tribe Pharnaciini Günther, 1953 Pharnaciini Günther, 1953 Type genus: Pharnacia Stål, 1877a: 40. Pharnaciini Günther, 1953: 555 (in part). Bradley & Galil, 1977: 193 (in part). Bragg, 2001: 376. Otte & Brock, 2005: 13 (in part). Acrophyllini Redtenbacher, 1908: 436 (in part). Giglio-Tos, 1910: 43. Acrophyllidae Bruner, 1915: 41. Bacteridae Brunner v. Wattenwyl, 1893: 80 & 83 (in part). Baculini Günther, 1953: 555 (in part). Clitumnini Brunner v. Wattenwyl, 1907: 181 (in part). Giglio-Tos, 1910: 16 (in part). Carl, 1913: 12 (in part). Hermarchini Zompro, 1997: 38 (in part).


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Lonchodidae Brunner v. Wattenwyl, 1893: 80–81 (in part). Phasminae Karny, 1923: 240 (in part). Phryganistriinae Kirby, 1904a: 358 (in part). Kirby, 1904b: 432 (in part).

Description: Large to very large (body length 113–357 mm), stick-like ± slender Phasmatinae; body cylindrical. Sexual dimorphism obvious. No ocelli. Antennae shorter than fore legs but distinctly longer than head and pronotum combined and consisting of less than 45 segments, antennomeres longer than wide. Antennae of %% considerably longer in &&. Scapus simple, not considerably dilated laterally. Mesothorax elongate, longer than metanotum and at least 1.5x longer than head and pronotum combined, usually unarmed but rarely covered with minute granules or single spines. Mesosternum simple or with a ± distinct median keel. && apterous. %% either apterous, brachypterous or with well developed alae; alae not reaching apex of abdomen. Anal region of alae plain transparent grey or brown, no conspicuous markings or colouration. Tegmina rudimentary, scale-like or spatulate and at best covering bases of alae. Median segment longer than metanotum in alate %%, as long as or shorter than metanotum in apterous %% and &&; > 1/8 the length of metanotum. Abdominal segments II–VI ± parallel-sided, of uniform width in %%. Tergite VII usually with more or less prominent lateral lobes or foliaceous dilations (&& in particular). Sternum VII of && often with a ± distinct praeopercular organ; if present then formed by one or two tubercles, spines or ± prominent, triangular lobes at posterior margin. Supraanal plate very small and indistinct, much shorter than anal segment; with a median carina or keel. Anal segment of && with a distinct excavation or notch posteromedially, posterolateral angles ± elevated. Semi-tergites of anal segment of %% usually longer than tergite IX with the apex rounded, triangular or ± strongly elongated and finger-like. Interior surfaces of semi-tergites densely covered with minute teeth in apical portion. Vomer of %% strongly reduced and hardly visible from externally, rarely produced as a filiform organ hidden underneath paraproct. Gonapophyses of && elongate, filiform and up-curving, usually reaching to or slightly projecting over posterior margin of anal segment. Cerci small and considerably shorter than anal segment, cylindrical or oval in cross-section, may be conspicuously laterally compressed and sickle-shaped. Subgenital plate of && strongly keeled, either short and boat-shaped, or considerably elongated and lancetlike. Poculum of %% bulgy and cup-like, sometimes with a ± spine at the angle. Legs elongate, with most carinae ± strongly serrate and/or armed with single enlarged teeth or foliaceous lobes (&& in particular). Medioventral carina of profemora (at least in &&) distinct, strongly elevated and ledge-like. Anterodorsal carina of meso- and metatibiae of && with a ± prominent rounded apical lobe. Basitarsi longer than following three tarsomeres combined, often with a ± strongly raised dorsal carina. Eggs: Capsule medium-sized to large, spherical or lens-shaped, occasionally with a dorsoventral keeled. Surface smooth and glossy, distinctly punctured or otherwise structured. Internal micropylar plate open, median line present. Capitulum hat-like and with a distinct stalk. Distribution: Spread over almost the complete Oriental Region. The centre of distribution with the proportional highest density of taxa is Sundaland. Representatives have so far been recorded from: India, Sri Lanka, Bangladesh, Bhutan, Sikkim, Darjeeling, Myanmar, Southern China and the Hainan Peninsular, Vietnam, Thailand, Laos, Peninsular Malaysia, Singapore, Borneo, Palawan, Sumatra, Banka Island prope Sumatra, Java and the Philippines. Genera included: 1. Baculonistria gen. nov. 2. Pharnacia Stål, 1877a: 40. 3. Phobaeticus Brunner v. Wattenwyl, 1907: 183. = Baculolonga Hennemann & Conle, 1997a: 347. = Lobophasma Günther, 1935: 138. syn. nov. = Nearchus Redtenbacher, 1908: 448. syn. nov. REVISION OF PHARNACIINI


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4. Phryganistria Stål, 1875: 6 & 63. 5. Tirachoidea Brunner v. Wattenwyl, 1893: 83. stat. rev.

Keys to the genera of Pharnaciini Günther, 1953

&& 1. Median segment > ½ the length of metanotum ............................................................................................3 - Median segment < ½ the length of metanotum ............................................................................................2 2. Median segment transverse, about 1/8 the length of metanotum; head with a pair of minute spines between the eyes; mesonotum indistinctly longer than metanotum; Central China ...............Baculonistria gen. nov. - Median segment longer than wide, at least 1/5 the length of metanotum; head unarmed; Mesonotum > 1.5x longer than metanotum .................................................................................................... Phryganistria 3. Massive insects; praeopercular organ indistinct or lacking ..........................................................................4 - Slender insects; praeopercular organ formed by a pair of ± prominent spines or lobes .............Phobaeticus 4. Vertex conically or spherically raised and with a pair of small granules, humps or spines; praeopercular organ formed by a pair of diverging ridges ................................................................................ Tirachoidea - Vertex flat or slightly convex and smooth; without granules, humps or spines; praeopercular organ lacking .......................................................................................................................................................Pharnacia

%% 1. Median segment > ½ the length of metanotum (usually longer than metanotum); fully alate or brachypterous, rarely apterous.......................................................................................................................................3 - Median segment < ½ the length of metanotum; apterous.............................................................................2 2. Median segment transverse, about 1/8 the length of metanotum; mesonotum indistinctly longer than metanotum; antennae distinctly shorter than head, pro- and mesonotum combined; Central China .................. ..................................................................................................................................Baculonistria gen. nov. - Median segment longer than wide, at least 1/5 the length of metanotum; mesonotum > 1.5x longer than metanotum; antennae longer than head, pro- and mesonotum combined ............................... Phryganistria 3 Vertex flat to globose; smooth ......................................................................................................................4 - Vertex conically or spherically raised and with a pair of small granules, humps or spines ....... Tirachoidea 4. Slender insects, mesonotum > 2.5x longer than head and pronotum combined; profemora with prominent black, ± triangular serrations; cerci elongate, ± constricted and in-curving apically; may be brachypterous or apterous ..................................................................................................................................Phobaeticus - More robust insects, mesonotum < 2.5x longer than head and pronotum combined; profemora at best with irregular teeth dorsally; cerci obtuse; alae at least reaching to posterior of abdominal tergite V .Pharnacia

1. 2. 3. 4. -

Eggs External micropylar plate covering more than 2/3 of capsule length ...........................................................3 External micropylar plate covering less than ½ of capsule length ...............................................................2 Capsule laterally compressed, lens-shaped and with a ± distinct dorsoventral keel ..................Phobaeticus Capsule ovoid, without a dorsoventral keel; surface with prominent impressions ..Baculonistria gen. nov. External micropylar plate shaped like an inverted “Y” ................................................................................4 External micropylar plate elongate, lance-shaped and tapered towards anterior end.............. Phryganistria Capsule surface smooth and shiny; polar-area with a black spot and a minute central spine .... Tirachoidea Capsule surface granulose; dull, not shiny; polar-area simple or with a slight dorso-polar depression......... .......................................................................................................................................................Pharnacia


HENNEMANN & CONLE

TABLE 2. Comparison of the genera of Pharnaciini [Key features are highlighted by italic letters]. Pharnacia

Tirachoidea

Phobaeticus

Phryganistria

Baculonistria gen. nov.

Body shape (&&)

Broad to very broad

Moderately slender Slender to very to very broad slender and elongate

Head

Oval, vertex slightly convex & unarmed

Vertex conically raised & bi-tuberculate / spinose

Elongate and oval Oval, distinctly to almost spherical longer than wide & unarmed & unarmed

Oval, distinctly longer than wide & with two minute spines between the eyes

Antennae (%%)

Longer than head, pro- and mesonotum combined




Shorter than head, pro- and mesonotum combined

Mesonotum

> 1.5x longer than metanotum

> 1.5x longer than > 1.5x longer than > 1.5x longer metanotum metanotum than metanotum

Indistinctly longer than metanotum

Metasternum (%%)

± keeled

Keeled

Distinctly keeled

Smooth

Smooth

Median segment (&&)

> 2/3 of metanotum

> 2/3 of metanotum

> 2/5 of metanotum

< 2/5 of metanotum

1/8 of metanotum

Median segment (%%)

Distinctly longer than metanotum

Distinctly longer than metanotum

> 2/5 of metano< 2/5 of metanotum (or longer than tum metanotum)

1/8 of metanotum

Tegmina (%%)

Oval, scale-like & Oval, scale-like & Elongate & spatuwith a ± promiwith a ± promilate; central hump nent, central hump nent, central hump indistinct

Alae (%%)

Fully developed; projecting over tergite IV

Praeopercular organ (&&)

Lacking

Moderately slen- Very slender and der and elongate elongate

Lacking

Lacking

Fully developed; projecting over tergite IV

Fully developed, Lacking reduced or lacking

Lacking

Very indistinct

± prominent; two Very prominent; Indistinct; a single, transverse tubercles, spines or two tubercles, pointed lobes spines or pointed lobe lobes

Tergite VIII (&&) Shorter than IX and X combined

Longer than IX and X combined

Shorter than IX and X combined

Subgenital plate (&&)

At best slightly projecting over apex of anal segment (never lanceolate)

At best slightly projecting over apex of anal segment (never lanceolate)

Variable (occasion- Variable (occaally strongly lan- sionally strongly lanceolate) ceolate)

Poculum (%%)

No distinct central With a distinct hump or spine-like central hump or process spine-like process

Variable; with or without a central hump or spine-like process

About as long as IX and X combined

No distinct central hump or spine-like process

Shorter than IX and X combined Very long and lanceolate

Centre conically elevated

......continued.



83

TABLE 2. (contiund) Pharnacia

Tirachoidea

Anal segment (%%)

Semi-tergites short, broad and rounded or roughly triangular apically

Semi-tergites short Semi-tergites very and broad to mod- elongate, slender and ± finger-like erately slender with apex ± elongated

Vomer (%%)

Rudimentary and Rudimentary and Rudimentary and Rudimentary and Well developed and sclerotised, hidden underneath hidden underneath hidden underneath hidden underY-shaped with neath paraproct paraproct paraproct paraproct apex split

Cerci (%%)

Short & obtuse (occasionally sickle-shaped)

Short & obtuse

Elongate & distinctly in-curving apically

Of moderate length & ± conical

Of moderate length & ± conical

Profemora (%%)

Dentate or irregularly serrate dorsally

Dentate or irregularly serrate dorsally

Prominently serrate dorsally

Serrate dorsally

Smooth dorsally

Probasitarsus (&&)

Short; at best as long as remaining tarsomeres combined

Short; at best as long as remaining tarsomeres combined

Elongate; longer Elongate; longer than remaining tar- than remaining someres combined tarsomeres combined

Probasitarsus (%%)

As long as or slightly longer than remaining tarsomeres combined

As long as or slightly longer than remaining tarsomeres combined

Very elongate; distinctly longer than remaining tarsomeres combined

Very elongate; distinctly longer than remaining tarsomeres combined

Very elongate; longer than remaining tarsomeres combined

Meso- and metabasitarsus

Dorsal carina not Dorsal carina disconsiderably raised tinctly raised and or rounded rounded

Simple or dorsal carina raised and rounded

Simple or dorsal carina gently rounded dorsally

Simple

Mesofemora (length)

Longer than metanotum and median segment combined



Longer than met- As long as metanotum anotum and median segment combined

Egg capsule

Globose & distinctly oval in cross-section

Spherical to slightly longer than wide & ± cylindrical in cross-section

Lens-shaped with a distinct dorsoventral keel; elliptical in cross-section

Oval, longer than Oval, longer than wide & ± ellipti- wide & oval in cal in cross-sec- cross-section tion

Capsule surface

Dull, not shiny & distinctly granulose

Strongly shiny & almost smooth

± shiny to dull & unevenly granulose to strongly sculptured

Slightly shiny & unevenly granulose (40x)

Granulose with numerous, prominent impressions

Polar-area

Rounded or with a With a minute cen- ± distinctly very slight impres- tral tubercle or impressed sion & unarmed spine

Rounded

Impressed

Micropylar plate

Inverted “Y”, apices of extensions ± broadened & truncate

Inverted “Y”, apices of extensions slender & rounded


Phobaeticus

Phryganistria

Baculonistria gen. nov.

Semi-tergites of moderate length, distinctly tapered and slightly elongated apically

Semi-tergites of moderate length, distinctly tapered and slightly elongated apically

Elongate; as long as remaining tarsomeres combined

Heart-shaped; Transverse to elon- Elongate; > 2/3 the length of cap- less than ½ the gate, often bisule length of capsule lobed; < 2/3 the length of capsule

HENNEMANN & CONLE

FIGURES 83–86. Geographical distribution of the genera of Pharnaciini 83. Pharnacia Stål, 1877 84. Phobaeticus Brunner v. Wattenwyl, 1907 85. Phryganistria Stål, 1875 and Baculonistria gen. nov. 86. Tirachoidea Brunner v. Wattenwyl, 1907

6.1 Genus Baculonistria gen. nov. Type species: Baculum album Chen & He, 1990: 54, by present designation. Baculum, Chen & He, 1990: 54. Phobaeticus, Cai & Liu, 1993: 469. Chen & He, 1997: 114, fig. 3a, b.



85

Brock, 1996: 28 (in part). Otte & Brock, 2005: 268 (in part). Chen & He, 2007: 347.

Description (%, &): Medium-sized to long, elongate and moderately slender Pharnaciini, both sexes apterous. Sexual dimorphism moderate. %% either much smaller and more slender than &&. Body surface unarmed and strongly shiny in %%. Head longer than wide and narrowed towards the posterior, vertex flat and smooth. In && with two minute spines between the eyes. Antennae of %% slightly shorter than head, pro- and mesonotum combined, in && conspicuously shortened and just about 1.5x longer than head and pronotum combined. Antennomeres IV–XII of && distinctly shortened, indistinctly longer than wide. Mesothorax of && short, about as long as metanotum and median segment combined. All sterna and pleurae simple, mesosternum not keeled. Median segment very short and wider than long, about 1/8 the length of metanotum. Abdominal tergites II–VIII longer than wide and parallel-sided. Sternum VII of && with a distinct praeopercular organ, formed by a gently rounded, transverse lobe. Semi-tergites of anal segment of %% raised anterodorsally, of moderate length and rounded apically. Vomer of %% produced and sclerotised, roughly Y-shaped with the apex split. Poculum of %% convex and at the angle conically elevated; ± acute. Subgenital plate of && keeled, boat-shaped and not reaching to the tip of the anal segment, or very elongate, lanceolate and projecting over the anal segment by more than 2x the combined length of tergites VIII–X. Cerci small, conical and cylindrical in cross-section. All legs of moderate length and minutely dentate ventrally; smooth dorsally. Anterodorsal carina of %% profemora unarmed. Meso- and metafemora of %% slightly broadened. Mesofemora as long as metanotum (&&) or metanotum and median segment combined (%%). Medioventral carina of meso- and metafemora distinct, sparingly spinose and occasionally with a prominent spine sub-apically. Hind legs of && short and staying clearly beyond apex of abdomen. Probasitarsus as long as (&&) or longer than remaining tarsomeres combined (%%); simple. Meso- and metabasitarsus at least as long as the following three tarsomeres combined; simple. Eggs (Figs. 192–193): Medium-sized. Capsule ovoid, distinctly longer than high and oval in cross-section. Polar area impressed and with longitudinal, keel-like extensions dorsally and ventrally. Anterior end of capsule with a prominent opercular collar. Capsule surface minutely punctate and all over covered with numerous, prominent impressions. External micropylar plate slightly less than ½ the length of capsule, broad and roughly heart-shaped, with a broad notch posteromedially. Open internally. Median line very distinct and conspicuously raised from capsule surface. Operculum flat, oval. Capitulum hat-like and distinctly stalked. Differentiation (Table 2): Distinguished from all other members of Pharnaciini and its closest relative Phryganistria Stål, 1875 by: the short mesonotum which is just indistinctly longer than the metanotum; extremely short and transverse median segment, which is only about 1/8 the length of the metanotum; distinct and spinose medioventral carina of the meso- and metafemora; short mesofemora which are about as long or indistinctly longer than the metanotum (distinctly longer than combined length of metanotum and median segment in all other genera); simple basitarsi and short antennae of both sexes, which are shorter than the head, pro- and mesonotum combined (&& in particular). && furthermore differ from all other genera of Pharnaciini by: having two minute spines between the eyes; distinctly shortened antennomeres IV–XII and having the praeopercular organ formed by a singe, gently rounded transverse lobe at posterior margin of tergite VII (paired lobes in all other genera). %% are additionally distinguished by the unarmed anterodorsal carina of the profemora. The eggs are distinctive by having a very prominent opercular collar, the capsule surface with numerous large impressions and a broad, heart-shaped micropylar plate which covers less than ½ of the dorsal egg surface. Etymology: Feminine. Combined from “baculus” (lat. = twig) and the ending “-nistria” of Phryganistria Stål, 1875 to mirror the close relation with this genus. Distribution (Fig. 85): Central and east China (Sichuan Prov., Chongqing Prov., Henan Prov. & Anhui Prov.). This is the ony genus of Pharnaciini which has dispersed as far north as to the southeastern regions of


HENNEMANN & CONLE

the Palearctic realm. Species included: 1. Baculum album Chen & He, 1990: 54, figs. 1a–c. = Phobaeticus sichuanensis (Cai & Liu, 1993: 469). syn. nov. 2. Phobaeticus longicornis Bi & Wang, 1998: 10, figs. 5–6. 3. Phobaeticus yuexiensis Chen & He, 1993: 54, figs. 1–2.

[China (Sichuan & Chongqing] [China (Henan Prov.)] [China (Anhui Prov.)]

Keys to the species of Baculonistria gen. nov.

&&* 1. Subgenital plate short, staying clearly beyond apex of anal segment; East China........................ yuexiensis - Subgenital plate long and lanceolate, prominently projecting over apex of anal segment; Central China .... ................................................................................................................................................................ alba * && of B. longicornis Bi & Wang, 1998 are not known

%%* 1. Antennae with 15 segments; femora with a longitudinal row of pale elliptical spots laterally; meso- and metafemora without a distinct sub-apical spine ventrally; E-China............................................ longicornis - Antennae with 25 segments; femora without lateral spots; meso- and metafemora with a distinct, sub-apical spine on medioventral carina; Central China .................................................................................... alba * %% of B. yuexiensis Chen & He, 1993 are not known

Baculonistria alba (Chen & He, 1990) comb. nov. (Figs. 87–88, 192–193, 266–268, 351, 358, 368, 416, 429–430) Baculum album Chen & He, 1990: 54, figs. 1 a (%), b–c (&). HT, %: China, Chongqing, Zhongxian County, leg. Zeng Lin 9.X.1989 (BFU); AT, &: China, Chongqing, Zhongxian, leg. Zeng Lin 9.X.1989 (BFU). Phobaeticus albus, Chen & He, 1997: 114, fig. 3a (%) & b (&). Chen, 1999b: 34. Otte & Brock, 2005: 268. Chen & He, 2007: 347, figs. 315 a-d. Baculonistria alba (in press), Hennemann et al., 2008: 9. Phobaeticus sichuanensis Cai & Liu, 1993: 469, figs. 1–4. HT, &: China, Chongqing, Zhongxian, leg. Zeng Lin & Hu Liancheng 9.X.1989 (DBNU); AT, %: China, Chongqing, Zhongxian, leg. Zeng Lin & Hu Liancheng 9.X.1989 (DBNU); PT: 2 %%, 2 &&: same data (DBNU). syn. nov. Zeng, Hu et al., 1990: 1. [Notes on biology and pest status] Brock, 1996: 30. Otte & Brock, 2005: 270. Further material [5 %%, 6 &&, eggs]: CHINA: 1 %, 1 &, 4 eggs: SW-China, Chongqing Municipality, Wanzhou County, Chongqing, nr. Zhongxian County, leg. Weiwei Zhang 1.–8.VIII.2006 (coll. FH, No’s 0614-1, 2 & E); 3 %%, 5 &&, eggs: ex Zucht: R. Galunder (Nümbrecht), Herkunft: SW-China: Chongqing, F1-Generation, 2008 (coll. FH, No’s 0614-3 to 9 & ED).



87

FIGURES 87-88: Baculonistria gen. nov. alba Chen & He, 1993) 87. & SW-China, Chongqing, Zhongxian Co. (coll. FH, No. 0614-1) 88. % SW-China, Chongqing, Zhongxian Co. (coll. FH, No. 0614-2)

Diagnosis: && are easily distinguished from B. yuexiensis (Chen & He, 1993) from E-China by the very long, lanceolate subgenital plate which projects over the apex of the anal segment by more than 2x the combined length of tergites VIII–X, and distinct sub-apical spine on the medioventral carina of the meso- and metafemora. %% differ from B. longicornis (Bi & Wang, 1998) by: the relatively shorter antennomeres; antennae with 25 segments; medioventral carina of meso and metafemora with a distinct sub-apical spine and femora lacking pale elliptical spots. Etymology: The name (lat. albus = white) refers to the white portions of the head and thorax of %%. Description: The descriptions presented below is based on the % and & in coll. FH. The colouration is described from colour-photographs of live specimens taken by Weiwei Zhang (Chongqing, China). && (Fig. 87, 416, 430): Long to very long (body length 159.0–194.5 mm, including subgenital plate 172.0–221.0 mm) and slender for the genus with a very long and lanceolate subgenital plate (33.4–50.5 mm). Body entirely smooth and slightly shiny. General colouration of body and legs greyish or greenish mid brown with paler grey, yellowish or whitish speckles. Meso- and metasternum mid to dark brown with pale yellow markings. Lateral surfaces of the head with a fine dark brown postocular line, the cheeks creamish white. Leg armature black. Antennae brown dorsally and black ventrally. Larger spines of the legs black. Eyes very dark purplish brown. Head: Oval, about 1.4x longer than wide and narrowed towards the posterior, broadest at the eyes. Between the eyes with a pair of minute, forward-pointing black spines. Eyes circular and projecting hemispherically, their length contained almost 3x in that of cheek. Antennae slightly longer than head and pronotum combined and distinctly shorter than profemora; with 25 segments. Scapus dorsoventrally compressed, 2.5x longer than wide and narrowed towards the base. Pedicellus 2/5 the length of scapus, cylindrical. III 1.5x longer than pedicellus, cylindrical. Following antennomeres cylindrical and increasing in length; IV–X conspicuously shortened. Thorax: Pronotum shorter and narrower than the head, narrowed anteriorly and with posterior margin 1.5x broader than anterior margin. Anterior margin raised and with an impression medially. Median transverse depression short, widely V-shaped and not reaching lateral margins of segment. Mesothorax very gently, gradually widening towards the posterior; as long as metanotum and median segment combined. Metanotum indistinctly shorter than mesonotum, parallel-sided and about 4.5x longer than wide. Abdomen: Median segment just slightly more than 1/8 the length of metanotum. Segments II–VI slightly increasing in length; III 2x, VI 2.5x longer than wide. VII gently narrowing towards the posterior and almost


HENNEMANN & CONLE

3x longer than wide. Sternites II–VII smooth. Praeopercular organ formed by a single gently rounded and thickened transverse lobe at posterior margin of sternum VII. Tergite VIII slightly narrower than previous, parallel-sided, 2.5x longer than wide and about 2/3 the length of VII. IV half the length of VIII, slightly longer than wide and strongly convex. Anal segment elongate, 1.4x longer than IX and flattened. Posterior margin with a wide, triangular median excision and the lateral angles rounded (Fig. 268). Supraanal plate prominent, broadly rounded and with a faint longitudinal median keel. Cerci small, finely bristled and distinctly projecting over posterior margin of anal segment. Subgenital plate very elongate, lanceolate and almost 2.5x longer than combined length of tergites VIII–X; apex pointed (Fig. 266). Legs: Of moderate length and slender. Profemora almost as long as head, pro- and mesonotum combined, mesofemora about as long as metanotum and metatibiae slightly projecting posterior margin of abdominal segment VI. Anterodorsal carina of profemora with 12–15 sharp, triangular serrations which decrease in size towards apex of femur, posteroventral carina with 10–12 minute serrations. Protibiae unarmed. Dorsal carinae of meso- and metafemora and tibiae unarmed. Ventral carinae of femora minutely dentate; medioventral carina distinct, armed with 5–7 moderately sized spines and a prominent sub-apical spine. Apical ¾ of ventral carinae of meso- and metatibiae serrate. Probasitarsus elongate, simple and longer than remaining tarsomeres combined. Meso- and metabasitarsu as long as following two tarsomeres combined, simple and unarmed. %% (Fig. 88, 416, 429): Medium-sized (body length 138.5–164.0 mm) and moderately slender insects. Body surface entirely smooth and strongly shiny. General colouration of the body pale greyish green and yellowish along the lateral surfaces, posterior margin of each body segment broadly pale brown. Dorsal portions of meso- and metanotum with a washed white longitudinal median stripe; meso- and metapleurae often whitish. Head white with a distinct dark brown postocular stripe (Fig. 429). Tergites VIII–X pale greyish brown; VIII and IX with a white spot anterolaterally (sometimes lacking on VIII). Fore legs, all tibiae, apices of femora and coxae greyish brown. Remaining portions of meso- and metafemora pale greyish green. Antennae very dark brown with the apex slightly paler. Eyes very dark purplish brown. Head: Generally as in && but lacking minute spines between the eyes. Antennae relatively longer than in &&, almost reaching posterior of mesonotum and with antennomeres relatively longer; 25-jointed. Eyes more prominent. Thorax: Pronotum slightly shorter and narrower than the head, anterior portion gently narrowed and anterior margin slightly raised. Transverse median depression shallow, widely V-shaped and not reaching lateral margins of segment. Mesothorax slightly broadened at the posterior; indistinctly longer than metanotum and median segment combined. Metanotum very gently constricted medially and about 8.5x longer than wide. Abdomen: Median segment about 1.3x longer than wide with the anterior margin slightly rounded; less than 1/7 the length of metanotum. Segments II–VI increasing in length, VI and VII slightly decreasing; V longest. II about 3x, V 4.5x longer than wide. II–VI very slightly constricted medially, VII gently widening toward posterior margin. Tergite VIII trapezoidal, slightly widening toward the posterior and about half the length of VII. IX slightly shorter than VIII, strongly convex and gently constricted medially. Anal segment longer than VIII, laterally compressed, strongly keeled and split, forming two broad, roughly triangular apically rounded semi-tergites (Fig. 267). These slightly swollen and club-like apically and covered with minute black teeth on their interior surfaces. Below anterior margin with a distinct, longitudinal furrow. Cerci elongate, less than half the length of anal segment, cylindrical and finely bristled. Vomer well developed and sclerotised, visible from externally and roughly Y-shaped with the apex split. Poculum small, cup-like, hardly reaching posterior margin of tergite IX and with a distinct, conical hump in the centre. Legs: Of moderate length, the meso- and metafemora conspicuously thickened and distinctly broader than corresponding tibiae. Profemora slightly longer than head, pro- and mesonotum combined, mesofemora about as long as metanotum and median segment combined, metatibiae slightly exceeding apex of abdomen. Profemora with a few very minute teeth on posteroventral carina, otherwise unarmed. Protibiae unarmed. Mesoand metafemora unarmed dorsally, the antero- and posteroventral carinae each irregularly dentate. Medioven-



89

tral carina prominent and armed with 5–6 small spines and a prominent, black sub-apical spine. Meso- and metatibiae smooth dorsally, the three ventral carinae minutely dentate. Probasitarsus very elongate, unarmed and longer than remaining tarsomeres combined. Meso- and metabasitarsi about as long as remaining tarsomeres combined except claw. Eggs (Figs. 192–193): Medium-sized. Capsule ovoid, distinctly longer than high and oval in cross-section. Polar area impressed and with longitudinal, rounded and keel-like extensions dorsally and ventrally. Anterior end of capsule constricted and with a prominent, broad opercular collar. Capsule surface minutely punctate and all over covered with numerous, prominent impressions of variable size and shape. External micropylar plate slightly less than ½ the length of capsule, broad and roughly heart-shaped, with a broad notch posteromedially. In the centre with a broad, roughly heart-shaped and impressed region. Median line very distinct and prominently raised from capsule surface. Operculum flat, oval. Capitulum prominent, hat-like and with a distinct stalk. Capsule greyish brown with the impressions and central, heart-shaped impression of the micropylar plate black. Opercular collar very dark brown. Operculum dark grey to black, capitulum orange. Measurements [mm]: length 4.0, length (including capitulum) 4.7, width 2.5, height 2.9, length of the micropylar plate 1.9. Cai & Liu (1993: 471) cited the following dimensions or specimens from Zhongxian County: length 3.68 mm, width 2.32 mm, height 2.64 mm. Comments: Chen & He (1990: 54) described Baculum album from Zhongxian County, Sichuan Province. Subsequently, Chen & He (1997) recorded it from the Three Gorge Reservoir at the Yangtze River and transferred it to Phobaeticus Brunner v. Wattenwyl, 1907. In 1993 Cai & Liu described Phobaeticus sichuanensis also from Zhongxian County, Sichuan Province, unaware that this species had already been described by Chen & He (1990) based on specimens collected from the same locality. The type specimens of Baculum album Chen & He in BFU and Phobaeticus sichuanensis Cai & Liu, 1993 (syn. nov.) in DBNU could unfortunately not be examined for the present study. However, an adult couple and eggs collected from Wanzhou County, Chongqing were kindly presented to the first author by Weiwei Zhang (Chongqing, China) for examination and match very well with the descriptions of both species, except for being comparatively longer (→Table 3). TABLE 3. Measurements [mm] of Baculonistria alba (Chen & He, 1990) comb. nov.

%, HT (BFU)*

&, AT (BFU)*

&, HT of sichuanensis (DBNU)**

%, AT of sichuanensis (DBNU)**

%%

&&

(coll. FH)

(coll. FH)

Body (incl. subgen. plate) -

-

-

-

172.0–221.0

-

Body

145.6

163.6

176.0

138.6

159.0–194.5

138.5–164.0

Pronotum

5.2

6.0

7.9

5.0

5.8–6.2

4.9–6.3

Mesonotum

30.4

31.6

33.4

29.5

27.6–34.0

25.7–30.2

Metanotum

26.2

28.6

27.9

24.5

22.5–30.8

23.6–28.2

Median segment

3.2

4.2

4.3

3.6

3.7–4.1

3.1–3.7

Subgenital plate

-

-

38.8

-

33.4–50.5

-

Profemora

42.0

29.0

46.0

46.8

37.6–46.3

39.5–48.2

Mesofemora

25.5

27.0

31.7

29.5

25.5–30.0

26.8–31.2

Metafemora

29.0

26.0

34.8

34.0

27.0–35.2

30.3–38.7

Protibiae

-

-

-

-

39.0–60.0

44.6–58.5

Mesotibiae

-

-

-

-

25.0–35.4

27.2–33.8

Metatibiae

-

-

-

-

30.0–42.4

34.2–43.4

Antennae

-

-

-

-

20.7-28.5

40.0-44.0

* according to Chen & He (1990: 55)


HENNEMANN & CONLE

** accordingt to Cai & Liu (1993: 470)

Chen & He (1994) and Cai & Liu (1993) reported this species to have caused considerable damage on a variety of plants in the Zhongxian County in 1986 (Sichuan Province, now Chonqing municipality) and that an area of several hundred hectars was almost completely destroyed. They stated more than one hundred specimens were counted per tree and the following plants to have shown considerable defoliation: Vernicia fordii (Euphorbiaceae), Cupressus funebris (Cupressaceae), Platycarya strobilacea, P. orientalis (Juglandaceae), Sapium sebiferum (Euphorbiaceae), Cyclobalanopsis sp. (Fagaceae), Salix cheilophila (Salicaceae), Zea mays (Poaceae) and Solanum tuberosum (Solanaceae). Zeng et al. (1990: 1ff.) discussed the biology and life-cylce in Chongqing and western Hubei Province and reported B. alba to take three years to complete a generation. Eggs are laid between early July and late August. They take about 11–13 months to hatch, the nymphs will reach 2nd instar in late autumn, over-winter and become adult in the third year. Culture stock from Chongqing municipality was introduced to Germany in 2007 and successfully reared in captivity. At least bramble (Rubus spp., Rosaceae), rose (Rosa spp., Rosaceae) and oak (Quercus robur, Fagaceae) are frequently accepted as alternative foodplants. Eggs have an incubation time of approximately three months and nymphs reach maturity in about 4 months. The insects prefer rather well ventilated conditions and a large cage is necessary for successful breeding. Captive reared && of the F1-generation produced an average of 10 eggs per day (personal communication with R. Galunder). Distribution (Fig. 368): SW-China (Chongqing municipality: Zhongxian County; Yangtze River, Three Gorge Reservoir 160–1200 m & Wanzhou County, Chongqing) and Hubei Province. Number of specimens examined: 11.

Baculonistria longicornis (Bi & Wang, 1998) comb. nov. (Fig. 369) Phobaeticus longicornis Bi & Wang, 1998: 10, figs. 5 & 6 (%). HT, %: China, Henan Province, Luanchan County, Longyuwan, 10.VII.1996 leg. Liu Zurao, Zhang Weinian & Yin Haisheng (SIES). Otte & Brock, 2005: 269. Baculonistria longicornis, Hennemann et al., 2008: 9.

Diagnosis: Differing from %% of B. alba (Chen & He, 1990) by: the relatively longer antennomeres; antennae with only 15 segments; medioventral carina of meso and metafemora without a distinct sub-apical spine and posterolateral surface of the profemora, as well as the antero- and posterolateral surfaces of the meso- and metafemora with a longitudinal row of pale elliptical spots. Etymology: The specific name longicornis is a composition of the Latin words “longus” (= long) and “cornus” (= horn or antenna) and refers to the elongate antennomeres of this species. Description: The following is an improved translation of the original description provided by Bi & Wang (1998: 10), which was kindly provided by Weiwei Zhang (Chonqing, China). %%: Rather robust. General colouration yellowish brown, slightly shiny. Lateral surfaces of head pale yellow. Legs straw the posterolateral surface of the profemora, as well as the antero- and posterolateral surfaces of the meso- and metafemora with a longitudinal row of pale elliptical spots. Head: Oval, the posterior slightly truncate and with a pair of small tubercles between the eyes. Eyes rather small, strongly convex and with a brown longitudinal median line. Dorsal surface of head with two small pits, a fine, impressed coronal line and two short longitudinal impressed lines posteriorly. Two further, deep impressions between the bases of the antennae. Antennae about 2/3 the length of the profemora and consisting of 15 segments. Scapus almost 2x, III almost 3x and IV 2x longer than pedicellus; remaining antennomeres



91

elongate. Thorax: Pronotum shorter than the head, rectangular and with an impressed longitudinal median line. Transverse median depression not reaching lateral margins of segment. Mesothorax moderately broadened posteriorly. Mesonotum as long as combined length of metanotum and median segment, smooth and with a faint longitudinal line which becomes more distinct towards the posterior of segment. Mesopleurae with a few minute granules anteriorly. Metathorax slightly broadened posteriorly. Abdomen: Median segment very short, slightly broader than wide and less than 1/7 the length of metanotum. Segments II–VII almost cylindrical, tergite IX and posterior part of VIII broader. Anal segment laterally compressed, strongly keeled and split, forming two moderately elongate, apically triangular semi-tergites. These strongly laterally compressed towards the apex and covered with minute black teeth on their interior surfaces. Poculum small, convex, cup-shaped and extending about half way along tergite IX. Posterior margin flattened and slightly truncate. Cerci distinct, oval in cross-section, slightly incurving apically and densely bristled. Legs: Bases of profemora curved. Profemora longer than mesonotum, mesofemora almost as long as mesonotum and tibiae longer than corresponding femora. Anterodorsal carina with 8 irregularly placed distinct serrations, ventral carinae unarmed. Protibiae entirely unarmed. Meso- and metafemora as well as the tibiae with all three ventral carinae dentate. Teeth identical in number and arrangement on mid and hind legs. Basitarsi with a few minute teeth ventrally. Comments: The pair of tubercles between the eyes, short antennae, which consist of only 15 segments, and very short median-segment place this species in Baculonistria gen. nov.. Unfortunately, the unique HT in SIES could not be examined for the present study, but the quite detailed original description of Bi & Wang (1998: 10) allows identification and differentiation of this species. With quite some probability B. longicornis is the % of B. yuexiensis (Chen & He) described from a & from the Anhui Province. This is however impossible to decide without seeing the type specimens of both species, and having material of both sexes from the same locality available for examination. This is the northernmost distributed species of the entire tribe Pharnaciini. Eggs unknown. Distribution (Fig. 369): Eastern Central China (Henan Province, Luanchan County, Longyuwan). Number of specimens examined: None TABLE 4: Measurements [mm] of Baculonistria longicornis (Bi & Wang, 1998) comb. nov.

% HT (SIES)* Body

130.0

Pronotum

5.0

Mesonotum

26.0

Metanotum

23.0

Median segment

3.0

Profemora

42.0

Mesofemora

26.0

Metafemora

32.0

* according to Bi & Wang (1998)

Baculonistria yuexiensis (Chen & He, 1993) comb. nov. (Fig. 370)


HENNEMANN & CONLE

Phobaeticus yuexiensis Chen & He, 1993: 54, figs. 1 & 2 (&). HT, Kujingyuan forest farm, 1985, Yu Fangbei leg. (BFU). Baculonistria yuexiensis, Hennemann, Conle & Zhang, 2008: 9.

&:

E-China, Anhui Province, Yuexi County,

Diagnosis: Distinguished from && of B. alba (Chen & He, 1990) from Chongqing and Hubei by: the much shorter subgenital plate which does not reach to the tip of the anal segment, longer and more slender cerci and lack of a prominent sub-apical spine on the medioventral carina of the meso- and metafemora. Etymology: The name refers to the type-locality, Yuexi County in the Anhui Province of east China. Description: The following is a translation of the original description, which was kindly provided by Weiwei Zhang (Chongqing, China). &&: Robust, body smooth and brown. Head: Much longer than wide and conspicuously narrowed towards the posterior. Vertex flat and with an impressed, longitudinal median line and two short lateral impressed lines at posterior margin. Laterally with a dark brown longitudinal postocular line, which reaches to the posterior of the head. Between the eyes with two short horns and a pair of shallow impressions in front. Antennae slightly longer than 1/3 the length of profemora. Antennomeres except caput short. Scapus 2x longer than wide, parallel-sided and with a longitudinal median carina dorsally. Pedicellus short, about ¼ the length of scapus, club-shaped. Thorax: Pronotum shorter than the head and posterior margin broader than anterior margin. Lateral margins indented anteriorly. Transverse median depression distinct and slightly curved to the anterior. Another transverse depression just behind anterior margin and an oblique depression on each side of the transverse median depression. Mesonotum gently gradually broadened towards the posterior and with a faint longitudinal median carina, almost 5x longer than pronotum. Metanotum and median segment together about 8/9 the length of mesonotum. Metanotum with a faint longitudinal median carina in anterior 2/3 of segment. Abdomen: Longer than combined length of head and complete thorax. Median segment shorter than broad and with a pair of indistinct, curved indentions. Segments II–V broader than following, V longest, II–IV of qual length. VIII shorter than VII and IX more than half the length of VIII. Tergites IX and X combined longer than VIII. Pleurae angular. Sternum VII with a prominent praeopercular organ formed by a prominent, spinelike median hump at posterior margin. Anal segment slightly tectiform with a distinct median carina anteriorly a small median indention at posterior margin. Supraanal small, roughly semicircular and with a ± spine-like median hump. Subgenital plate rather short, strongly keeled and boat-shaped, tapered apically and staying slightly beyond the tip of the anal segment. Laterally with a distinct longitudinal carina. Cerci distinct, longer than the supraanal plate and distinctly tapered towards the apex. TABLE 5: Measurements [mm] of Baculonistria yuexiensis (Chen & He, 1993) comb. nov.

& HT (BFU)* Body

164.0

Pronotum

6.4

Mesonotum

34.0

Metanotum

27.6

Median segment

3.2

Abdomen

85.8

Subgenital plate

16.6

Profemora

46.0

Mesofemora

30.0

Metafemora

33.0

* according to Chen & He (1993: 54)



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Legs: All three pairs long. Profemora longer than meso- and metafemora and longer than mesonotum, but shorter than protibiae. Anterodorsal carina with 12 black serrations. Mesofemora slightly shorter than metafemora. All three ventral carinae of the meso- and metafemora and tibiae with black serrations. A few minute teeth on the ventral carinae of the meso- and metabasitarsus. Comments: This species was not listed in Otte & Brock (2005) and Chen & He (2007). The spines between the eyes, short antennae, short mesothorax, very short and transverse median segment and praeopercular organ which is formed by a single median structure on abdominal sternum VII clearly place this species in Baculonistria gen. nov.. Unfortunately the unique HT in BFU could not be examined for the present study, but the quite detailed original description allows identification and differentiation of this species. The unknown % might be represented by B. longicornis (Bi & Wang) comb. nov. from Henan Province. Eggs unknown. Distribution (Fig. 370): East China (Anhui Province, Yuexi County, Kujingyuan forest farm). Number of specimens examined: None

6.2 Genus Pharnacia Stål, 1877 Type species: Pharnacia ponderosa Stål, 1877a: 40, by subsequent designation of Kirby, 1904a: 359. Pharnacia Stål, 1877a: 40. Kirby, 1904a: 359 (in part—only species No. 3). Redtenbacher, 1908: 449 (in part). Giglio-Tos, 1910: 43 (in part). Karny, 1923: 240 (in part). Weidner, 1966: 231. Brock, 1996: 27 (in part). Brock, 1998a: 53. Brock, 1999: 132, figs. 24 & 26 (in part). Seow-Choen, 2000: 35 (in part). Otte & Brock, 2005: 264 (in part). Baculolonga, Bragg, 2001: 377 (in part). Bacteria, Burmeister, 1838: 563 (in part). Cladoxerus, Gray, 1835: 42 (in part). De Haan, 1842: 131 (in part). Clitumnus Brunner v. Wattenwyl, 1907: 187 (in part). Lobophasma, Günther, 1935b: 138 (in part). Phibalosoma, Westwood, 1859: 71 (in part). Stål, 1875: 63 (in part). Wood-Mason, 1877: 161. Phobaeticus Brunner v. Wattenwyl, 1907: 183 (in part). Brock, 1996: 28 (in part). Otte & Brock, 2005: 268 (in part). Tirachoidea, Kirby, 1904a: 359 (in part—only species No. 1a). [Not: Pharnacia, Chen & He, 2007: 349, figs. 316-317 = Tirachoidea Brunner v. Wattenwyl, 1893]

Description (%, &): Moderately robust to very massive Pharnaciini (&& in particular) with strong sexual dimorphism. Head at least 1.5x longer than wide, oval with the vertex flat or gently rounded, unarmed. Antennae of && shorter than mesonotum, in %% as long as or longer than head and thorax combined; antennomeres elongate. Pronotum considerably shorter and narrower than head. Mesothorax moderately elongate, 1.7–2.4x longer than head and pronotum combined, in && ± decidedly constricted anteriorly. Mesonotum smooth or more rarely set with a few minute granules. Mesosternum of %% occasionally with a ± distinct longitudinal median carina; simple in &&. Alae of %% well developed, at least reaching to posterior margin of abdominal tergite V. Tegmina ovate or scale-like, at best 2x longer than wide. Median segment of && at least 2/3 the


HENNEMANN & CONLE

length of metanotum, usually of ± equal length but never longer; conspicuously longer than metanotum in %%. Lateral margins of abdominal tergite VII dilated in posterior portion, forming a ± prominent, rounded or truncate lateral lobe. No obvious praeopercular organ on sternum VII of &&. Tergite VIII of && shorter than XI and X combined. Semi-tergites of anal segment of %% at best slightly longer than tergite IX, rather broad and often ± triangular with the apex broadly rounded, roundly truncate or triangular. Cerci short, oval to almost round in cross-section, occasionally distinctly compressed laterally and sickle-shaped. Subgenital plate of && strongly keeled, boat-shaped and at best slightly projecting over apex of anal segment; never lanceolate. Poculum of %% strongly convex and cup-like with posterior margin ± truncate; centre rounded and without a distinct spine or spine-like process at the angle. Legs of moderate length, femora of && ± broadened towards the base and broader than corresponding tibiae. All carinae ± strongly dentate, serrate and/or armed with single conspicuously enlarged spines or lobe-like teeth. Anterodorsal carina of profemora ± serrate. Mesofemora distinctly longer than metanotum and median segment combined. Medioventral carina of meso- and metafemora distinct and ± prominently spinose (&& in particular). Probasitarsus of && at best as long as remaining tarsomeres combined usually shorter; of %% as long as or slightly longer than remaining tarsomeres. Meso- and metabasitarsus about as long as following three tarsomeres combined with all carinae minutely serrate; dorsal carina not conspicuously raised. Eggs (Figs. 145–156): Large. Capsule globose slightly longer than wide, sub-cylindrical in cross-section, with flat impressions and surrounded by a very faint dorsoventral bulge. Capsule surface rough, punctated or granulated, not shiny. Polar-area smooth or with a slight dorso-polar depression. External micropylar plate covering more than 2/3 of the capsule length, shaped like a bold inverted Y. Apices of posterior arms of plate broadened and ± truncate. Operculum slightly convex, oval to almost circular and with an irregularly shaped hat- or knob-like capitulum on a distinct stalk. Diagnosis (Table 2): Very closely related to Tirachoidea Brunner v. Wattenwyl, 1893 but easily distinguished by the following characters: vertex of both sexes flat or at best slightly convex and unarmed; tergite VIII of && shorter than IX and X combined; dorsal carina of meso- and metabasitarsus of %% at best slightly raised but never prominently rounded and serrate. Furthermore, the eggs are distinguished by: the sub-cylindrical cross-section of the capsule, which exhibits a very faint dorsoventral bulge and more or less distinct depressions near the operculum and polar-area; rough and punctated not shiny capsule surface; broadened apices of the posterior arms of the micropylar plate; lack of a blunt central spine or tubercle at the polar-area and the plain colouration of the capsule which lacks a conspicuous dark lateroventral marking. Certain species may at first glance show striking similarity to some of the more massive taxa of Phobaeticus Brunner v. Wattenwyl, 1907. From this genus it is however easily distinguished by: the lack of a praeopercular organ of &&; shorter antennae of && which are shorter than the mesonotum; much shorter, broader and triangular lobes of the anal segment and less elongate, obtuse cerci of %%. Furthermore, the subgenital plate of && is never strikingly elongated or lanceolate as in certain species of Phobaeticus and does never project considerably over the anal segment, and the median segment of && is at least 2/3 the length of the metanotum (as short as half the length of metanotum in certain && of Phobaeticus). The eggs differ obviously from those of Phobaeticus by being globose, not prominently keeled or lens-shaped and having the micropylar plate shaped like an inverted Y. Comments: Stål (1877a: 40) originally established Pharnacia for his new Philippine species Pharnacia ponderosa Stål and Phibalosoma hypharpax Westwood, 1859 from Sri Lanka. Kirby (1904a: 359) selected Stål’s species as the type of Pharnacia Stål, 1877. Brunner v. Wattenwyl (1893: 83) split the genus and transferred Phibalosoma hypharpax Westwood, 1859 and three further species (Phibalosoma cantori Westwood, 1859, Phibalosoma tirachus Westwood, 1859 and Cyphocrania tamyris Westwood, 1859) to his new genus Tirachoidea. The latter species, C. tamyris Westwood, is not closely related and has subsequently been selected as the type species of Diesbachia Redtenbacher, 1908, a genus of the subfamily Necrosciinae (Brock, 1995). From the remaining Kirby (1904a: 359) designated Phibalosoma cantori Westwood as the type species



95

of Tirachoidea Brunner v. Wattenwyl, 1893. Redtenbacher (1908: 449) synonymised Tirachoidea Brunner v. Wattenwyl with Pharnacia Stål and listed 16 species in the genus. The distinguishing key for Pharnacia Stål however listed the type species P. ponderosa Stål as “speciae incertae”, most certainly because Redtenbacher had not examined Stål’s type specimens. Karny (1923: 240) recognized Pharnacia sensu Redtenbacher to be polyphyletic and re-established Tirachoidea Brunner v. Wattenwyl, transferring the first four more massive species in Redtenbacher’s key, which have a distinctly swollen or conical head, to Tirachoidea Brunner v. Wattenwyl. This included the type species of Tirachoidea, T. cantori (Westwood, 1859). Brock (1996: 25) recognized the erroneous placement of Phobaeticus Brunner v. Wattenwyl, 1907 and transferred it from Baculini Bradley & Galil, 1977 (= Clitumnini) to Pharnaciini. The author proposed a new arrangement of the genera Pharnacia, Tirachoidea and Phobaeticus, and in contrast to Karny (1923 : 240) confirmed Redtenbacher’s synonymy of Pharnacia Stål and Tirachoidea Brunner v. Wattenwyl , synonymizing Tirachoidea yet again. Brock (1996: 25) stated “Tirachoidea to be a synonym of Pharnacia on the basis that T. cantori belongs to the same genus as Pharnacia ponderosa Stål, the type species of Pharnacia.” and mainly based the synonymy on %% of both species having fully developed alae. Furthermore, his key states Pharnacia ponderosa Stål to show the same strongly rounded or elevated head seen in Ph. cantori (Westwood) “Back of head rounded or elevated; often with two swollen tubercles. (1996: 26)”, which is not the case; it is actually just gently rounded and entirely smooth in Ph. ponderosa Stål (Fig. 290). Thus, the strongly convex or conical and bi-tuberculate head (Figs. 296–304) is not a typical character of Pharnacia Stål, but one of the key features that distinguish Tirachoidea Brunner v. Wattenwyl from Pharnacia Stål. Consequently, all those taxa placed in Pharnacia by Brock (1996), which have a distinctly convex and bi-tuberculate head like the type species T. cantori (Westwood), actually belong in Tirachoidea Brunner v. Wattenwyl. Pharnacia Stål has the vertex smooth, completely destitute of spines or tubercles and at best gently rounded. Apart from the shape of the head, there are several other obvious characters of the insects and eggs which clearly distinguish Pharnacia and Tirachoidea (→ Table 1 & comments on Tirachoidea). As a result, Kirby’s and Karny’s treatment of the genera is here confirmed and Tirachoidea Brunner v. Wattenwyl re-established as a valid genus (stat. rev.). Based on egg-morphology and features of the genitalia two species are here removed from Phobaeticus Brunner v. Wattenwyl, 1907 and re-transferred to Pharnacia: Pharnacia heros Redtenbacher, 1908 and Phibalosoma tirachus Westwood, 1859 (stat. rev.). Distribution (Fig. 83): Complete Sundaland (Peninsular Malaysia, Sumatra, Java, Borneo & Palawan) and Philippine Islands. Species included: 1. Pharnacia borneensis spec. nov. [Borneo] 2. Pharnacia heros Redtenbacher, 1908: 453. [Java] 3. Pharnacia kalag Zompro, 2005a: 73, figs. 3–6. [Philippines (Mindanao Id.)] 4. Pharnacia palawanica spec. nov. [Palawan] 5. Pharnacia ponderosa Stål, 1877a: 40. [Philippines] = Pharnacia magdiwang Lit & Eusebio, 2008: 117, figs, 2c, 2f, 3a, 4, 5. syn. nov. 6. Phobaeticus sumatranus Brunner v. Wattenwyl , 1907: 184. [Sumatra, Peninsular Malaysia & Java] = Pharnacia rigida Redtenbacher, 1908: 453. syn. nov. 7. Phibalosoma tirachus Westwood, 1859: 75, pl. 37: 3. [Peninsular Malaysia & Java] = Clitumnus irregularis Brunner v. Wattenwyl , 1907: 195. syn. nov.


HENNEMANN & CONLE

FIGURES 89-93: Pharnacia spp. 89. Pharnacia ponderosa Stål, 1877, & Philippines, Panay Id., Mount Nangtud (coll. FH, No. 0192-11) 90. Pharnacia ponderosa Stål, 1877, % Philippines, Panay Id., Mount Nangtud (coll. FH, No. 0192-7) 91. Pharnacia palawanica spec. nov., & PT, SE-Palawan, Brook’s Point (coll. FH, No. 0455-1) 92. Pharnacia sumatrana (Brunner v. Wattenwyl , 1907), & West Malaysia, Tapah Hills (coll. FH, No. 0042-2) 93. Pharnacia sumatrana (Brunner v. Wattenwyl , 1907), % West Malaysia, Tapah Hills (coll. FH, No. 0042-4)



97

Keys to the species of Pharnacia Stål, 1877

&& 1. Cerci laterally compressed, up-curving and sickle-shaped...........................................................................2 - Cerci cylindrical or oval in cross-section, not up-curving or sickle-shaped.................................................3 2. Large species (body length 189.5–225.0 mm); meso- and metasternum with bold red or purplish spots (Fig. 420); legs armed with prominent spines (Figs. 315 & 319); Peninsular Malaysia, Sumatra & Java .... ...................................................................................................................................................... sumatrana - Smaller species (body length 180.0–184.3 mm); no red spots on meso- and metasternum; legs minutely dentate; Borneo............................................................................................................ borneensis spec. nov. 3. Anterodorsal carina of protibiae without conspicuous armature; mesofemora without distinctly enlarged teeth or lobes; Philippines & Palawan ..........................................................................................................6 - Anterodorsal carina of protibiae distinctly serrate and/or with single enlarged teeth or lobes; mesofemora with distinctly enlarged teeth or lobes ..........................................................................................................4 4. Anterodorsal carina of protibiae not raised but with 1–4 conspicuously enlarged lobes .............................5 - Anterodorsal carina of protibiae distinctly raised; without conspicuously enlarged lobes; Java ..........heros 5. Head oval, < 2x longer than wide; anterodorsal carina of protibiae serrate and at best with one enlarged lobe; Philippines ....................................................................................................................................kalag - Head elongate, almost 2.5x longer than wide (Fig. 294); anterodorsal carina of protibiae with 3–4 prominent + foliaceous lobes; Peninsular Malaysia & Java ...................................................................... tirachus 6. Very robust species (body width 14.0–18.0 mm); tergites IV–VII < 2x longer than wide; tergite VII with + prominent posterolateral lobes; Philippines...................................................................................ponderosa - Slender species (body width 9.0–10.0 mm); tergites IV–VII > 2x longer than wide; posterolateral lobes of tergite VII very indistinct; Palawan ............................................................................palawanica spec. nov.

%%* 1. 2. 3.

Semi-tergites of anal segment ± triangular, and tapered apically .................................................................2 Semi-tergites of anal segment broad and rounded apically (Fig. 196); Philippines ......................ponderosa Head globose, ± 1.5x longer than wide vertex convex.................................................................................3 Head elongate, > 2x longer than wide, vertex flat (Fig. 295); Peninsular Malaysia & Java ............ tirachus Robust insects, tergites II–VI < 5x longer than wide; alae at least reaching half way along abdominal tergite VI .............................................................................................................................................................4 - Slender insects, tergites II–VI > 5x longer than wide; alae shorter, at best reaching to posterior of abdominal segment V ...............................................................................................................................................5 4. Small (body length 103.8–110.0 mm); cerci elongate, apically pointed and slightly projecting over apex of anal segment (Fig. 199); abdominal tergite VII with a distinct, rounded posterolateral lobe; Borneo .......... ..................................................................................................................................... borneensis spec. nov. - Large (body length 125.5–149.0 mm); cerci short, obtuse and not reaching apex of anal segment (Fig. 202); posterolateral angles of abdominal tergite VII at best gently rounded; Peninsular Malaysia, Sumatra & Java ........................................................................................................................................... sumatrana 5. Alae reaching about half way along abdominal segment V; semi-tergites of anal segment with apex short and roughly triangular; vertex with blackish markings (Fig. 426); Philippines ....................................kalag - Alae reaching to posterior of abdominal segment V; apex of semi-tergites of anal segment strongly tapered with the tip gently elevated (Fig. 204); vertex without conspicuous dark markings; Java.......heros * %% of Ph. palawanica spec. nov. are not known.


HENNEMANN & CONLE

Pharnacia borneensis spec. nov. (Figs. 94–95, 153–154, 197–199, 352, 371) HT, %: N-Borneo, Mt. Kinabalu, Mesilau Camp., 1.–5.IV.1964, 5000 ft., Royal Soc. Exped., Coll. S. Kueh, BM 1964250 (BMNH). PT, &: Brunei, Badas, swamp forest, leg. M. Herbert 1995 (coll. MH). PT, &: Borneo Exped. Dr. Nieuwenhuis, Mahakam Riv, 1894, Pharnacia rigida K. Günther det. (RMNH). PT, %: Sabah, just outside Kinabalu Park HQ, found dead on ground, killed by domestic cat, P.E. Bragg 26-8-2001 (coll. PEB, No. 3152). PT, %: Sabah, just outside Kinabalu Park HQ area, P.E. Bragg, 11.XII.2006 (coll. PEB, No. 3550). PT, %: Sabah, Kinabalu N. P. 26.IX.–15.XI.83 A. Harman, 5,000 ft. (coll. AJH). PT, %: Sabah (N. Borneo), Kudasang, 24. May 1984, Coll. C. L. Chan & J. Cocking (coll. CLC). PT, %: Kinabalu N.P., May 1986, C.L. Chan (coll. CLC). PT, %: Sabah, Kinabalu Park, Sabah E. Malaysia, 18.10.1993, leg. Patrick Gindal (KNP, No. SP-EPH-00017). PT, %: P.H.Q., Kinabalu, 9.11.2001, Coll. Sinail. D. (KNP, No. SP-EPH-00018). PT, %: Kinabalu Park HQ, Alt 1650 m, 08/May/2006, Coll: Sinail Dunsul (KNP, No. SP-EPH-01288). PT, 10 eggs: Borneo, Brunei, Badas, swamp forest, leg. M. Herbert 1995 (coll. FH, No’s 0402-E & 0402-MP). PT, 2 eggs: Borneo, Brunei, Badas, swamp forest, leg. M. Herbert 1995, ex coll. F. Hennemann (coll. PEB). Pharnacia rigida, Günther, 1943: 155. [Misidentification] Baculolonga rigida, Bragg, 2001: 385, map 24B. [Relating to misidentified specimen of Günther, 1943: 155] Further material [2 &&]: BORNEO (SABAH): 1 &: Sabah, Tenom, Ulu Kallang, May 2004, Mornin B., Richard L.A. & Willibrord J. (FRCS); 1 &: Sabah, Bandukan Park, Keningau, 20.07.1993, coll. Matth. Allai (FRCS).

Diagnosis: Closely related to Ph. sumatrana (Brunner v. Wattenwyl, 1907) from Peninsular Malaysia, Sumatra and Java and the type species Ph. ponderosa Stål, 1877 endemic to the Philippines. Apart from being endemic in Borneo it is easily distinguished from Ph. sumatrana by: the much smaller size of both sexes; distinctly less prominent armature of the legs and lack of red or purplish sports on the meso- and metasternum of &&; as well as the slightly more elongate and apically less strongly constricted, semi-tergites of the anal segment and much longer and more slender cerci of %%. The eggs differ by the smaller dimensions, more slender micropylar plate and having the polar-area flattened. From Ph. ponderosa is can be distinguished by: the smaller size and more slender body and more distinct triangular dorsal lobe of the meso- and metatibiae of of both sexes (may sometimes be present in ponderosa); the smaller lateral lobes of tergite VII; much longer cerci, and more elongate, apically tapered, roughly triangular semi-tergites of the anal segment of %%. The eggs are smaller and have the polar-area not conspicuously indented and the micropylar plate differently shaped with the anterior apex rounded instead of elevated and truncate as in Ph. ponderosa. Etymology: The name refers to the distribution of this species. It is endemic in Borneo and the only representative of Pharnacia Stål, 1877 known from the island so far. Description: && (Fig. 92): Moderately sized (body length 180.0–184.3 mm) and rather robust species (maximum body width 10.0–11.0 mm). General colouration of body and legs dull green with irregular brown mottling, or pale to dark yellowish or greyish brown with lighter brown to pale greyish mottling and speckles; often with shallow whitish portions on the head, pronotum, mesonotum and abdomen. The & PT in RMNH is dark greyish brown and has a broad whitish longitudinal median stripe running along the complete dorsal surface of the head and body. Eyes greyish brown. Antennae mid brown. Head: Oval, 1.7x longer than wide, oval in cross-section, vertex gently convex, posterior margin slightly pointed medially. A fine transverse impressed line between the bases of the antennae. Eyes circular and moderately convex; their length contained almost 4x in that of cheek. Antennae almost reaching to middle of



99

mesonotum, segments increasing in length towards apices. Scapus dorsoventrally flattened, parallel-sided, almost 3x longer than wide. Pedicellus cylindrical, about 1/3 the length of scapus.

FIGURES 94-97: Pharnacia spp. 94. Pharnacia borneensis spec. nov., 1908, & PT, Brunei, Badas (coll. MH) 95. Pharnacia borneensis spec. nov., % PT, Sabah, Mount Kinabalu (coll. PEB, No. 3550) 96. Pharnacia tirachus (Westwood, 1859), & West Malaysia, Perak (ZMUH) 97. Pharnacia tirachus (Westwood, 1859), % Java, Mount Gede, Tjibodas (MNHU)


HENNEMANN & CONLE

Thorax: Pronotum rectangular, almost parallel-sided and about 1.4x longer than wide, distinctly shorter and narrower than head. Median transverse depression distinct and {-shaped, median line strongly impressed in anterior half of segment. Mesothorax of moderate length, about 2x longer than of head and pronotum combined. Mesonotum parallel-sided and very slightly constricted anteriorly. Metanotum parallel-sided, 2x longer than wide, less than half the length of mesonotum. Abdomen: Median segment slightly more than 2/3 the length of metanotum, 1.3x longer than wide and parallel-sided. Segments II–VI increasing in length, II indistinctly longer than wide, III 1.5x, IV 2x longer than wide, all parallel-sided. Tergite VII slightly narrower and shorter than previous, Tergite VII ¾ the length of VI and in posterior half expanded into a narrow, very widely triangular lobe, which laterally extends by less than ¼ the width of the segment. Sternites II–VII smooth except for a fine longitudinal carina along lateral margins. Tergite VIII equal in length to previous but distinctly narrower, constricted medially, convex and 2.5x longer than wide. IX less than half the length of VIII, strongly convex and roughly quadrate. Anal segment as long as IX and with a faint median carina; posterior margin with a broad, [-shaped median excavation, posterolateral angles short and slightly truncate (Fig. 198). Supraanal plate short, roundly truncate, keeled and just not reaching apex of anal segment. Cerci small, laterally compressed, gently up-curving and slightly projecting over apex of anal segment. Gonapophyses elongate, slender, distinctly up-curving and projecting over anal segment. Subgenital plate strongly keeled, boat-shaped and projecting over apex of abdomen by about half the length of anal segment (Fig. 197). Legs: All moderately long and of moderate strength for the genus, profemora a little longer than mesothorax, metafemora slightly projecting over posterior margin of abdominal segment IV and hind tarsi just not reaching tip of abdomen. All carinae minutely but densely serrate (more sparingly on dorsal carinae of midand hind legs). Meso and metafemora very slightly broadened basally. Anterodorsal carina of profemora with 12–15 acute triangular serrations, posteroventral carina with a similar number of rather pointed teeth. Medioventral carinae of meso- and metafemora with a row of 7–9 small spines. Posterodorsal carina of meso- and metatibiae with a distinct triangular lobe about 1/3 off the base and apically forming a slightly rounded lobe. All carinae of basitarsi minutely serrate, dorsal carina slightly raised (very gently rounded on meso- and metabasitarsus). Probasitarsus as long as remaining tarsomeres combined except claw. Meso- and metabasitarsus as long as combined length of following three tarsomeres. %% (Fig. 93): Rather small (body length 103.8–110.0 mm) for the genus, moderately slender and with very long alae (62.0–67.4 mm). General colouration of body and legs variable, ranging from yellowish over greenish to reddish mid brown. Tegmina and costal region of alae mid brown and sometimes greenish, the latter with irregular whitish markings. Head with irregular darker brown to black markings on the cheeks and five more or less parallel lines of small black spots on the vertex. Meso- and metasternum mid to dark green. Metapleurae green with posterior portion brown. Sternites of abdomen with irregular white markings and speckles. Anterior margin of tegmina and alae with a yellowish white longitudinal line, which only runs about 1/3 the way along the alae. Anal region of alae transparent grey with darker veins. All femora with indistinct greyish mottling. Antennae pale to mid brown and becoming darker brown towards the apex. Head: Generally as in &&, 1.6x longer than wide but eyes more prominent, strongly convex and projecting hemispherically; their length contained about 2.5x in that of cheek. Antennae consisting of 30 segments and reaching at least half way along abdominal tergite III. Segments distinctly increasing in length towards apices. Scapus und pedicellus as in &&. Third antennomere about 2.5x longer than pedicellus. All antennomeres finely bristled. Thorax: Pronotum as in &&, 1.5x longer than wide, distinctly narrower and shorter than head. Mesonotum very slightly broadened at posterior margin. Mesonotum and metapleurae irregularly set with a few small granules. Metanotum very short. Mesosternum with a very faint longitudinal median carina which terminates about half the weavy along segment. Tegmina oval, constricted basally, almost 2x longer than wide and with a blunt conical central hump. Alae ± reaching posterior margin of abdominal tergite VI.



101

Abdomen: Median segment 2x longer than metanotum. Segments II–VI ± 4x longer than wide and very slightly constricted medially. Tergite VII ¾ the length of VI and posteriorly dilated into a rounded lobe, which laterally extends by almost half the width of the segment. VIII slightly shorter than previous, strongly swollen and widening towards posterior; posterior margin almost 2x broader than anterior margin. IX distinctly narrower than previous, 2x longer than wide, constricted medially. Anal segment about 1.2x longer than IX. Semi-tergites rather slender, triangular and gradually tapered towards an acute apex (Fig. 199). Cerci very elongate, about 2/3 the length of anal segment and slightly projecting over apex of abdomen; cylindrical, slightly up-curving and laterally compressed at the apex. Poculum strongly convex, cup-like, with a conical hump at the angle and reaching about 2/3 the way along tergite IX. Legs: Generally as in && but relatively longer, more slender and with the spination more decided. Profemora about as long as head, pro- and mesonotum combined, mesofemora longer than mesonotum, metafemora reaching about 1/3 the way along abdominal tergite V and hind tarsi projecting over apex of abdomen. Mesoand metatibiae often lacking the triangular median lobe on posterodorsal carina. Eggs (Figs. 153–154, 352): The following description is based on a number of eggs which were laid by the & PT from Brunei (Badas) in coll. MH. Unfortunately, the capitulum is missing in all examples at hand. Medium-sized, capsule slightly laterally compressed and oval in cross-section, longer than high. Seen laterally the polar-area is slightly truncate. Capsule surrounded by a faint dorsoventral keel, beginning and ending at operculum. General colouration of capsule and micropylar plate greyish mid brown, operculum blackish. Capsule surface densely but very minutely granulated. Micropylar plate gently raised with the outer margin darker brown. General shape like a bold inverted Y; all three extension rounded at apices. Median line short, not extending over posterior of micropylar plate. Micropylar cup small, rounded and placed in centre of the posteromedian gap of the plate. Operculum flat, oval and with a white cone-shaped, capitular stalk in the centre. Measurements [mm]: length 3.9–4.1, width 3.2–3.3, height 3.7–3.8, length of micropylar plate 2.8–3.0. TABLE 6. Measurements [mm] of Pharnacia borneensis spec. nov.

%, HT (BMNH)

%, PT (coll. AJH)

&&, PT

Body

103.8

110.0

180.0–184.3

Head

4.5

5.3

12.0–12.2

Pronotum

3.2

3.8

7.0–7.1

Mesonotum

15.0

17.1

35.2–37.0

Metanotum

-

6.8

14.9–16.0

Median segment

-

11.5

8.8–10.0

Tegmina

8.8

10.1

-

Alae

63.0

67.4

-

Profemora

28.9

29.5

39.0–39.5

Mesofemora

20.8

21.4

29.0–31.5

Metafemora

29.8

29.8

36.0–39.6

Protibiae

32.5

34.2

41.0–41.8

Mesotibiae

20.4

20.9

29.5–30.5

Metatibiae

28.8

28.9

40.2–41.2

Antennae

> 39.0

52.0–54.0

31.5


(RMNH & MH)

HENNEMANN & CONLE

Comments: Examination of the & from Mahakam, Borneo in RMNH, which was erroneously identified as Pharnacia rigida Redtenbacher by Günther (1943: 155), has proven this to represent a specimen of Ph. borneensis spec. nov., here designated a PT. Ph. rigida Redtenbacher, 1908, a synonym of Ph. sumatrana (Brunner v. Wattenwyl , 1907 (syn. nov.)., is restricted to Sumatra, Peninsular Malaysia and Java and although closely related easily distinguished from the Bornean Ph. borneensis spec. nov. by numerous morphological features (see diagnosis above). Already Bragg (2001: 385) doubted this Bornean specimen really was Ph. rigida and provided a brief characterization and measurements. During a one-day visit at Mulu National Park (Sarawak) in August 1994 the first author observed a dark green & on the hand rail of the main plank-walk, that leads from the park headquarters to “Deer Cave”. As seen from the recorded localities Ph. borneensis spec. nov. has a wide distributional range in the northern half of Borneo, which comprises Sabah, Sarawak, Brunei and the northern portions of Kalimantan, but appears to be rather uncommon. Distribution (Fig. 371): Northern and Central Borneo, endemic. Sabah (Mt. Kinabalu, Mesilau 5000 ft. & Kinabalu Park Headquarters 1500 m), Brunei (Badas, swamp forest), Sarawak (Mulu National Park, near Dear Cave & Kuching) and Kalimantan (Mahakam River). Number of specimens examined: 13

Pharnacia heros Redtenbacher, 1908 stat. rev. (Figs. 98–99, 203–204, 314, 372) Pharnacia heros Redtenbacher, 1908: 453. LT [by present designation], &: Java orient. Montes Tengger 4000´ 1890 H. Fruhstorfer; Java H. Fruhstorfer vend. 10.II.1894: 3, Tirachoidea heros Redt. & Typus. Jos. Redtenbacher determ. 1899, public. 1906-08; Bestimm.-Verz. Nr. 3 „T. Hypharpax Ww.“ (ZMUH); PLT, &: Java occident. Sukabumi, leg. Fruhstorfer 2000´ 1893; H. Fruhstorfer vend. 15.1.1895; 4; Tirachoidea heros var. Redt. & (Typus) Jos. Redtenbacher determ. 1899, public. 1906-08, Bestimm.-Verz. Nr. 4 (ZMUH). [Not: PLT, &: Sumatra, Indragiri, leg. Mechel 2. 1895 (NHMB). This is the PLT of Pharnacia sumatrana (Brunner v. Wattenwyl, 1907); see comments below] Weidner, 1966: 231. Zompro, 2002a: 189. Phobaeticus heros, Brock, 1996: 28. Otte & Brock, 2005: 268. ? Pharnacia nigricornis, Caudell, 1927: 19. [Misidentification] Further material [1 &, 1 %]: JAVA: 1 %: Volcan Gede, Java, 2. 1893, Collection A. Finot, Phryganistria acanthopus de Haan (MNHN, coll. Finot, BoxNo. 280); 1 &: Volcan Gede, Java, Collection A. Finot, Phryganistria acanthopus de Haan (MNHN, coll. Finot, Box-No. 280).

Diagnosis: Close to Ph. tirachus (Westwood, 1859) but distinguished by: the slightly more robust body and legs; relatively shorter body segments; shorter, more globose and convex head; densely serrate anterodorsal carina of the protibiae; lacking considerably enlarged lobes on the same carina, and less prominent armature of the mid and hind-legs of both sexes. Furthermore, && differ by the more slender lateral lobes of tergite VII and %% by the broader, apically constricted and rounded semi-tergites of the anal segment. Etymology: “Heros” (gr. = hero). A noun in apposition to the generic name. Description: && (Fig. 98): Small to medium-sized (body length 164.0–180.0 mm) and rather slender for the genus (maximum body width 9.0–10.0 mm). General colouration of body and legs dark greyish brown and irregularly set with numerous pale greyish or yellowish markings and speckles. Meso- and metafemora each with a bold dark sub-basal band Eyes reddish brown. Antennae dark brown.



103

Head: Oval, 1.5x longer than wide, vertex gently convex. Between the bases of the antennae with a small, curved impression. Eyes small, convex and slightly truncate at anterior margin; their length contained a little more than 3x in that of cheek. Antennae reaching about 2/3 the way along the mesonotum, antennomeres increasing in length towards apices of antennae. Scapus dorsoventrally flattened, parallel-sided and about 2.3x longer than wide. Pedicellus less than half the length of scapus, cylindrical. Thorax: Pronotum 2/3 the length and narrower than head, with posterior margin broader than anterior margin, 1.2x longer than wide. Anterior margin with a slightly raised transverse carina, median transverse depression indistinct, gently curved and reaching to lateral margins of segment. Mesothorax very slightly constricted at the anterior, about 2.5x longer than head and pronotum combined. Mesonotum slightly widened posteriorly and with a fine, longitudinal median carina. Mesosternum smooth. Metanotum 2/5 the length of mesonotum, 2x longer than wide, parallel-sided, anterior or posterior margins slightly raised. Abdomen: Median segment slightly shorter than metanotum, about 2x longer than wide and rectangular. Segment II slightly shorter than median segment, 1.5x longer than wide. III–VI slightly increasing in length, III 2.3x, VI 2.7x longer than wide. Tergites II–VIII each with a small, rounded posteromedian tubercle. VII shorter than previous, 2x longer than wide and posteriorly dilated into a broad, rounded lobe which laterally projects by about half the body width and posteriorly slightly projects the margin of the tergite. VIII shorter and narrower than VII, strongly convex and slightly constricted medially. IX as wide as VIII, but considerably shorter and just indistinctly longer than wide. Anal segment slightly longer than IX and with a fine median carina. Posterior margin with a shallow, roundly triangular median excavation, outer angles tapered and rounded at the apex. Supraanal plate small, rounded and with a distinct dorsal keel. Cerci small, oval in crosssection and tapered towards the apex. Gonapophyses elongate, up-curving and slightly projecting over apex of abdomen. Subgenital plate strongly keeled, boat-shaped and reaching to apex of anal segment (Fig. 203). Legs: All rather long and moderately slender. Profemora almost as long as pro-, and mesothorax combined, mesofemora slightly shorter than mesothorax, metafemora projecting over posterior margin of abdominal tergite IV and hind tarsi slightly projecting over apex of abdomen. All carinae except ventral carinae of protibiae minutely serrate or dentate. Anterodorsal carina of profemora with 20–24 acutely triangular serrations, posteroventral carina with a similar number of considerably smaller teeth. Anterodorsal carina of protibiae elevated and distinctly and densely serrate (Fig. 314). Mesofemora with a broad triangular, dentate sub-basal lobe on posteroventral carina and a single enlarged tooth about ¼ off the base on posterodorsal carina. The latter carina protruded into a distinct, black bi-, or tri-dentate apical lobe. Posterodorsal carina of meso- and metatibiae with a rounded lobe about 1/3 of the base on posterodorsal carina (less decided on metatibiae) and a rounded, dentate lobe at the apex. Basitarsi with all carinae slightly elevated; slightly longer than following three tarsomeres combined. %% (Fig. 99): Medium-sized (body length 112.3 mm) and very slender for the genus, with long, slender legs but comparatively short alae (50.0 mm). General colouration of body and legs mid brown, legs with indistinct pale greyish transverse bands. Posterior portion of anal segment very dark brown, cerci pale brown. Abdominal sternites II–VII with irregular white speckles. Mesonotum and mesosternum green, metasternum yellowish with a few elongate white spots. Tegmina mid brown with a large black marking on the central hump and costal region of alae slightly greenish mid brown; both with anterior margin white. Anal region of alae transparent grey. Armature of legs black. Antennae mid brown. Head: Generally as in && but vertex more convex and eyes relatively larger and projecting hemispherically; their length contained a little more than 2x in that of cheek. Antennae reaching to posterior margin of tergite II, otherwise as in &&. Thorax: Pronotum as in &&. Mesothorax almost 3x longer than head and pronotum combined, mesonotum parallel-sided and smooth. Mesosternum with a faint longitudinal median carina, metasternum smooth. Tegmina oval, strongly constricted towards the base and with a prominent, rounded and slightly forward-pointing central hump. Alae reaching to posterior margin of abdominal tergite V.


HENNEMANN & CONLE

FIGURES 98-102: Pharnacia / Phobaeticus spp. 98. Pharnacia heros Redtenbacher, 1908, & PLT, Java, Sukabumi (ZMUH) 99. Pharnacia heros Redtenbacher, 1908, % Java, Mount Gede (MNHN, coll. Finot) 100. Pharnacia kalag Zompro, 2005, & HT, Philippines, Mindanao Id., Lantapan (coll. OZ) 101. Phobaeticus ingens (Redtenbacher, 1908), & LT, SW-India, Malabar Coast, Meppadi (MHNG) 102. Phobaeticus ingens (Redtenbacher, 1908), % N-Myanmar, Highland of Tenasserim (coll. FH, No. 0251-2)

Abdomen: Median segment distinctly longer than metanotum, smooth. Segments II–VI slightly increasing in length II 5x, VI about 6.5x longer than wide, all gently constricted medially. Tergites V–VII with a minute posteromedian tubercle. VII a little shorter than VI, slightly widening towards the posterior, in posterior por-



105

tion with a small and narrow, rounded lateral lobe. VIII ¾ the length of VII, strongly swollen, convex, widened towards the posterior and 1.5x broader than previous segments. IX slightly longer than VIII but distinctly narrower and decidedly constricted medially; about 2.5x longer than wide. Anal segment about as long as IX. Semi-tergites with posterior portion strongly constricted and the apex roundly truncate (Fig. 204). Interior surfaces minutely dentate. Cerci small and rather short, cylindrical and tapered towards a pointed apex, not reaching tip of anal segment. Poculum reaching half way along tergite IX, strongly convex, cup-like and angled. Legs: All very long and slender. Profemora longer than head, pro-, and mesonotum combined, mesofemora slightly exceeding posterior margin of abdominal segment II, metafemora reaching to apex of abdominal segment V and metatibiae projecting considerably over apex of abdomen. Anterodorsal carina of profemora with 15–17 triangular serrations of average size, posteroventral carina with a similar number of small teeth. Anterodorsal carina of protibiae very minutely but densely dentate. Probasitarsus almost as long as remaining tarsomeres combined, with a smooth but slightly elevated dorsal carina. All carinae of mid and hind legs set with minute teeth, more sparingly on dorsal carinae although. Posteroventral carina of mesofemora with a shallow dilation basally and three slightly more prominent, triangular teeth. Medioventral carina of meso- and metafemora gently raised and set with a few very minute spines. Posterodorsal carina of meso- and metatibiae with a distinct and rounded, dentate apical lobe. Meso- and metabasitarsus slightly longer than remaining tarsomeres combined and with a very slightly elevated dorsal carina. Comments: Redtenbacher (1908: 453) originally described Pharnacia heros from three syntypic &&, two in ZMUH and one in NHMB. The & from the Tengger Mountains in ZMUH, which bears the green label “No. 3”, is here selected as the LT. Presumably Redtenbacher erroneously referred to the same Sumatran specimen in NHMB which had already been described as Phobaeticus sumatranus by Brunner v. Wattenwyl (1907: 184). Therefore, “Sumatra” is here regarded an erroneous record due to based on a different species. Weidner (1966: 231) erroneously listed the & from Tengger-Gebirge as HT and the one from Sukabumi as PT. The collection of M. Finot in MNHN contains a nice & and % from Mount Gedeh in Western Java, which were erroneously identified as “Phryganistria acanthopus de Haan”. This % in MNHN is the only one known of this species and served for the description above. Eggs unknown. Distribution (Fig. 372): Java (Tengger Mountains 4000 ft.; Mount Gede & Sukabumi 2000 ft.). Number of specimens examined: 4 TABLE 7. Measurements [mm] of Pharnacia heros Redtenbacher, 1908 stat. rev.

&&,

&

%

LT & PLT (ZMUH)

( MNHN)

( MNHN)

Body

164.0–178.5

180.0

112.3

Head

-

9.4

4.7

Pronotum

6.0–6.5

5.8

3.2

Mesonotum

34.5–38.0

38.5

22.5

Metanotum

15.0–16.8

16.3

-

Median segment

10.5–11.7

11.5

-

Tegmina

-

-

8.4

Alae

-

-

50.0

Profemora

39.5–42.5

44.7

35.0

Mesofemora

30.5–31.0

33.2

27.7

Metafemora

38.0–40.0

40.0

34.8

Antennae

35.0

28.5

55.5


HENNEMANN & CONLE

Pharnacia kalag Zompro, 2005 (Figs. 100, 147–148, 208, 320–321, 374, 425–426) Pharnacia kalag Zompro, 2005a: 73, figs. 3–6. HT, &: Mindanao Id, Bukidnon Prov., Katanglad Mts, Lantapan, 1000– 1300 m, IV.1991, leg. A. Buenafe (coll. OZ); PT, &: same data as holotype (ANIC); PT, &: same data as holotype (BMNH); PT, &: same data as holotype (MCSN); PT, &: same data as holotype (MHNG); PT, &: same data as holotype (MNHN); PT, &: same data as holotype (NHMW); PT, &: same data as holotype (ZMUH); PT, 5 &&, 1 egg: same data as holotype (UPPC); PT, 8 &&, 1 egg: same data as holotype (coll. OZ). Otte & Brock, 2005: 265. Lit & Eusebio, 2008: 117, figs. 2b, e. Further material: [3 &&, 1 male, eggs]: PHILIPPINES (Mindanao Id.): 1 &: Philippines, Mindanao, Bukidnon Province, 1000–1300 m, Katanglad Mountains, Lantapan, IV.1991, leg. A. Buenafe, ex coll. OZ (coll. FH, No. 0449-1); 18 eggs: Philippines, Mindanao Id., South Cotabato Prov., Tampakan 1300 m, leg. B. Mabanta III.2008 (coll. FH, No. 0449-E); 2 &&: Philippines: Malaybalay, Bukidnon, II.1987, BMNH(E) 200598, Phobaeticus sp. det. P. Brock 2005 (BMNH). 1 %: Tampakan field trails, South Cotabato, Mindanao, Philippines; ex culture J. Bresseel 2007)

Diagnosis: Closely related to P. tirachus (Westwood, 1859) from Java and Peninsular Malaysia, and Ph. heros Redtenbacher, 1908 from Java. From the it is distinguished by: the more robust body; more oval and less elongate head, and smaller but more acute dorsal serrations of the profemora of both sexes, as well as having at best one ± distinctly enlarged lobe on the anterodorsal carina of the protibiae (3–4 in tirachus); distinctly dentate ventral carinae of the mesofemora and differently shaped lateral lobes of abdominal tergite VII in && and a relatively longer anal segment of %%. The eggs differ considerably in being less globose and having the micropylar plate almost triangular, with the posterolateral extensions very short. From Ph. heros && may be distinguished by: the longer and more elongate body; relatively longer body segments; broader lateral lobe of abdominal tergite VII; not densely serrate dorsal carina of the protibiae and different armature of the mid and hind legs (Figs. 320–321). %% are very similar ot those of Ph. heros but differ by: the less globose and conspicuously marked vertex (Fig. 426); slightly shorter alae; less distinct apical lobe of the meso- and metatibiae and more acutely triangular apex of the semi-tergites of the anal segment. Etymology: “Kalag” is a Cebuano word meaning soul, ghost or spirit (Zompro, 2005a: 74). Description: && (Fig. 100): Long to very long (body length 196.0–228.0 mm) and slender (maximum body width 8.0–9.0 mm) for the genus. General colouration of body and legs mid to dark brown with irregular whitish speckles and markings. Meso- and metafemora usually with a ± decided black and white sub-basal band, and corresponding tibiae often with a faint, irregular pale transverse band pre-medially. Eyes dark greyish or reddish brown. Two basal segments and apical half of antennae dark brown, remaining portion reddish brown Head: Oval, almost 1.8x longer than wide, vertex slightly convex. Between the bases of the antennae with a small, slightly curved depression. Eyes small, circular and moderately convex; their length contained almost 4x in that of cheek. Antennae reaching about 1/3 the way along mesonotum, with about 27 segments. Scapus dorsoventrally flattened, parallel-sided and about 2.5x longer than wide. Pedicellus one third the length of scapus, round in cross-section. Thorax: Pronotum narrower and about 2/3 the length of head, about 1.3x longer than wide; anterior margin narrower than posterior margin. Median transverse depression short, slightly curved and not reaching lateral margins of segment. Mesothorax moderately constricted anteriorly and about 2.4x longer than head and pronotum combined. Mesonotum parallel-sided and with a faint longitudinal median line. Metanotum slightly less than half the length of mesonotum, 3x longer than wide, parallel-sided. Meso- and metasternum smooth. Abdomen: Median segment slightly shorter than metanotum but longer than tergite II, about 2.5x longer



107

than wide and parallel-sided. Segments II–VI increasing in length, II 2x, V and VI almost 4x longer than wide; all parallel-sided. Tergite V with a scale-like posteromedian tubercle. VII ¾ the length of VI and posterolaterally dilated into an roundly triangular lobe which projects by about 1/3 of the body width. Tergite VIII narrower than previous segments, about ¾ the length of VII, strongly convex and slightly constricted medially. IX parallel-sided, indistinctly longer than wide, 2/3 the length of VIII. Anal segment longer than IX with a fine median carina. Posterior margin with a small semi-circular median indention, posterolateral angles rounded. Supraanal plate small, transverse and with a fine median keel. Cerci small sub-oval in cross-section, tapered towards the apex. Gonapophyses elongate, up-curving and slightly projecting over anal segment. Subgenital plate distinctly keeled, boat-shaped and reaching to posterior margin of anal segment (Fig. 208). Legs (Figs. 320–321): All moderately long and slender, profemora slightly longer than mesothorax, mesofemora a little shorter than mesothorax, metafemora reaching half way along abdominal segment V and hind tarsi not or just reaching apex of abdomen. Anterodorsal carina of profemora with 15–20 prominent and acute serrations; posteroventral carina more minutely and sparingly serrate. Anterodorsal carina of protibiae densely serrate and occasionally with an enlarged, roughly triangular median lobe; ventral carinae minutely serrate. Meso- and metafemora with one, two or three ± prominent triangular teeth or lobes on the anterodorsal carina, otherwise sparingly dentate. Two outer ventral carinae roughly but sparsely dentate. Medioventral carina armed with a variable number ± prominent spines, which decrease in size towards the apex. Posterodorsal carina of meso- and metatibiae with a prominent, roundly triangular lobe just before the middle and a rounded, dentate lobe apically. Probasitarsus slightly shorter than remaining tarsomeres combined, dorsal carina slightly raised and all carinae minutely serrate. Meso-, and metabasitarsi as long as following three tarsomeres combined, with all carinae serrate and dorsal carina slightly raised and rounded. %% (Figs. 425, 426): The colouration is described from photos of a live % kindly provided by R. Krijns (Maastricht, Netherlands) two of which are here reproduced. Medium sized (body length 132.0 mm) and very slender for the genus, with long alae. Most of body and legs greenish mid brown, the terminal abdominal segments darker. Head ochracheous with a washed greenish marking between they eyes and irregular blackish markings on the vertex and cheeks. Eyes pale cream and irregularly marbled with dark brown. Mesothorax and metasternum dull green. Mesopleurae with a few whitish spots, metapleurae dark greyish. Tegmina and alae pale creamish brown, the anterior margin of the tegmina and basal 1/3 of alae broadly white. Tegmina with a small blackish spot on top of the central hump. Anal region of alae transparent pale brown with brown veins. Cerci pale cream. Apices of meso- and metafemora and bases of corresponding tibiae pale brown. Complete armature of legs black. Tarsi pale brown, the mesoand metabasitarsi whitish basally. Antennae mid brown, scapus ochracheous. Head: Oval, about 1.5x longer than wide, narrowing from eyes towards posterior margin, vertex flat. Between the bases of antennae with a slight transverse impression, followd by a slightly raised oval area between the eyes. Eyes very prominent, strongly convex and projecting hemispherically from head capsule; their length only about 1.6x in that of cheeks. Antennae projecting considerably over median segment. Scapus dorsoventrally flattened, parallel-sided and 2.5x longer than wide. Pedicellus 1/3 the length of scapus, cylindrical. Following antennomeres first short but then distinctly increasing in length. Thorax: Pronotum distinctly shorter and narrower than head, the anterior portion slightly narrowed. Median transverse depression faint, short and not reaching lateral margins of segment. Mesothorax about 2.6x longer than head and pronotum combined, mesonotum with a few minute granules along the lateral margins. Mesosternum with a very faint longitudinal median carina, metasternum smooth. Tegmina elongate, oval and with a moderately distinct, rounded central hump. Alae ± reaching about half way along abdominal tergite V. Abdomen: Segments II–VI roughly of equal length and about 5x longer than wide. VII ¾ the length of VI and only about 3.5x longer than wide. Tergite V with a minute posteromedian tubercle. Sternites II–VII smooth. Tergite VIII a little less than 3/4 the length of VII, 3x longer than wide, and gradually widening towards the posterior. IX about as long but considerably narrower than VIII, strongly convex, constricted


HENNEMANN & CONLE

medially and about 3x longer than wide. Anal segment a little shorter than IX, strongly keeled and laterally compressed. Semi-tergites broad with the apex roughly triangular. Interior surfaces apically set with small black teeth. Cerci cylindrical, slightly tapered towards a narrow tip and reaching to apex of anal segment. Poculum strongly convex, roundly cup-like and reaching about half way along tergite IX, the central portion slightly elevated. Legs: All very long and slender, profemora considerably longer than head, pro- and mesonotum combined, mesofemora a little longer than pro- and mesonotum combined, metafemora reaching to posterior margin of abdominal segment V and metatibiae extending well beyond apex of abdomen. Anterodorsal carina of profemora with about 16 distinct and acutely triangular black serrations, which increase in size towards apex of femur. Posteroventral carina with a about 20 very minute teeth. Anterodorsal carina of protibiae minutely but quite densely serrate, posteroventral carina very sparsely serrate, anteroventral carina smooth. Two outer ventral carinae of meso- and metafemora sparsely dentate, dorsal carinae each with only 3–7 irregularly set minute teeth. Medioventral carina indistinct and with a few very minute spines. All carinae of meso- and metatibiae minutely serrate, the dorsal carinae more sparsely although. Probasitarsus a little longer than following tarsomeres combined, with all carinae moderately elevated and smooth. Meso- and metabasitarsi about equal in length to combined length of remaining tarsomeres, all carinae gently elevated, the dorsal smooth and the ventral with a few very minute teeth. Variation: The type series (all &&) shows some variation concerning to the size of the two enlarged teeth of the anterodorsal carina of the mesofemora. These may be indistinct and just slightly larger than the remaining teeth of that carina (Fig. 321), or prominently enlarged to form two roughly triangular lobes (Fig. 321). One of the two && from Malaybalay in BMNH is remarkable for the strikingly developed leg furniture, having the meso- and metafemora each supplied with three large foliaceous lobes and the corresponding tibiae with the triangular median lobe on the posterodorsal carina very large (all comparatively smaller on hind legs). The second & widely corresponds to the type series, which shows considerable variation to occur within a single colony. Eggs (Figs. 147–148): Medium-sized, capsule slightly laterally compressed, oval in cross-section, longer than high. In lateral aspect the polar-area has a shallow ventro-polar depression. Complete capsule surrounded by a very faint dorsoventral bulg, beginning and ending at the operculum. General colouration of capsule and operculum plain dark brown, capitulum black. Complete capsule surface very minutely granulated and leather-like; slightly glabrous. Micropylar plate slightly raised and with a darker brown outer margin. Generally shaped like a very bold inverted “Y”, the posterolateral extensions very short and roughly triangular. Median line almost extending to polar-area. Micropylar cup small and placed in centre of posteromedian indention of the micropylar plate. Operculum flat and oval. In its centre with an large, roughly cone-shaped capitulum which covers almost the complete operculum. Measurements [mm]: length (including capitulum) 4.5–4.6, length 3.8–4.0, width 2.7–2.8, height 3.5–3.7, length of micropylar plate 3.0. Comments: There are several inaccuracies in the original description provided by Zompro (2005a) which in particular concerns to the measurements given for the type series. Some years before publication of Ph. kalag the entire type series, including the HT and eggs, were kindly given to the first author on loan for examination. As indicated in Table 8 below measuring of these specimens provided body lengths of 196.0–228.0 mm in contrast to 168.0–178.0 given by Zompro. Unfortunately, the HT was not measured separately but the 175.0 mm given by Zompro are very doubtful, since it is 21 mm shorter than the shortest body length taken from the type series by the first author. A & in coll. FH (No. 0449-1) originating from the same locality and source was kindly presented to the first author by O. Zompro but not designated a PT. It measures 218.0 mm, hence exceeding the maximum length recorded by Zompro by as much as 40 mm. Furthermore, Zompro described and illustrated the micropylar plate of the eggs as being heart-shaped. In fact however it is triangular with the posterolateral angles slightly elongated and triangular, forming the inverted “Y” typical for Pharna-



109

cia Stål (Fig. 147). Culture stock originating from Tampakan in the South Cotabato Province (Mindanao Id.), has recently been imported to Europe and is currently being attempted to rear on oaks (Quercus spp., Fagaceae) and bramble (Rubus fruticosus, Rosaceae). Distribution (Fig. 374): Southern Philippines, Mindanao Island (Bukidnon Province: Katanglad Mountains, Lantapan 1000–1300 m & Malaybalay; South Cotabato Province: Tampakan). Number of specimens examined: 25 TABLE 8. Measurements [mm] of Pharnacia kalag Zompro, 2005.

&, HT (coll. OZ)*

&&, PT*

Body

175.0

168.0-178.0

Head

8.3

Pronotum

&

%

(coll. FH)

(coll. JB)

196.0-228.0

218.0

132.2

7.3-8.8

12.8-14.6

13.5

6.2

5.4

5.1-6.9

8.0-9.3

8.0

4.4

Mesonotum

34.5

32.4-39.4

39.5-46.8

46.0

27.5

Metanotum

8.9

8.5-9.9

14.7-18.9

18.1

4.0

Median segment

7.6

6.8-8.9

11.5-14.2

14.0

13.1

-

-

-

-

9.6

Tegmina Alae

&&, PT**

-

-

-

-

55.6

Profemora

40.0

38.1-48.6

44.0-52.9

46.4

48.9

Mesofemora

29.5

28.9-36.8

32.5-42.1

37.2

33.5

Metafemora

35.5

33.7-42.0

40.8-52.0

46.0

45.1

Protibiae

45.2

43.5-55.2

-

57.2

43.8

Mesotibiae

31.3

30.5-39.6

-

38.2

32.2

Metatibiae

39.1

38.5-47.8

-

51.0

39.5

-

-

> 30.0

82.5

Metatibiae * according to Zompro (2005a: 74). ** measurements taken by the first author

Pharnacia palawanica spec. nov. (Figs. 91, 149–150, 209, 375) HT, &: Philippines, Palawan, Brooks Point, 7.10.1974, R. Rodriguez, BM 1975-193 (BMNH). PT, & (subadult nymph): Philippines, Palawan, Brooks Point, 7.10.1974, R. Rodriguez, BM 1975-193 (BMNH). PT, 2 &&: Philippines, SE-Palawan, Brook’s Point, 1998 (coll. FH, No’s 0455-1 & 2). PT, 1 egg: Philippines, SE-Palawan, Brook’s Point, 1998, ex ovipositor FH 0455-1 (coll. FH, No. 0455-E). PT, &: Philippines, Palawan, IV.1985, leg. C. Finetti (coll. MG). PT, female: Philippinen, Nord Palawan, 20.XII.1997 (coll. OC).

Diagnosis: Closely related to the type species Ph. ponderosa Stål, 1877 but at once distinguished by: the more slender body and legs; relatively longer body segments; much smaller posterolateral lobes of abdominal tergite VII; shape of the anal segment; less punctured surface of the egg capsule; shape of the micropylar plate and smaller dimensions of the eggs. Etymology: The name refers to the type locality of this new species, which is the so far only known representative of Pharnacia Stål, 1877 in Palawan. Description: && (Fig. 91): Long (body length 191.5–195.0 mm) and moderately slender for the genus (maximum width 9.0–10.0 mm). General colouration of body and femora plain pale to dark brown (abdomen


HENNEMANN & CONLE

darkened due to preservation in all specimens at hand), tibiae and carinae of femora greenish mid brown. Mesonotum with a broad longitudinal band of irregular black speckles along lateral margins. Vertex with a slightly darker brown coronal-line and four fine sub-parallel longitudinal lines dorso-laterally. Antennae dark brown with the two basal segments slightly paler brown and the ventral surfaces black. Eyes reddish brown. Head: Large and oval, 1.6x longer than wide, vertex gently convex. Between the bases of the antennae with a small sickle-shaped, transverse impression. Eyes of moderate size, conspicuously convex and with the anterior margin slightly angled; length contained about 3x in that of cheek. Antennae at least reaching half the way along mesonotum (broken in all specimens at hand), antennomeres increasing in length towards apices of antennae. Scapus dorsoventrally flattened, very slightly outcurving, 2.5x longer than wide. Pedicellus less than half the length of scapus, 1.5x longer than wide, cylindrical. Third antennomere very indistinctly longer than pedicellus. Thorax: Pronotum, 1.4x longer than wide, considerably shorter and narrower than head, lateral margins gently concave and posterior margin a little broader than anterior margin. Anterior margin slightly raised, median transverse depression short, {-shaped and not reaching lateral margins of segment. Mesonotum very slightly widened towards the posterior, about 2.2x longer than head and pronotum combined. Metanotum parallel-sided, a little more than 2x longer than wide, less than half the length of mesonotum. Meso- and metasternum smooth. Abdomen: Median segment a little more than 2/3 the length of metanotum, 1.5x longer than wide, parallel-sided. Segments II–III 2x, IV–VII 2.5x longer than wide. Tergite VII posterolaterally with a small, roughly triangular tooth. VIII narrower than previous, strongly convex, parallel-sided, 2.5x longer than wide. IX half the length of VIII, indistinctly longer than wide, strongly convex. Anal segment slightly longer than IX, with a very faint, slightly impressed median line. Posterior margin with a broad concave median excavation, the outer angles rounded. Supraanal plate small, rounded and not extending to apex of anal segment. Cerci small, elongate, round in cross-section and slightly tapered towards the apex. Gonapophyses elongate, slender, upcurving and just reaching to posterior margin of anal segment. Subgenital plate distinctly keeled, boat shaped and slightly projecting over apex of abdomen (Fig. 209). Legs: All rather long and slender with all carinae very minutely dentate (more sparingly on dorsal carinae). Profemora slightly longer than pro-, and mesonotum combined, mesofemora shorter than mesonotum, metafemora reaching 1/3 the way along abdominal segment V and hind tarsi slightly projecting over tip of abdomen. Anterodorsal carina of profemora with 16–18 well decided serrations. Medioventral carina of mesoand metafemora indistinct and set with 7–10 minute spines. Basitarsi as long as following three tarsomeres combined, all carinae very minutely serrate. Eggs (Figs. 149–150): Medium-sized, capsule slightly laterally compressed, oval in cross-section, longer than high. Seen laterally the polar-area has a distinct dorso-polar depression. Capsule surrounded by an indistinct blunt dorsoventral bulg, beginning and ending at operculum. General colouration of capsule and operculum creamish mid brown, capitulum black, micropylar plate greyish brown. Complete capsule surface very minutely granulated and leather-like. Micropylar plate gently raised from capsule and with a broad, dark brown outer margin. Shaped like a bold inverted “Y”, with all three extensions of moderate length. Anterior extension rounded, the two posterior extensions slightly truncate apically. Median line projecting distinctly over posterior margin of plate. Micropylar cup small, cup-like and placed in centre of the posteromedian gap of the plate. Operculum flat and oval with a large, irregularly cone-shaped capitulum in the centre. Measurements [mm]: length including capitulum 5.1, length 4.4, width 3.1, height 4.0, length of micropylar plate 2.7. Comments: The & penultimate instar nymph (PT) in BMNH measures a body length of 140.0 mm. %% unknown. Distribution (Fig. 375): Palawan (Brooks Point). Number of specimens examined: 6



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TABLE 9. Measurements [mm] of Pharnacia palawanica spec. nov.

&, HT (BMNH)

&&, PT

Body

195.0

191.5–194.0

Head

13.0

12.6–12.9

Pronotum

7.3

7.2

Mesonotum

39.5

36.0–37.5

Metanotum

17.5

16.0–16.5

Median segment

11.0

9.5–10.0

Profemora

50.0

47.5–49.5

Mesofemora

35.5

35.0–37.5

Metafemora

43.5

44.0–45.0

Protibiae

55.0

53.5–55.5

Mesotibiae

31.2

35.0–35.5

Metatibiae

48.0

45.3–46.2

Antennae

41.0

> 30.0

(coll. FH)

Pharnacia ponderosa Stål, 1877 (Figs. 89–90, 145–146, 194–196, 269, 290–291, 318, 376, 427, 436–438) Pharnacia ponderosa Stål, 1877a: 40. LT, %: Ins. Phil., Semper; Pharnacia ponderosa Stål [darker brown specimen] (NHRS); PLT, %: Ins. Phil., Semper; Pharnacia ponderosa Stål (NHRS). Kirby, 1904a: 359. Redtenbacher, 1908: 455. Sjöstedt, 1933: 6. Brock, 1996: 27. Brock, 1999: 181. [Designation of LT] Bresseel, 2004: 15, figs. 3–5. Otte & Brock, 2005: 265. Lit & Eusebio, 2008: 120, figs. 2a, d, 3a & table 1. Pharnacia magdiwang Lit & Eusebio, 2008: 117, figs, 2c, 2f, 3a, 4, 5. HT, &: ex Mangifera indica, Sibuyan Island, Brangay Ipil, Magdiwang, 31.X.2006, S.R. Rabo (UPPC, No. PHA-00130); AT, %: reared from egg of holotype, died 15.IV.2007, UPLB MNH Central Lab. (UPPC, No. PHA-00132); PT, %, &: same data as AT (UPPC, No’s PHA-00131 & 00133); PT, 10 eggs: laid by HT (UPPC). syn. nov. [Not: Pharnacia ponderosa, Kirby, 1896: 451; Misidentification, = Phasmotaenia sanchezi (Bolívar, 1897)] Further material: [45 &&, 14 %%, 2 nymphs, eggs]: PHILIPPINES: 1 &, 3 %%, 2 && (nymphs), 4 eggs: Philippinen, Marinduque Id., ex. Zucht F. Hennemann, 1992 (coll. FH, No’s 0192-1 to 6 & 0192-E1); 3 &&, 1 egg: Philippinen, Mindoro Island, Mt. Halcon, leg. Mohagan 2.IV.–5.V.1996 (coll. FH, No’s 0192-27 to 29 & 0192-E3); 1 &: Philippinen, N-Luzon Id., Mountain Province, Nueva Viscaya, Balite, leg. I.O. Lumawig VII.1996 (coll. FH, No. 0192-30); 15 &&, 4 %%: Philippinen, Panay Island, Mt. Nangtud, 1500 m, leg. Mohagan VII. 1997 (coll. FH, No’s 0192-7 to 25 & 0192-E2); 1 &: without data (coll. FH, No. 0192-26); 1 %, 1 &: ex Zucht; R. Galunder (Nümbrecht) III.2008, Philippinen (coll. FH, No’s 0192-31 & 32); 1 %, 1 &: ex Zucht. F. Hennemann 2008, Herkunft: Philippinen (coll. FH, No’s 0192-33 & 34); 5 &&: Philippinen, Panay Island, Mt. Nangtud, 1500 m, leg. Mohagan VII.1997 (coll. OC); 1 &: Philippinen, Mindoro, Mt. Halcon, leg. Mohagan 15.–26.V.1996 (coll. OC); 1 &: Philippines, Luzon, from UP Los Baños, without data, X. 1995 (coll. OZ, No. 0084-2); 1 &: Philippines, Sibuyan Id., Romblon Prov., Magdiwang, Pawala River, Tampayan, Camp New St. Gallen area, 20.–30.VII.1986, 100–400 m, Roland


HENNEMANN & CONLE

A. Müller leg. (coll. OZ, No. 0084-3); 1 & (nymph): Philippines, Cebu Id., Minglanilla, Camp 7, 300 m, 21./22.V.1991, Roland A. Müller leg., Coll. R. A. Müller (coll. OZ, No. 0084-4); 1 %: Zucht O. Zompro, Philippines, Marinduque Id. (coll. OZ, No. 0084-1); 1 %: Philippinen, Aroroy (MHNG); 1 &: Mt. Makiling, Luzon, Baker (ANSP); 1 &: Mt. Isarog, Tiaong, camarines, Sur, Elev. 2000 feet, May 20, 1924, M.A. Mariano, P.C.A. (ANSP); 1 &, 1 %: Island Sibuyan, Baker (ANSP); 1 %: N.W. Panay, 20773 (ANSP); 1 &: Los Baños, P.I., August 1928 J.R. Arena, P.C.A. (ANSP); 2 &&: Philippines: Sierra Madre, Quezon, E. Luzon, VI.1999, I.O. Lumawig; BMNH(E) 2005-98 (BMNH); 1 &: Philippines: Panay Is., 12.IV.1998; BMNH(E) 2005-98 (BMNH); 1 &: Philippines: Panay Is., IV.1998, BMNH(E) 2005-98 (BMNH); 2 &&: Philippines: Mountain Prov., N. Luzon, VII.96, I. Lumawig, BMNH(E) 2005-98 (BMNH); 1 %, 1 &: Philippines, Sorsogon, S. Luzon, VIII.1998, I. Lumawig; BMNH(E) 2005-98 (BMNH); 1 &, 9 eggs: Philippines, Marinduque Island, Reared by H. Probst, March 1990; BMNH(E) 2005-98 (BMNH); 1 %: Canarines S. Libuanan, Maranedo (MNCN). NO DATA: 1 &: 94.138 (BMNH).

Diagnosis: Closely related to Ph. palawanica spec. nov. from Palawan and Ph. borneensis spec. nov. from Borneo. From the first && differ by: the considerably broader and more robust body and legs; relatively shorter body segments; much broader lateral lobes of abdominal tergite VII; different shape of the anal segment and longer cerci. The eggs differ by the larger dimensions and different shape of the micropylar plate, which has the three extensions relatively longer and more slender. From the Bornean Ph. borneensis spec. nov. it differs by: the larger size; more robust body and broader lateral lobes of abdominal tergite VII of both sexes; deeper posteromedian excavation of the anal segment and more slender cerci of &&; considerably shorter and smaller cerci and much broader, posteriorly broadly rounded semi-tergites of the anal segment of %%. The eggs differ by the larger dimensions, more decidedly indented polar-area and relatively larger micropylar plate, which has the three extension longer and the anterior one slightly elevated and truncate at the apex. Etymology: From the Latin “ponderosus” (= weighty) and referring to the robust body of this species. Description: The colouration is described from preserved wild specimens and live captive reared specimens. && (Fig. 89): Large (body length 170.0–205.0 mm), very robust and massive species of cigar-like appearance (maximum body width 14.0–18.0 mm). General colour of body and legs variable, ranging from straw over pale to dark brown, more rarely mid to dull green. Colouration either plain, or body more or less decidedly mottled with pale greyish, yellowish or whitish speckles and legs to a variable degree furnished with annulations and markings of the same colour. Head usually with six dark, ± decided, sub-parallel longitudinal lines. Abdominal tergite III sometimes paler than remaining tergites and with distinct pale greenish or whitish mottling. Spination of the legs dark greyish green, the spines of the medioventral carina of the meso- and metafemora ochre. Antennae mid to dark brown with ventral surface very dark brown to black. Eyes dark greyish or reddish brown. Head (Fig. 290): Prominent, oval, 1.6x longer than wide with vertex slightly convex. Vertex with a ± distinctly impressed coronal-line. Between the bases of antennae with a small but distinct curved impression. Eyes of moderate size, higher than long, gently convex with anterior margin slightly angled; length contained almost 5x on that of cheek. Antennae reaching almost 2/3 the way along the mesonotum, with 30 segments. Antennomeres slightly increasing in length with the apical 5–7 segments conspicuously shortened. Scapus dorsoventrally flattened, slightly oval in dorsal aspect and about 2.3x longer than wide. Pedicellus less than half the length of scapus, slightly longer than wide, cylindrical. Third antennomere slightly longer than pedicellus. Thorax: Pronotum 2/3 the length and distinctly narrower than head, posterior margin broader than anterior margin, about 1.2x longer than wide. Anterior margin raised and anterior half of segment with a prominently impressed median line. Median transverse depression distinct but short and at best covering half the width of segment. Mesothorax less than 1.8x longer than head and pronotum combined, decidedly constricted anteriorly, very gently swollen pre-medially and slightly widened at posterior margin. Mesonotum with a very faint



113

longitudinal median line. Metanotum 2/5 the length of mesonotum, almost 2x longer than wide and parallelsided. Meso- and metasternum smooth. Abdomen: Median segment slightly shorter than metanotum, 1.5x longer than wide and roughly rectangular with the lateral margins gently concave. Segments II–VI indistinctly increasing in length, II about 1.3x, VI 1.5x longer than wide; III and IV broadest segments. Tergite VII slightly shorter than VI, the lateral margins gradually widened to form a ± distinct, rounded or roundly triangular lobe in posterior half, which may project by as much as half of the body width. VIII narrower than VII, 2x longer than wide, constricted medially and strongly convex; anterior margin slightly broader than posterior margin. IX a little more than half the length of VIII, roughly quadrate, strongly convex. Anal segment slightly longer than IX and with a fine longitudinal median carina. Posterior margin with a deep, almost semi-circular median excavation, the outer angles each forming a rounded lobe (Fig. 195). Supraanal plate rounded, keeled and not extending beyond posterior margin of anal segment. Cerci obtuse, moderately elongate, oval in cross-section, slightly tapered towards apex and reaching about to apex of anal segment. Gonapophyses elongate, slender, up-curving and variable in length; either staying slightly beyond or considerably projecting over anal segment. Subgenital plate strongly keeled, boat-shaped and slightly projecting over apex of anal segment (Fig. 194). Legs: All rather short and robust, the meso- and metafemora slightly swollen sub-basally. Profemora slightly longer than pro-, and mesothorax combined, mesofemora very slightly shorter than mesothorax, metafemora projecting over posterior margin of abdominal segment IV and hind tarsi at best slightly exceeding apex of abdomen. Anterodorsal carina of profemora with 17–23 distinct, pointed serrations; posteroventral carina with a similar number of considerably smaller teeth. All carinae of protibiae very indistinctly dentate. Meso- and metafemora with all carinae rather minutely dentate; teeth more numerous on ventral carinae (Fig. 318). Dorsal carinae of mesofemora sometimes with a slightly enlarged, triangular tooth about 1/3 off the base; occasionally there is a further much smaller, triangular tooth some 2/3 the way along posterodorsal carina and a small tooth 1/3 off the base anterodorsal carina. Metafemora usually without conspicuously enlarged teeth, but rarely with a slightly enlarged, triangular tooth 1/3 off the base on posterodorsal carina. Ventral surfaces of meso- and metafemora slightly convex and with a fine medioventral carina, set with 6–8 rather distinct and pointed spines; 2–4 largest. Ventral carinae of meso- and metatibia very densely but minutely serrate, more sparsely and minutely on dorsal carinae. Occasionally there is a slightly enlarged or even lobe-like, triangular tooth about 1/3 off the base on posterodorsal carina. Probasitarsus with a uniformly raised but dorsal carina, all carinae smooth and about as long as remaining tarsomeres combined. Meso- and metabasitarsi with all carinae minutely dentate, dorsal carina very slightly raised and as long as following three tarsomeres combined. %% (Fig. 90): Medium-sized to large (body length 111.0–134.5 mm) and robust species (maximum body width 4.5–5.5 mm) with long alae (56.0–67.5 mm). Colouration variable. Body, legs, tegmina and costal region of alae uniformly yellowish, greenish or greyish pale to dark brown; more rarely dull green. Head occasionally with a broad dark brown marking or postocular band. Meso- and metasternum often dull green with several white, elongate spots. Anterior margin of tegmina and alae with a bold white exterior and a more indistinct green interior longitudinal line. Tegmina occasionally with a short, white diagonal band, which begins at the posterior margin and terminates in the centre of tegmen. Anal region of alae transparent greyish brown with darker veins. Abdominal tergite IX with a ± decided longitudinal white patch close to lateral margins. Antennae dark brown with ventral surfaces darker to black. Eyes greyish or reddish mid to dark brown. Head (Fig. 291): Generally as in &&, but eyes more prominent, circular and projecting hemispherically; their length contained out 2.5x in that of cheek. Antennae with 30 segments and slightly projecting over posterior margin of abdominal tergite II. Antennomeres distinctly increasing in length with the apical 7–8 antennomeres conspicuously shortened. Scapus und pedicellus as in &&. Thorax: Pronotum as in &&, 1.3x longer than wide. Mesonotum 2.1x longer than head and pronotum combined, parallel-sided and very slightly broadened at posterior margin. Meso- and metasternum smooth. Teg-


HENNEMANN & CONLE

mina oval, almost 2x longer than wide, constricted basally and with a distinct, roundly conical central hump; projecting over posterior margin of metanotum. Alae reaching about half way along tergite VI. Abdomen: Median segment 2x longer than metanotum, 3x longer than wide and slightly narrowing towards the posterior. Segments II–IV slightly increasing, V–VII decreasing in length, IV longest segment and about 3.5x longer than wide. Tergite VII ¾ the length of VI and posterolaterally dilated into a distinct, rounded lobe, which laterally extends by as much as half the width of segment. Tergites VIII and IX prominently swollen. VIII broadest segment and conspicuously widening towards the posterior, posterior margin almost 2x wider than VI. IX about as long as VIII, slightly narrowing towards the posterior, gently constricted medially and strongly convex. Anal segment slightly shorter than IX. Semi-tergites very broad, 1.3x longer than high and rounded apically (Fig. 196). Interior surfaces densely covered with minute hooked spines. Cerci small, moderately elongate, cylindrical , gently in-curving and reaching to apex of anal segment. Poculum strongly convex, cup-like and with a blunt, conical hump at the angle; reaching about ¾ the way along tergite IX. Legs: Generally as in && but relatively longer, much more slender and with spination slightly more decided. Profemora longer and mesofemora as long as than combined length of head, pro-, and mesonotum, metafemora almost reaching posterior margin of abdominal tergite V and metatibiae reaching about to apex of abdomen. Ventral surfaces of meso- and metafemora not conspicuously convex, dorsal carinae without enlarged teeth. Meso- and metatibiae occasionally with an enlarged triangular tooth some 1/3 off the base on a posterodorsal carina. Tarsi generally as in && but more slender and segments relatively longer. Eggs (Figs. 145–146): Large, capsule slightly laterally compressed, oval in cross-section, longer than high. Seen laterally the polar-area has a ± distinct impression (depends on locality). Capsule surrounded by a blunt dorsoventral keel, which begins and ends at the operculum. General colouration of capsule dull blackish brown or grey, micropylar plate greyish brown, operculum black and capitulum dark reddish brown. Capsule surface minutely but densely granulated. Two broad oval impression near operculum anterodorsally- and ventrally and two anteroventral depressions near polar end of capsule. Micropylar plate gently raised from capsule and with a slightly darker brown outer margin. Generally shaped like a bold inverted “Y” with all three extensions rather long. Anterior extension slightly broadened and truncate at the apex, the two posterior extensions with apices rounded. Median line short, not extending over micropylar plate. Micropylar cup small, placed in centre of posteromedian gap of the plate. Operculum flat, oval and with a knob-like capitulum on a short stalk in its centre. Measurements [mm]: length including capitulum 5.6–5.9, length 4.9–5.1, width 3.1–3.3, height 4.0–4.1, leng of micropylar plate 3.3. Variation: In accordance to its wide distributional range in the Philippines Ph. ponderosa shows quite some deal of intraspecific variation. Features that underlie considerable variation are the size, width and shape of the lateral lobes of abdominal tergite VII, armature of the dorsal carinae of the mid and hind legs and colouration. Most of these features appear to vary from locality to locality. A 200.0 mm & from N-Luzon (Mountain Province) in coll. FH (No. 0192-30) has the lateral lobes of tergite VII very broad, truncate and elevated towards the posterior (Fig. 269) and exhibits two conspicuously enlarged teeth about one third the way along the antero- and posterodorsal carina of the mesofemora and a distinct triangular pre-medial dorsal lobe on the meso- and metatibiae. In contrast, enlarged teeth or lobes on the legs are completely lacking or just very poorly developed and the lateral lobes of abdominal tergite VII considerably smaller and rounded in specimens from Panay, Mindoro or Marinduque. These two features appear to be typical for specimens from Luzon only. Slight variation is also seen in the shape of the semi-tergites of the anal segment in %%, which have the apex very broad in specimens from Panay or Luzon (Fig. 196) but comparatively more narrow in specimens from Marinduque. && from Mindoro Island are comparatively smaller (173.0–182.0 mm, coll. FH) than && from Luzon, Marinduque, Panay or Sorsogon (184.0–204.0 mm, coll. FH). Captive reared %% from Marinduque are typical for having a distinct blackish postocular marking on the cheeks and a whitish, diagonal line



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along the posterior margin of the tegmina, both features lacking in %% from other localities. Wild && are mostly various shades of brown, more rarely green, and often prettily mottled with white or pale brown. Captive reared && that were raised on alternative foodplants in Europe, are exceptionally mid to dull green. Not only the insects but also the eggs show slight variation, those from the islands of Panay and Mindoro being somewhat smaller with the polar-area less distinctly impressed than eggs from Luzon or Marinduque. Furthermore, slight variation is observed in the shape of the micropylar plate. Comments: Stål (1877a: 40) originally described Pharnacia ponderosa from two %% with the unprecise location “Philippinen” in NHRS of which Brock (1999: 181) designated the darker brown specimen as the LT. The fact that both specimens have a triangular pre-medial dorsal lobe on the meso- and metafemora suggests both specimens are from Luzon (see comments on variation above). Already Redtenbacher (1908: 455) doubted that the & which Kirby (1896: 451) described to have 7.0 mm long tegmina was the corresponding & of Stål’s species. Günther (1933: 159) correctly stated this specimen to be a & of Phasmotaenia sanchezi (Bolívar, 1897), which was confirmed by examination. Although Ph. ponderosa has a wide distributional range in the Philippines and is quite frequently encountered on most of the larger islands of the archipelago (see distribution below), being rather abundant in some localities, the & and egg have so far remained formally undescribed. Pharnacia magdiwang Lit & Eusebio, 2008 was described upon both sexes and the eggs from Magdiwang in the Romblon Province, Sibuyan Island. Lit & Eusebio (2008: 117) distinguished this species from Ph. ponderosa Stål, 1877 mainly on the basis that both sexes lack an obvious pre-medial dorsal lobe on the meso- and metatibiae and slight differences of the egg-morphology. Based on examination of specimens from various Philippine islands the pre-medial dorsal lobe of the meso- and metatibiae is here shown to underlie considerable intraspecific variability (see comments on variation above) and to be frequently well developed only in specimens from Luzon, the only island from which Lit & Eusebio (2008) had specimens of Ph. ponderosa at hand for comparison with their Ph. magdiwang. As stated above, even the eggs of Ph. ponderosa show slight variation depending on the locality, thus the differences summarized by Lit & Eusebio (2008: 117) for the eggs of Ph. magdiwang lie well within the range of variation of Ph. ponderosa. The “distinct sculpturing” and “fontanelle-like depression” of the head in Ph. magdiwang mentioned as distinctive features upon Ph. ponderosa by Lit & Eusebio (2008: 117) are questionable. Measurements and length relations of the insects generally correspond to other specimens of Ph. ponderosa examined for the present study (→ Table 10). The fact that Sibuyan Island shows a very high degree of endemicity and is hence often recognized as a separate biogeographical subregion of the Philippines (see Heaney & Regalado, 1998), can also not support to the validity of Ph. magdiwang, since Ph. ponderosa is known from most of the larger islands that surround Sibuyan (e.g. Luzon, Marinduque, Mindoro, Panay and Cebu) and the genus Pharnacia Stål, 1877 itself is apart from the Philippines distributed throughout entire Sundaland. Consequently, as none of the distinguishing features mentioned by Lit & Eusebio, 2008: 117) for Ph. magdiwang Lit & Eusebio, 2008 holds true this species is obviously a junior synonym of Ph. ponderosa Stål, 1875 (syn. nov.). In the Philippines Lit & Eusebio, 2008: 121) stated Ph. ponderosa to feed on Canarium ovatum (Burseraceae), Mangifera indica (Anacardiaceae), Terminalia microcarpa (Combretaceae) an guava (Psidium guajava, Myrtaceae). In captivity in Europe it accepts oaks (Quercus robur, Q. petraea, Q. rubra & Q. ilex, Fagaceae), bramble (Rubus spp., Rosaceae), eucalyptus (Eucalyptus spp., Myrtaceae), Salal (Gaultheria shallon, Ericaceae) and rose (Rosa spp., Rosaceae) as alternative foodplants. Culturing in Europe has been attempted several times already from stock originating from various localities but none was maintained for more than a few generations. A current culture-stock from an unknown locality however is rather easy to rear in large and well ventilated cages. Eggs have an incubation time of 4–5 months and nymphs reach maturity in about 6–8 months. A & produces an average of 8–10 eggs per day and a total of 400–700 during her lifetime. Distribution (Fig. 376): Philippines (endemic): Aroroy Id., Cebu Id. (Minglanilla 300 m), Luzon Id. (Mount Makiling; Sorsogon; Sierra Madre, Quezon; Los Baños; Camarines Sur Province, Mount Isarog: Tia-


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ong 2000 ft. & Mountain Province: Nueva Viscaya, Balite), Marinduque Id., Mindoro Id. (Mount Halcon), Panay Id. (Mount Nangtud), Sibuyan Id. (Romblon Province: Magdiwang, Pawala River, Tampayan 100–400 m & Brangay Ipil) & Libuanan Id. Number of specimens examined: 61 TABLE 10. Measurements [mm] of Pharnacia ponderosa Stål, 1875.

&&

%%

&, HT of

%, AT of

Ph. magdiwang (UPPC)*

Ph. magdiwang (UPPC)*

Body

173.0–205.0

111.0–134.5

170.00

125.30

Head

11.9–15.0

5.5–6.9

14.30

7.59

Pronotum

6.5–9.2

3.8–4.3

7.00

4.50

Mesonotum

33.0–38.5

20.5–24.5

34.20

23.57

Metanotum

14.8–18.0

6.5 –9.0

13.00

12.65

Median segment

9.8–11.0

13.5–15.0

10.75

7.75

Alae

-

56.0–67.5

-

59.50

Profemora

43.3–51.5

34.0–41.5

44.05

35.85

Mesofemora

32.0–36.8

25.2–29.0

32.00

25.69

Metafemora

39.5–42.8

33.0–38.5

38.00

35.13

Protibiae

49.5–56.0

37.0–48.5

47.45

41.56

Mesotibiae

33.5–35.0

24.5–27.5

30.75

28.67

Metatibiae

41.0–44.0

33.5–38.0

41.50

36.53

Antennae

42.0–45.0

50.5–67.0

-

-

* according to Lit & Eusebio (2008: 119, 120)

Pharnacia sumatrana (Brunner v. Wattenwyl , 1907) (Figs. 92–93, 151–152, 185, 200–202, 292–293, 315, 319, 353, 377, 415, 417, 419) Phobaeticus sumatranus Brunner v. Wattenwyl , 1907: 184. LT [by present designation], &: Musée de Genève No. 4; Lakat Sumatra, acheté J. M. Schneider (MHNG); PLT, &: Sumatra, Indragiri, leg. Mechel 2. 1895 (NHMB). Hennemann & Conle, 1999: 8. Pharnacia sumatranus, Brock, 1996: 27 Seow-Choen, 2000: 37, pl. 93 (%, &). [Descriptions and illustrations of % and &] Pharnacia sumatrana, Zompro & Brock, 2003: 24. Otte & Brock, 2005: 265. Pharnacia cantori, Seow-Choen, 1997: 52, fig. 25 (&). Brock, 1999: 132 (in part—only description of & and egg, figs. 24 (egg), 26 (&)). Lobophasma rex, Günther, 1935b: 139 (in part—only % from Sumatra). Pharnacia rigida Redtenbacher, 1908: 453. HT, &: Sumatra, Montes Battak, ex Coll. Fruhstorfer; Collectio Br. v. W.; det. Br. v. W.; 23.331; Pharnacia rigida Redt. Type! (NHMW, Nr. 862). syn. nov. Brock, 1997: 45. Brock, 1998a: 53. Otte & Brock, 2005: 265. Phobaeticus rigidus, Brock, 1996: 29. Seow-Choen, 1998a: 41. Pharnacia westwoodi, Giglio-Tos, 1910: 43. [Misidentification] Monandroptera gigliolii Westwood, in litt. 1 &: Java, „det. Monandroptera gigliolii Westwood in litt.“, Viaggio di Filippi (MRSN). [Not: Pharnacia rigida, Günther, 1943: 155. Misidentification, = Pharnacia borneensis sp. nov.] REVISION OF PHARNACIINI


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[Not: Baculolonga rigida, Bragg, 2001: 385. Misidentification, = Pharnacia borneensis sp. nov.] Further material: [14 &&, 4 %%, 1 & (nymph), eggs]: PENINSULAR MALAYSIA: 1 &, 1 & (penultimate instar nymph): W-Malaysia, Perak, Tapah Hills, 600 m, VIII.1994, via M.K.P. Yeh (coll. FH, No’s 0042-1 & 3); 1 &: W-Malaysia, Perak, Tapah Hills, ca. 600 m, IV.1994, via M.K.P. Yeh (coll. FH, No. 0042-2); 1 &, 1 egg: W-Malaysia, Perak, Tapah Hills, ca. 600 m, VII.1996, via Wong Tet Fatt (coll. FH, No’s 0042-5 & E1); 3 eggs: ex Zucht J.T.C. Sellick (England), Herkunt: West Malaysia, PSG No. 78 (coll. FH, No. 0042-E2); 1 %: W-Malaysia, Tapah Hills, ca. 600 m, 1993, via M.K.P. Yeh (coll. FH, No. 0042-4); 1 &: W-Malaysia, Tapah Hills, 200–600 m, via. M.K.P. Yeh, ex coll. FH, det. F. Hennemann X 1996 (ZMUH); 1 &: Cameron Highlands, 1981, C.L. Chan, BM 1985-463 (BMNH); 1 %, 1 &: W. Malaysia: Cameron Highlands, 2000ft., 19.VI.1972, C.C.Chua, BM 1975-604 (BMNH); 2 &&, eggs: Captive reared (BMNH); 2 &&: Cameron Highlands, Perak, Malaysia, 1200–1600 m, achat Pfanner 1970 (MHNG); 1 %: Malaisie, Perak, Cameron Highl., 1200–1600 m, leg. Pfanner (MHNG); 1 &: Cameron Highlands, Tapah Hills, Malaysia 1974 (ZSMC). SUMATRA: 1 %: E-Coast Sumatra, Doerian Moelan, Brindjei, Lt. R. Coughtrie (BMNH); 1 %: Sumatra, Redjang-Lebong, H. Kubale S. leg. (MNHU); 1 &: P.O. Stolz, Solok (Sum), 1914 (RMNH). JAVA: 1 &: Java, „det. Monandroptera gigliolii Westwood in litt.“, Viaggio di Filippi (MRSN); 1 &: Java, Phibalosoma acanthopus Burm.* (MLUH).

Diagnosis: && of this characteristic species is easily distinguished from all other members of the genus by: the very prominent and long ventral spines of the meso- and metafemora; the prominent, saw-like ventral spination of all three tibiae; obtuse, sickle-shaped cerci and large dark red to purplish spots on the meso- and metasternum. %% and eggs are similar to Ph. borneensis spec. nov. from Borneo and the type species Ph. ponderosa Stål, 1877 from the Philippines. %% differ from both species by the larger size and more elongate body; relatively longer body segments and lacking the conspicuous rounded lateral lobe of abdominal tergite VII. From the first they additionally differ by: the lack of a dorsal lobe on the meso- and metatibiae; shorter and broader cerci as well as the shorter and broader, apically less slender and acute semi-tergites of the anal segment. The eggs differ from Ph. borneensis spec. nov. by the larger dimensions, rounded polar-area and broader micropylar plate. From Ph. ponderosa %% may also be distinguished by: the more slender, roughly triangular, apically tapered and constricted semi-tergites of the anal segment and more obtuse cerci. Eggs differ from those of Ph. ponderosa in the shape of the micropylar plate, rounded polar-area and the lack of conspicuous dorso-polar impressions of the capsule. Etymology: Referring to the type locality of this species, the island of Sumatra. Description: The colouration is described from photos of live specimens. && (Fig. 92, 417, 419): Large to very large (body length 189.5–225.0 mm) and robust species (maximum body width 13.0–15.0 mm) with conspicuously broadened and heavily spinose legs. Colour variable, general colouration of body and legs either greyish to mid brown with darker brown speckles and occasionally with dark green mottling, or dark green with pale green and brownish speckles. Meso- and metanotum as well as pleurae with a broad longitudinal band of irregular black speckles along lateral margins. A similar but much more narrow longitudinal lateral stripe on abdominal tergites. An irregularly punctured, black V-shaped marking in posterior portion of mesonotum. Abdominal tergites II–III with a pair of short and curved, black lines just behind middle. Meso- and metasternum with several distinct, bold rounded dark red or purplish spots (Fig. 419). Abdominal sternites all over furnished with small black speckles. Antennae mid to dark reddish or greyish brown with the two basal antennomeres paler. Eyes reddish brown. Head (Fig. 292): Elongate, almost 2x longer than wide, oval in cross-section, vertex very slightly convex (Fig. 274). Posteriorly with a very faint darker coronal-line and four indistinct, slender and subparallel longitudinal lateral lines; the interior ones diverging towards the posterior and forming a bold “V”. Occasionally


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these lines may be impressed along the posterior margin of the head capsule. Between the bases of antennae with a small but decided, curved transverse impression, followd by a pair of slightly raised and smooth, oval areas. Eyes large, oval, higher than long and just slightly projecting from head capsule; their length contained about 3x in that of cheek. Antennae with 32 segments and reaching half way along mesonotum; antennomeres increasing in length towards the mid, then decreasing in length towards apices. Scapus dorsoventrally flattened, almost 2.5x longer than wide and tapered towards the base. Pedicellus half the length of scapus, 1.3x longer than wide, cylindrical. Third antennomere slightly longer than pedicellus. Thorax: Pronotum distinctly shorter and narrower than head, about 1.5x longer than wide and trapezoidal being gently narrowed towards the anterior. Anterior margin raised to form a fine transverse carina. Median transverse depression distinct, but very short and not reaching lateral margins of segment. Anterior portion with a distinctly impressed median line. Mesothorax decidedly constricted anteriorly and considerably broadened posteriorly, about 1.7x longer than head and pronotum combined. Mesonotum and mesosternum each with a very faint pale longitudinal median line. Mesopleurae with a distinct longitudinal carina and swollen near coxae. Metanotum parallel-sided, 1.5x longer than wide, less than half the length of mesonotum; longitudinal median line indistinct. Metapleurae distinctly widening and convex towards coxae. Abdomen: Median segment about 2/3 the length of metanotum, slightly longer than wide, gently constricted medially, anterior margin very slightly concave. Segments II–VI increasing in length, II 1.4x, IV 1.8x and almost VI 2.3x longer than wide. Tergite VII slightly shorter than VI, posterior half dilated into a rounded lateral lobe which extends by about ¼ of the body width. Sternites II–VII smooth. Tergite VIII as long as VII but distinctly narrower, strongly convex, 3x longer than wide and slightly constricted medially. IX strongly convex half the length of VIII, indistinctly longer than wide and gently widened towards the posterior. Anal segment as long as IX and with a distinct median keel; posterior margin with a deep truncate median excavation, the outer angles each forming a roundly triangular lobe (Fig. 197). Supraanal plate roundly truncate, strongly keeled but not projecting over apex of anal segment. Cerci of a conspicuously sickle-like appearance, strongly flattened laterally, with a distinct dorsal and ventral carina and a narrow up-curving apex (Fig. 196). Gonapophyses moderately elongate, slender, up-curving and slightly projecting over apex of anal segment. Subgenital plate strongly keeled, boat-shaped and projecting over posterior margin of anal segment by almost half of its length. Legs: All moderately long but very robust and heavily armed with long, pointed spines; meso- and metafemora prominently swollen towards the base with ventral carinae increasingly elevated. Profemora about as long as pro- and mesonotum combined, mesofemora very slightly longer than mesonotum, metafemora slightly projecting over posterior margin of abdominal segment IV and hind tarsi just projecting over apex of abdomen. Anterodorsal carina of profemora armed with 12–14 distinct, pointed serrations. Posteroventral carina with 8–10 long, pointed triangular spines and occasionally 2–5 smaller ones near base of femora (Fig. 315). Posterodorsal carina with 7–10 small teeth. Dorsal carinae of protibiae with only a very few minute teeth, ventral carinae densely armed with laterally flattened, pointed triangular serrations. Dorsal carinae of meso- and metafemora sparingly dentate. Two outer ventral carinae each armed with 8–14 very prominent, long and pointed spines. Ventral surfaces of meso- and metafemora convex, tectiform and medioventral carina armed with 4–7 very strong and prominent, slightly hooked spines; 2nd and 3rd usually being the largest (Fig. 319). Dorsal carinae of meso- and metatibiae sparingly dentate, antero- and posteroventral carinae raised and each densely armed with prominent, pointed triangular serrations. Medioventral carina with a much smaller number of comparatively longer and more slender spines. Probasitarsus with dorsal carina smooth and very gently elevated; all remaining carinae of basitarsi minutely serrate. Basitarsi slightly longer than following three tarsomeres combined. Nymphs: The early instars are green and only take on a brownish colouration towards the last two instars. The & penultimate instar nymph in coll. FH measures a body length of 161.5 mm and has the armature of legs distinctly less developed than the adults. The general colouration is plain yellowish brown.



119

%% (Fig. 91): Large (body length 125.5–149.0 mm) and rather slender (maximum body width 4.7–5.1 mm) for the genus with long alae (68.0–79.0 mm). General colouration of body and legs pale greenish brown to dull green, meso- and metasternum and abdominal sternites straw or pale grey. Head with a broad dark brown to black postocular marking on cheeks. Tegmina greenish brown and occasionally with an indistinct pale central spot. Costal region of alae plain greenish brown or dull green; anal region transparent grey with brown veins and slightly darker at the apices. Anterior margin of tegmina and basal 1/3 of alae with a longitudinal white exterior and a brown interior stripe. Antennae pale to mid brown and slightly darker brown ventrally. Head (Fig. 293): Generally as in && but eyes much more prominent and projecting hemisphericall; their length contained just a little more than 2x in that of cheek. Coronal line and dorso-lateral lines very faint. Antennae with 32 segments, densely sethose and projecting over posterior margin of median segment, length relations of antennomeres as in &&. Scapus and pedicellus as in &&. Thorax: Pronotum generally as in &&. Mesothorax almost 2x longer than head and pronotum combined. Mesosternum with a distinct longitudinal median keel, metasternum with a very indistinct median carina. Tegmina oval, 2x longer than wide, slightly projecting over posterior margin of metanotum and with a blunt, rounded central hump. Alae reaching half way along abdominal tergite VI. Abdomen: Median segment 2.5x longer than wide, distinctly longer than metanotum. Segments II–V slightly increasing in length, II 4x V almost 5x longer than wide. VI slightly shorter than previous, about 4.5x longer than wide. Tergite VII about ¾ the length of VI, posterolateral angles slightly expanded and with a minute triangular tooth. VIII slightly shorter than VII, strongly swollen and broadening towards the posterior; posterior margin almost 2x wider than anterior margin. IX strongly convex, narrower than previous and decidedly constricted medially, about 2.3x longer than wide. Anal segment strongly keeled and laterally compressed, truncate anterodorsally. Semi-tergites rather broad, apical half roughly triangular and gradually constricted with the apex slender and rounded (Fig. 198). Interior surfaces densely covered with minute black teeth. Cerci prominent, laterally compressed, carinate, up-curving and but staying beyond apex of anal segment. Poculum convex, cup-like with a transverse and longitudinal median carina in apical half; angle with a slight rounded hump. Legs: All long and slender, with all carinae densely but minutely serrate (more sparingly on dorsal carinae). Profemora longer and mesofemora about as long as combined length of head, pro- and mesonotum, metafemora reaching about ¼ the way along abdominal segment V and metatibiae reaching apex of abdomen. Medioventral carina of meso- and metafemora fine and armed with 4–9 small spines. Carinae of probasitarsus smooth, those of meso- and metabasitarsus minutely serrate; as long as remaining antennomeres combined except claw. Eggs (Figs. 151–152, 353): Large, capsule very gently laterally compressed, slightly oval in cross-section, and indistinctly longer than high. Capsule surrounded by very faint dorsoventral keel, beginning and ending at the operculum. Polar-area rounded if seen laterally. General colouration of capsule, micropylar plate and operculum dull mid to dark brown, capitulum dark reddish brown. Capsule densely but minutely granulose (Fig. 415). An oval, flat depression near operculum anterodorsally- and ventrally and near polar-area. Micropylar plate slightly raised from capsule and with a fine slightly darker brown outer margin. Generally shaped like a bold inverted “Y”, with all three extensions rather short. Apices of posterior extensions slightly truncate, anterior extension broadened and slightly tapered apically. Median line short but prominently raised. Micropylar cup small, knob-like and positioned centrally in posteromedian gap of the plate. Operculum flat and oval with a broad, knob-like capitulum in the centre. Measurements [mm]: length including capitulum 5.6, length 4.8, width 3.9, height 4.3, length of micropylar plate 3.8. Comments: Brunner v. Wattenwyl (1907: 184) originally described Phobaeticus sumatranus from two && ST from Sumatra in MHNG and NHMB. Due to the NHMB specimen is in very poor condition and the 120 · Zootaxa 1906 © 2008 Magnolia Press

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measurement clearly concern to the & in MHNG, the latter is here selected as the LT. Redtenbacher (1908: 453) described his Pharnacia rigida from a single & from Sumatra in NHMW not aware it was the same as Brunner v. Wattenwyl’s species. Examination and comparison of both types clearly shows Ph. rigida to be a junior synonym of Ph. sumatranus (Brunner v. Wattenwyl) (syn. nov.). Although these two species are synonymous, Brock (1996: 29) erroneously transferred Pharnacia rigida Redtenbacher to the genus Phobaeticus Brunner v. Wattenwyl, 1907 and Phobaeticus sumatranus Brunner v. Wattenwyl to Pharnacia Stål. Subsequently, Brock (1997: 45) revised his decision but only transferred Ph. rigida back to Pharnacia. Examination of the % from Sumatra in MNHU which Günther (1935b : 139) believed to be the opposite sex of Phobaeticus rex (Günther, 1928) has proven this to be a typical specimen of Ph. sumatrana. A single & from Java in the collection of H. Burmeister in MLUH bears a hand-written determination label by Burmeister stating “Phibalosoma acanthopus Burm.*”, but the HT of this species is in MNHU. Brock (1999: 132, fig. 88) listed Pharnacia cantori (Westwood, 1859) (here transferred to the genus Tirachoidea Brunner v. Wattenwyl, 1893) as the only species of Pharnacia Stål to be recorded from Peninsular Malaysia and reproduced the illustrations of both sexes by Westwood (1859, pl. 37: 1, pl. 38: 1). The photographs of the live & (Brock, 1999: pl. 26) and egg (Brock, 1999: pl. 24) and description of the egg however relate to Ph. sumatrana. Ph. sumatrana itself is omitted in the author’s book on Malaysian Phasmatodea, but Brock (1999: 133) briefly stated there may be a second species in Malaysia, commenting on T. cantori (Westwood): “…adults are virtually identical and although two distinct types of eggs have been obtained […], a series would need to be reared to establish precise details of any variation.” Brock (1999: 133) characterized the “second type of eggs” as “glossy whitish brown with a black capitulum”; these actually being T. cantori and indeed a separate species. The confusion around Ph. sumatrana and T. cantori was discussed and partly clarified by Seow-Choen (2000: 35), who provided illustrations, characterizations and a list of characters which distinguish Ph. sumatrana and T. cantori. Seow-Choen (2000: 37) cited body-lengths of 198.0-225.0 mm for && and 131.0-134.0 mm for %% of Ph. sumatrana. TABLE 11. Measurements [mm] of Pharnacia sumatrana (Brunner v. Wattenwyl, 1907).

&, LT MHNG

&&

%%

Body

196.0

189.5–218.5 (225.0)*

125.5–149.0

Head

14.2

13.1–15.0

6.4–7.0

Pronotum

8.1

8.0–9.3

4.0–4.8

Mesonotum

41.2

39.5–40.5

19.3–24.8

Metanotum

16.3

16.0–18.3

9.5–10.5

Median segment

10.8

9.5–11.0

11.5–14.0

-

-

68.0–79.0

Profemora

45.5

48.5–50.0

33.0–40.0

Mesofemora

39.8

40.0–42.5

28.0–32.5

Metafemora

46.3

45.5–50.25

43.0–45.5

Protibiae

49.5

50.0–53.7

39.6–45.0

Mesotibiae

36.0

38.0–40.0

29.5–33.0

Metatibiae

45.5

49.0–50.5

40.0–41.7

Antennae

33.2

32.0–33.5

56.0

Alae

* according to Seow-Choen (2000: 37)



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Ph. sumatrana is being cultured in Europe since the early 1980’s from stock originating from the Tapah Hills, Peninsular Malaysia. Additional stock was imported on several occasions subsequently. The original culture was misidentified as “Pharnacia cantori (Westwood)” and included on the Phasmid Study Group (PSG) culture-list as culture No. 30. In captivity in Europe this species readily accepts oak (Quercus robur & Q. petraea, Fagaceae) and bramble (Rubus fruticosus, Rosaceae) as alternative foodplants. Seow-Choen (2000: 37) stated Mangifera indica (Anacardiaceaee), Psidium guajava (Myrtaceae), Rubus fruticosus and Rubus moluccanus (Rosaceae) to be taken as foodplants in Peninsular Malaysia. Distribution (Fig. 377): Sumatra (Lakat; Mount Battak; Redjang-Lebong; Solok; Doerian Moelan: Brindjei & Indragiri), Peninsular Malaysia (Perak: Tapah Hills & Cameron Highlands) and Java. Number of specimens examined: 22

Pharnacia tirachus (Westwood, 1859) stat. rev. (Figs. 96–97, 155–156, 205–207, 294–295, 316–317, 322, 337, 378) Phibalosoma tirachus Westwood, 1859: 75, pl. 37: 3. HT, %: Cantor leg., Malacca (OXUM). Pharnacia tirachus, Brock, 1995: 95. Phobaeticus tirachus, Brock, 1996: 30. Seow-Choen, 2000: 38, pl. 97 (%). Otte & Brock, 2005: 270. Tirachoidea tirachus, Brunner v. Wattenwyl, 1893: 83. Kirby, 1904a: 359. Clitumnus irregularis Brunner v. Wattenwyl, 1907: 195. HT / ST &(&) nymph(s): Java, Buitenzorg, Pedastshenko (ZMAS – not traced). syn. nov. Baculum irregularis, Brock, 1998a: 37. Otte & Brock, 2005: 303. Ramulus irregularis, Brock, 2006: 55. [Syntypes retained in NHMW] Further material [1 &, 2 && (nymphs), 7 %%, eggs]: PENINSULAR MALAYSIA: 1 &, 3 eggs: Malacca, Perak, A. Grubauer leg.; B. Jachau vend. 15.12.1901 (ZMUH); 2 %%: Penang Hills, 1800– 2500ft., S.S. Flower; 99-248 (BMNH). JAVA: 1 & (nymph): Buitenzorg, Java occident., Pedaschenko 1901; Coll. Br. v. W.; 26.235 (NHMW, No. 335); 1 & (nymph): Buitenzorg, Java occident., Pedaschenko 1901; Coll. Br. v. W. (NHMW, No. 335); 2 && (nymphs): W. Java, Gedeh, Tjibodas, Dr. Siebert (MNHU); 3 %%: W. Java, Gedeh, Tjibodas, Dr. Siebert; Pharnacia hypharpax Westwood, 1859, K. Günther det. (MNHU); 1 %: Kendeng-Geb., O.-Java, 3-4000´, A. Heyne, Berlin-Wilm. I; Pharnacia hypharpax Westwood, 1859, K. Günther det. (MNHU); 1 %: Java (ZSMC).

Diagnosis: This species is well characterized by the very elongate, posteriorly narrowed head and flat vertex (Figs. 294–295) and very slender body of both sexes. It is closely related to the Philippine Ph. kalag Zompro, 2005 and Ph. heros Redtenbacher, 1908 from Java. From the first && are easily distinguished by: the presence of at least 3 conspicuously enlarged lobes on the anterodorsal carina of the protibiae (Figs. 316–317); distinct sub-basal lobe of the antero and posteroventral carinae of the mesofemora; more prominent and less acute serrations or lobes of the anterodorsal carina of the profemora and differently shaped lateral lobes of abdominal tergite VII. The eggs differ considerably in being more globose and having the micropylar plate considerably larger with the posterolateral extensions much more elongate (Figs. 155–156). %% differ by the relatively shorter apically more acutely triangular semi-tergites of the anal segment (Fig. 207), different leg armature and more distinct apical lobe of the mesoand metatibiae. Apart from the characteristic features mentioned also differing from Ph. heros by: the presence of


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enlarged, ± foliaceous lobes on the anterodorsal carina of the protibiae; very prominently dentate or lobed anterodorsal carina of the profemora and broader lateral lobes of abdominal tergite VII of && as well as the less constricted but acutely distinctly triangular apex of the semi-tergites of the anal segment of %% (Fig. 207). Etymology: The specific name is possibly based on “Tirachus” a shortened form of Tiberius Sempronius Gracchus (162–133 a. Chr.), a great Roman politician. Description: && (Fig. 96): Long (body length 195.0 mm) and slender for the genus (maximum body width 8.0 mm). General colouration of body and legs drab to ochracheous brown with various darker markings. Head, pronotum and tergites IV–V darker than rest of body. Head with a bold oval whitish marking between the eyes. Mesonotum slightly darker brown at anterior margin. Median segment dark greenish brown along lateral margins. Abdominal tergite II with an indistinct elongate longitudinal dark greenish brown marking on both sides of the median line. Tergites IV–X irregularly spotted with pale and mid brown, lateral margins of VI–VII slightly whitish. Apical 1/3 of profemora marbled with white, remaining portion more or less uniformly coloured. Apex of protibiae pale greyish brown. Meso- and metafemora marbled with greyish white, tibiae darker brown with slightly paler apices. Bases of basitarsi greyish white. Eyes dark orangebrown. Antennae dark brown in basal portion and becoming reddish brown towards the apex. Head (Fig. 294): Elongate, almost 2.5x longer than wide, and narrowing towards posterior margin which is slightly angled and indistinctly projecting over anterior margin of pronotum; vertex flat. Between the eyes with an indistinct curved depression. Eyes small, circular and slightly projecting from head capsule; their length contained more than 4x in that of cheek. Antennae consisting of 26–27 segments and reaching about 1/ 3 the way along mesonotum. Scapus dorsoventrally flattened, parallel-sided and about 2.5x longer than wide. Pedicellus half the length of scapus, cylindrical. Thorax: Pronotum narrower and only half as long as head, about 1.5x longer than wide and gently narrowed towards the anterior. Median transverse depression indistinct, slightly curved and not reaching lateral margins of segment. Median line distinct. Mesothorax very elongate, about 2.2x longer than head and pronotum combined and not considerably constricted anteriorly. Mesonotum with an indistinct pale longitudinal median line and very sparsely set with a few minute granules. Metanotum slightly less than half the length of mesonotum, 3x longer than wide, parallel-sided and with a very indistinct longitudinal median line. Mesoand metasternum smooth. Abdomen: Median segment slightly shorter than metanotum, 2.5x longer than wide, parallel-sided and posterior margin with a minute median tubercle. Segments II–IV increasing in length, V–VII decreasing, IV longest and almost 3x longer than wide. Tergites II–VII each with a small, scale-like posteromedian tubercle. VII ¾ the length of IV and posterolaterally dilated into a rounded lobe, which projects by about 1/3 of the body width. Sternum VII with two minute granules close to posterior margin. Tergit VIII narrower and shorter than previous, strongly convex and slightly constricted medially. IX parallel-sided, indistinctly longer than wide, roughly rectangular. Anal segment considerably longer than IX and with a fine median carina, posterior margin with a rather deep, roundly truncate median excavation; outer angles slender and truncate (Fig. 206). Supraanal plate small, broadly rounded and with a distinct median keel. Cerci small, oval in cross-section and slightly up-curving. Gonapophyses moderately elongate, up-curving and staying beyond posterior margin of anal segment. Subgenital plate strongly keeled, boat-shaped and just not reaching apex of anal segment (Fig. 205). Legs: All long and slender, mid and hind legs very gently down-curving. Profemora almost as long as proand mesonotum combined, slightly shorter than mesothorax, metafemora almost reaching to posterior margin of abdominal segment IV and metatibiae reaching posterior of abdominal segment VII. Anterodorsal carina of profemora elevated into a roughly wave-like lamella which is protruded into 10–11 broad, triangular teeth; the basal ones minute, median ones very prominent and apical ones decreasing in size. Posteroventral carina very minutely and sparingly dentate. Anterodorsal carina of protibiae raised and protruded into three prominent, irregularly rounded lobes in great distance to another; the median one largest. Occasionally there is another



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more minute lobe between the second and third one (Fig. 317). Meso- and metafemora with a broad triangular and dentate sub-basal lobe on the two outer ventral carina and a single enlarged tooth about 1/3 the way along posterodorsal carina; otherwise very minutely and sparingly serrate. Meso- and metatibiae with a prominent rounded median and apical lobe on posterodorsal carina and a rounded apical lobe on posteroventral carina (Fig. 322). Basitarsi as long as following three tarsomeres combined, all carinae more or less distinctly elevated and smooth. Nymphs: Body more slender, head more prominent and armature of legs more decided than in adults. Abdominal tergites II–VII with a more decided posteromedial tubercle and lateral lobes of VII more slender than in adults, occasionally with lateral lobes on tergite VI as well. General colouration of body and legs dark greyish brown with irregular pale greyish speckles and markings on head, mesonotum, median segment, single abdominal segments and legs which generally correspond to those in the adult & described above. Head occasionally with a bold, wedge-shaped, greenish white marking and metanotum occasionally with a distinct white posteromedial spot. %% (Fig. 97): Medium sized (body length 109.5–136.0 mm) and very slender (maximum body width 2.5 mm) for the genus, with long alae (48.7–58.5 mm). General colouration of body and legs greyish, yellowish or greenish mid brown, ventral surface of abdomen to a various degree furnished with whitish speckles. Mesoand metasternum dull green, mesopleurae with a greyish white spot close to posterior margin. Lateral margins of tergite IX with a bold ± decided white longitudinal band in posterior portion. Sternum IX with a white lateral spot. Cerci greyish mid brown. Anterior margin of tegmina and basal third of anterior margin of alae with a longitudinal white band which is interiorly bordered by a fine black line (the latter very indistinct on alae). Costal region of alae slightly lighter brown than tegmina, anal region transparent greyish brown. All veins mid brown, transverse veins very indistinct. Meso- and metafemora and all tibiae with faint whitish transverse bands, bases of all femora pale brown. Complete armature of legs black. Meso- and metabasitarsus whitish basally. Eyes dark brown. Antennae yellowish to reddish mid brown. Head (Fig. 295): Elongate, oval, almost 2.5x longer than wide, narrowing from eyes towards posterior margin, vertex flat. Between bases of antennae with a distinct transverse impression, and two very slight impressions between the eyes. Eyes prominent, strongly convex and projecting hemispherically from head capsule; their length contained almost 3x in that of cheeks. Antennae reaching about half way along median segment, with 27 segments. Scapus dorsoventrally flattened, parallel-sided, 2.5x longer than wide. Pedicellus 1/3 the length of scapus, spherical. Following antennomeres distinctly increasing in length, the 5 apical ones strongly shortened. Thorax: Pronotum distinctly shorter and narrower than head, the anterior portion very slightly narrowed. Median transverse depression distinct but short and not reaching lateral margins of segment. Mesothorax about 2.8x longer than head and pronotum combined. Mesonotum with a very fine and indistinct longitudinal median carina and sparsely set with a variable number of minute granules, occasionally with a few slightly enlarged granules along the lateral margins. Mesopleurae with a longitudinal row of minute granules. Mesosternum with a faint longitudinal median carina, metasternum smooth. Tegmina elongate, oval and with a distinct roundly conical central hump. Alae ± reaching posterior margin of abdominal tergite V. Abdomen: Median segment 2.5x longer than metanotum, 4x longer than wide and slightly tapered towards the posterior. Segments II–VI about 6.5x, VII 5x longer than wide. V occasionally with a very minute transverse posteromedian tubercle. Sternites II–VII smooth. Tergite VIII 3/4 the length of VII, 3x longer than wide, slightly widening towards posterior margin and strongly convex. IX slightly longer than VIII, constricted medially and 4x longer than wide. Anal segment distinctly shorter than IX, strongly keeled and laterally compressed. Semi-tergites short with the apex acutely triangular and the tip pointed and gently downcurving (Fig. 207). Interior surfaces apically set with several small black teeth. Cerci short, round in crosssection and tapered towards a pointed tip, not reaching apex of anal segment. Poculum strongly convex and cup-like with the centre moderately conically elevated; reaching about half way along tergite IX.


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Legs: All very long and slender, profemora a little longer than combined length of head, pro- and mesothorax, mesofemora a little longer than pro- and mesonotum combined, metafemora almost reaching posterior margin of abdominal segment V and metatibiae projecting over apex of abdomen. Anterodorsal carina of profemora with 10–16 ± prominent, black triangular serrations; these occasionally strongly enlarged and saw- or lobe-like (Fig. 337). Posteroventral carina set with a similar number of considerably smaller triangular teeth. Anterodorsal carina of protibiae smooth or sometimes sparingly serrate; occasionally with three more or less distinctly enlarged teeth. Posteroventral carinae sparingly serrate, anteroventral carina smooth. Two outer ventral carinae of meso- and metafemora minutely serrate, dorsal carinae each with only 3–7 minute teeth. Medioventral carina with 4–8 very minute spines. Occasionally the mesofemora posses a distinctly enlarged, roundly triangular sub-basal lobe on the antero- and posteroventral carinae and two or three enlarged teeth about one quarter along the dorsal carinae. An enlarged, triangular lobe may sometimes also be present about one third the way along the posterodorsal carina of the metafemora. Ventral carinae of mesoand metatibiae minutely and sparsely serrate, dorsal carinae with only a very few minute teeth. The posterodorsal carina forming a ± distinct bi-dentate lobe at the apex of tibia and occasionally bears a ± prominent and broad, triangular lobe near the middle. Probasitarsi almost 1.5x longer, meso- and metabasitarsus about as long as remaining tarsomeres combined, all carinae gently elevated and smooth. Variation: Although only a very few specimens are know, Ph. tirachus is seen to exhibit considerable intraspecific variability which concerns to both sexes. The quite mutilated and incomplete % in ZSMC is remarkable for the very prominent armature of the legs (Fig. 337), but is otherwise a typical specimen. The teeth of the anterodorsal carinae of the profemora are distinctly enlarged and saw-like, the mesofemora bear a large basal lobe on both the antero- and posteroventral carinae and have two or three distinctly enlarged teeth about one quarter the way along the dorsal carinae. The left metafemora exhibits a large triangular lobe about one third the way along the anterodorsal carina. The two && nymphs in MNHU alone, clearly show the large range of variability within && of this species. One specimen has the complete leg armature strongly reduced (Fig. 316) while it is remarkably developed in the other. The latter specimen has the anterodorsal carina of the profemora dilated into three prominent, triangular teeth and the anterodorsal carina of the protibiae has the two more apical lobes prominently enlarged and foliaceous (Fig. 317). All of the lobes of the mid and hind legs are strongly enlarged and ± foliaceous. The lateral lobes of abdominal tergite are broader than the body width and there are two slightly smaller posterolateral lobes on tergite VI in this specimen. The latter lobes are lacking and those of tergite VII decidedly less developed in the other. Eggs (Figs. 155–156): There is a glass-tube next to the ZMUH & containing three eggs, one of which has been taken from the ovipositor and hence fully developed. Unfortunately, this unique sample has the capitulum broken off. Meduim-sized, capsule very gently laterally compressed and surrounded by a very faint, blunt bulge, beginning and ending at the operculum. Seen laterally the capsule is almost circular with bold impressions on both sides of the median line, anterodorsally near the operculum and ventrally close to the polar-area. Capsule surface densely granulated. General colouration of capsule, micropylar plate and operculum blackish brown. Micropylar plate broad, distinctly raised from capsule and shaped like an inverted “Y”. All the extensions moderately long and rounded apically. Micropylar cup very small, knob-like and placed in centre of posteromedian gap of the plate. Median line indistinct. Operculum slightly oval, flat. Capitular stalk short, whitish. Measurements [mm]: length 4.0, width 3.7, height 3.9, length of micropylar plate 2.9. Comments: Westwood (1859: 75) originally described Phibalosoma tirachus from a unique % in OXUM and provided a nice illustration of the HT (Westwood, 1859: pl. 37: 3). The HT was so far untraced and not included in the numbered type-collection of OXUM. A % in the main collection with the data “Cantor, Malacca” (= Peninsular Malaysia) however matches exactly the locality, description and illustration provided by Westwood, and hence undoubtedly represents the so far untraced HT of Phibalosoma tirachus Westwood.



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The MNHU collection contains four %% from Java, which were erroneously identified as “Pharnacia hypharpax Westw.” by K. Günther, and there is a further % from Java in ZSMC. Although from Java, examination of these specimens and comparison with the HT from Peninsular Malaysia have clearly proven all five specimens to represent Ph. tirachus. Along with the %% in MNHU there are two large & nymphs which bear exatly the same data. Comparison with the %% has proven these to be the opposite sex. Furthermore, these are undoubtedly identical to Clitumnus irregularis Brunner v. Wattenwyl, 1907, which consequently becomes a junior synonym of Pharnacia tirachus (Westwood) (syn. nov.). Brunner v. Wattenwyl (1907: 195) described Clitumnus irregularis based on a nymph from Buitenzorg [= Bogor], Java in ZMAS and stated the body length to be 140.0 mm. Brock (2006: 55) stated no type specimens of C. irregularis were traced in ZMAS and that the specimen was apparently retained in NHMW. Indeed, there are two && nymphs from Java in NHMW which are clearly this species. However, due to Brunner v. Wattenwyl (1907) did not mention any material from Vienna and the specimens are considerably shorter than the length recorded for the type (< 100.0 mm), they are definitely not the type specimens. One of the & nymphs from Java in MNHU bears a hand-written note by K. Günther which states: “Die Spezies ist als Larve beschrieben und als solche in vielen Sammlungen; die erwachsenen Tiere dazu dürften der auch in diese Gattung gehörige Eucarcharus inversus Br.v.W. oder Pharnacia semilunaris Redt. Sein” [This species was described from a nymph and is represented as such in numerous collections; the adults might be represented by Eucarcharus inversus Br.v.W. or Pharnacia semilunaris Redt., which belong in the same genus]. This shows at least Günther to have identified the generic position of Clitumnus irregularis. The ZMUH contains an adult & from Perak (Peninsular Malaysia), the only one known so far, and a glass-tube with three eggs which served for the descriptions above. Ph. tirachus is apparently rare in Peninsular Malaysia but appears to be not uncommon and rather widely distributed in Java. Seow-Choen (2000: 38, pl. 97) provided an illustration of the % and cited body-lengths of 115.0–136.0 mm. Distribution (Fig. 378): Peninsular Malaysia (Malacca; Perak & Penang Hills, 1800–2500ft.) and Java (Bogor; Mount Gede: Tjibodas & Kendeng Mountains 3000–4000 ft.). Number of specimens examined: 13 TABLE 12. Measurements [mm] of Pharnacia tirachus (Westwood, 1859) stat. rev.

%, HT (OXUM)

%%

&

&&, nymphs

(MNHU)

(ZMUH)

(MNHU)

Body

120.0

109.5–119.3

195.0

146.5–148.0

Head

5.3

5.1–5.3

12.0

8.6–8.7

Pronotum

3.5

2.9–3.2

6.5

4.2–4.4

Mesonotum

24.0

21.0–23.5

38.8

29.6–30.0

Metanotum

6.0

5.5–5.8

17.5

13.6–14.2

Median segment

12.0

11.5–12.5

14.5

9.6–10.0

Tegmina

9.5

8.4–9.5

-

-

Alae

58.5

48.7–56.2

-

-

Profemora

36.0

34.1–35.1

42.0

30.2–34.2

Mesofemora

31.8

28.7–29.4

34.5

24.2–26.0

Metafemora

36.0

33.6–35.7

38.5

26.6–31.5

Protibiae

44.0

42.3–44.1

41.0

32.8–37.5

Mesotibiae

29.0

28.1–30.1

31.8

22.0–25.0

Metatibiae

37.5

36.0–37.5

47.2

27.1–31.0

Antennae

42.0

42.2–45.8

-

13.4–14.2


HENNEMANN & CONLE

6.3 Genus Phobaeticus Brunner v. Wattenwyl, 1907

FIGURES 103-107: Phobaeticus spp. 103. Phobaeticus foliatus (Bragg, 1995), & HT, Midden-East-Borneo (RMNH). 104. Phobaeticus foliatus (Bragg, 1995), % Midden-East-Borneo (RMNH) 105. Phobaeticus mucrospinosus spec. nov., & HT, W-Sumatra, Solok (RMNH) 106. Phobaeticus sinetyi Brunner v. Wattenwyl , 1907, & PLT, S-India, Madras Prov., Shembagonor (NHMW) 107. Phobaeticus sinetyi Brunner v. Wattenwyl , 1907, % LT, S-India, Madras Prov., Shembagonor (NHMW) REVISION OF PHARNACIINI


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FIGURES 108-110: Phobaeticus spp. 108. Phobaeticus magnus nom. nov., & N-Thailand, Lemgao Petchabun (coll. FH, No. 0130-1) 109. Phobaeticus magnus nom. nov., % N-Myanmar, Highland of Tenasserim (coll. FH, No. 0130-2) 110. Phobaeticus redtenbacheri (Dohrn, 1910), & Sabah, Mount Kinabalu 1500 m (ZMUH)


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Type species: Phobaeticus sobrinus Brunner v. Wattenwyl, 1907: 184, pl. 7: 1a & b, by subsequent designation of Brock, 1996: 28. Phobaeticus Brunner v. Wattenwyl , 1907: 183, pl. 7: 1a–b (in part). Brock, 1996: 28 (in part). Seow-Choen, 1997: 120, figs. 102–103. Seow-Choen, 1998a: 40. Brock, 1999: 136 (in part). Seow-Choen, 2000: 38. Otte & Brock, 2005: 268 (in part). Baculolonga Hennemann & Conle, 1997a: 347. (Type species: Cladoxerus serratipes Gray, 1835, by original designation). [Synonymised by Otte & Brock, 2005: 268] Bragg, 2001: 377 (in part). Bacteria, Burmeister, 1838: 565 (in part). Bactridium, Rehn, 1920: 242. Cladoxerus Gray, 1835: 42 (in part). De Haan, 1842: 131 (in part). Eucarcharus, Carl, 1913: 12. Günther, 1928: 218. Otte & Brock, 2005: 136 (in part). Heteronemia, Otte & Brock, 2003: 290 (in part). Lobophasma Günther, 1935b: 138 (in part). (Type species: Eucarcharus rex Günther, 1928, by original designation of Günther, 1935b: 139) syn. nov. Günther, 1953: 555. Otte & Brock, 2005: 178. Nearchus Redtenbacher, 1908: 448. (Type species: Nearchus maximus Redtenbacher, 1908, by subsequent designation of Bragg, 1995: 273) syn. nov. Dohrn, 1910: 409. Günther, 1935a: 10. Bragg, 1995b: 273, 274 (in part). Brock, 1999: 131, 181. Bragg, 2001: 388. Otte & Brock, 2005: 209. Pharnacia, Kirby, 1904a: 359 (in part—only species No’s 1, 1a, 2). Redtenbacher, 1908: 449 (in part). Karny, 1923: 240 (in part). Günther, 1932: 315. Brock, 1995: 95 (in part). Phibalosoma, Westwood, 1859: 77 (in part). Kaup, 1871a: 19. Kaup, 1871b: 37. Wood-Mason, 1877a: 161 (in part). Wood-Mason, 1877b: 346 (in part). Phryganistria Stål, 1875: 6, 63 (in part). Tirachoidea Brunner v. Wattenwyl, 1893: 83 (in part). Kirby, 1904a: 359 (in part—only species No. 3). Karny, 1923: 240 (in part). Hypermergethes Brunner v. Wattenwyl , in litt.

Description (%, &): Long to very long, mostly very elongate and extremely slender Pharnaciini with distinct sexual dimorphism. Head longer than wide and to almost spherical; vertex flat or rounded, smooth. Antennae of && longer than mesonotum, of %% as long as or longer than head and complete thorax combined. Mesothorax very elongate and slender, 2.3–3.5x longer than head and pronotum combined, parallel-sided. Mesonotum usually smooth but occasionally covered with some minute granules or blunt spines in one species. Mesosternum with a distinct longitudinal median carina; occasionally median carina also present on metasternum. Median segment at least ½ the length to slightly longer than the metanotum. %% either completely apterous,



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with rudimentary or fully developed alae. Alae at beast reaching half way along abdominal tergite V. Tegmina (if present) elongate, spatulate and strongly constricted towards the base, central hump very indistinct; at least 2.5x longer than wide. Tergite VII of && either parallel-sided or with a ± prominent posterolateral lobe; in one case tergites II–IX each bear a dentate lateral lobe. Sternum VII of && with a ± distinct praeopercular organ, formed by a pair of more or less prominent tubercles, spines or triangular lobes. Cerci cylindrical or oval in cross-section, short in &&, but elongate, constricted and incurving apically in %%. Semi-tergites of %% anal segment strongly elongated, slender and conspicuously finger-like; occasionally down-curving. Subgenital plate of && strongly keeled, either not reaching or only slightly projecting over the apex of anal segment, or very elongate, lanceolate and projecting over anal segment by as much as the combined length of abdominal tergites VII–X. Poculum of %% strongly convex and with a ± conical, backward pointing central spine at the angle. All legs very long and slender, carinae more or less distinctly serrate and / or occasionally armed with single enlarged teeth or triangular lobes; in one case with numerous prominent foliaceous appendages. Anterodorsal carina of profemora distinctly raised with prominent saw-like serrations (&& in particular); serrations usually black in %%. Meso- and metafemora with a fine and ± spinose median carina. Mesofemora distinctly longer than metanotum and median segment combined. Probasitarsus of && at least as long as remaining tarsomeres combined usually distinctly longer; much longer than remaining tarsomeres in %%. Meso- and metabasitarsus longer than following three tarsomeres combined, in && dorsal carina simple or ± distinctly raised and rounded; never considerably elevated in %%. Eggs (Figs. 169–185): Medium-sized to large. Capsule strongly laterally compressed and elliptical in cross-section, basically lens-shaped and with a prominent keel on dorsal and ventral surfaces and polar-area. Keel with a depression at micropylar plate (sometimes faint) and at polar-area. Occasionally the keel may be considerably elevated, forming flap-like extensions. Capsule surface either almost smooth and shiny, minutely punctured or distinctly sculptured. External micropylar plate short, at best covering 2/3 of capsule length, usually transverse, bilobed and generally heart or kidney-shaped. Operculum elliptical and with a distinct conical, hat or knob-like capitulum on a short stalk. Variation: In Phobaeticus Brunner v. Wattenwyl, 1907 several morphological characters of the insects and eggs are subject to considerable intrageneric variability. This e.g. concerns to: the shape and length of the subgenital plate of &&; shape of the semi-tergites of the anal segment of %%; shape of the head; relative length of the median segment; armature of the legs and length of the tegmina and alae in %%. Even the eggs show considerable variability e.g. concerning to the sculpturing of the capsule and shape of the micropylar plate. All these features serve well for the distinction of species within the genus. The && subgenital plate varies considerably in length and shape. It is either short and does not reach the posterior margin of the anal segment (e.g. Ph. kirbyi Brunner v. Wattenwyl, 1907), roughly reaches to (e.g. Ph. serratipes (Gray, 1835) or Ph. sinetyi Brunner v. Wattenwyl, 1907) or slightly projects over the same (e.g. Ph. lumawigi Brock, 1997), or is conspicuously lanceolate and projects over the apex of the abdomen by as much as the combined length of tergites VII–X (e.g. Ph. magnus nom. nov. or Ph. redtenbacheri (Dohrn, 1910)). The semi-tergites of the anal segment of %% vary in shape from being slender and straight with the apex strongly elongated and finger-like (e.g. Ph. serratipes (Gray, 1835) and Ph. redtenbacheri (Dohrn, 1910)) to rather short, broad and roundly rectangular in lateral aspect (e.g. Ph. chani Bragg spec. nov.). The head ranges in shape from elongate-oval, being distinctly longer than wide with the vertex flat (e.g. Ph. serratipes (Gray, 1835) and Ph. chani Bragg spec. nov.), to globose and almost spherical (e.g. Ph. kirbyi Brunner v. Wattenwyl, 1907 or Ph. palawanensis spec. nov.). The mesonotum is usually destitute of spines or granules as typical for Pharnaciini but one species, Phobaeticus foliatus (Bragg, 1995), has the mesonotum armed with several prominent spines. The median segment varies in length from being only half as long as the metanotum (Ph. sinetyi Brunner v. Wattenwyl, 1907) to being considerably longer than the metanotum, especially in %% (e.g. Ph. palawanensis spec. nov.). %% are either apterous (e.g. Ph. sobrinus Brunner v. Wattenwyl , 1907 or Ph. sinetyi Brunner v. Wattenwyl, 1907), have vestigial tegmina and alae (Ph. palawanensis spec. nov.), brac-


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hypterous (e.g. Ph. serratipes (Gray, 1835) and Ph. kirbyi Brunner v. Wattenwyl, 1907) or are enabled of active flight with the alae well developed and covering most of the abdomen (e.g. Ph. hypharpax (Westwood, 1859) or Ph. magnus nom. nov). The legs shows a very wide range of armature. They are only very sparingly dentate or densely serrate, may bear single enlarged teeth or lobes or may even be covered with numerous large, foliaceous appendages (Ph. foliatus (Bragg, 1995b)). The medioventral carina of the meso- and metafemora is either smooth, minutely spinulose or armed with several prominent spines. Abdominal tergite VII of && is either almost parallel-sided, posterolaterally elevated to form a more or less distinct, rounded or roundly triangular lobe, or in one case (Ph. rex (Günther, 1928)) is protruded into a large, irregularly dentate appendix which laterally extends by as much as the length of the complete segment. As an extreme, Ph. foliatus (Bragg, 1995b) exhibits a more or less prominent, dentate lateral lobe on tergites II–X in && and II–VI in %%. The egg capsule is generally restricted to the typical lens-like shape but shows variation concerning to the shape and structure of the dorsoventral keel which surrounds the capsule, shape of the micropylar plate, operculum and capitulum as well as the sculpturing of the capsule surface. Unfortunately, up to now fully developed eggs of only nine species are known, which does obviously not allow to describe the complete existing variability that occurs in eggs of Phobaeticus. Diagnosis (Table 2): Phobaeticus Brunner v. Wattenwyl is easily distinguished from all other genera of Pharnaciini by: the strongly laterally compressed, keeled and lens-shaped egg capsule; short, mostly bilobed, heart or kidney-shaped external micropylar plate which usually covers less than 2/3 of the capsule length as well as the elongate, slender and finger-like semi-tergites of the anal segment and elongate, apically constricted and in-curving cerci of %%. It is very closely related to Phryganistria Stål, 1875, with which it shares the distinct praeopercular organ of &&. Apart from the typical features mentioned above, Phobaeticus differs from Phryganistria by: the longer median segment of both sexes which is longer or at least ½ the length of the metanotum, the keeled mesosternum of %% and relatively shorter antennomeres. Certain species of Phobaeticus may at first glance show striking similarity to the more slender taxa of Pharnacia Stål, 1877 but apart from the distinguishing features mentioned above, Phobaeticus differs from that genus by: the presence of a prominent praeopercular organ and antennae of &&, being longer than the mesonotum; not conspicuously elevated carinae of the basitarsi of %% as well as the shorter and differently shaped micropylar plate of the eggs (shaped like an inverted “Y” in Pharnacia). Furthermore, the subgenital plate of && is never strikingly elongated and lanceolate in Pharnacia Stål and the median segment of && is at least 2/3 the length of the metanotum. Comments: Brunner v. Wattenwyl (1907: 183) originally placed Phobaeticus in Clitumnini and characterized it as being apterous in both sexes. Consequently, he included species most of which were only known from the &&. Only the apterous %% of Ph. sinetyi Brunner v. Wattenwyl, 1907 and Ph. fruhstorferi Brunner v. Wattenwyl, 1907 were known to Brunner v. Wattenwyl, but the latter is obviously not a member of the genus and here transferred to Phryganistria Stål, 1875 (→ 6.4). Ph. kuehni Brunner v. Wattenwyl, 1907 from Roma Island (Indonesia) is also not a member of Phobaeticus and here transferred to the genus Nesiophasma Günther, 1934c (Phasmatidae: Stephanacridini, → 5.6). Ph. incertus Brunner v. Wattenwyl, 1907 is of uncertain systematic position and here removed from Phobaeticus; it may belong in Nesiophasma Günther, 1934c (→ see below). Brunner v. Wattenwyl was not aware that some of the corresponding and winged %% of the species he described in Phobaeticus were included in Pharnacia Stål, 1877 by Redtenbacher (1908: 449) in the third part of their well-known monography “Die Insektenfamilie der Phasmiden”. Brock (1996: 26) recognize the erroneous position of Phobaeticus and transferred it from Baculini Bradley & Galil, 1977 (= Clitumnini Brunner v. Wattenwyl, 1893) to Pharnaciini. From the species originally placed in Phobaeticus by Brunner v. Wattenwyl, Brock (1996: 28) designated Ph. sobrinus Brunner v. Wattenwyl, 1907 as the type species, since it is the only species that Brunner v. Wattenwyl illustrated (1907, pl. 7: 1a–b, %). Brock (1996) proposed a re-arrangement of the genera Pharnacia Stål, 1877 and Phobaeticus Brunner v.



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Wattenwyl, 1907 and synonymised Tirachoidea Brunner v. Wattenwyl, 1893 with Pharnacia Stål. Brock (1996: 28) re-defined Phobaeticus as containing apterous %% as well as ones with shortened or fully developed alae. Consequently, and on the basis of lacking a conically swollen or bi-tuberculate vertex, Brock (1996) transferred several of the more elongate species which were included in Pharnacia Stål, 1877 by Redtenbacher (1908) to Phobaeticus. The convex, swollen and/or bi-tuberculate vertex is however not a distinctive feature between Phobaeticus Brunner v. Wattenwyl and Pharnacia Stål but a key feature that distinguishes Pharnacia Stål from Tirachoidea Brunner v. Wattenwyl , 1893 (→ 6.2 and 6.5). The latter genus is here removed from synonymy with Pharnacia Stål and re-established as a valid genus (→ 6.5., stat. rev.). As genital features and egg-morphology were generally not taken into account Phobaeticus remained as yet polyphyletic. Examination of the genitalia has shown three of the species which Redtenbacher (1908) originally described in Pharnacia Stål are not members of the tribe Pharnaciini, this is Phobaeticus annulatus Redtenbacher, 1908, Ph. enganensis Redtenbacher, 1908 and Ph. nigricornis Redtenbacher, 1908. These three species belong to the genus Sadyattes Stål, 1875 now in Stephanacridini and are here transferred (comb. nov.). Based on egg-morphology like the typically Y-shaped micropylar plate and genital features such as lacking a conspicuous praeopercular organ in && two species need to be re-transferred to Pharnacia Stål, this is Pharnacia heros Redtenbacher, 1908 and Phibalosoma tirachus Westwood, 1859 (stat. rev.). Another four species do not belong to either genus but are members of Phryganistria Stål, 1875 or Tirachoidea Brunner v. Wattenwyl (→ 6.4, 6.5). There are obvious differences which clearly distinguish Phobaeticus from Pharnacia, such as the presence of a distinct praeopercular organ in &&, the comparatively much more elongate and finger-like semitergites of the anal segment and elongate and apically constricted, in-curving cerci of %% (→ Table 1 and diagnosis above). Furthermore, eggs of Phobaeticus differ significantly from those of Pharnacia, being laterally compressed, ± lens-shaped, surrounded by a distinct dorsoventral keel and having the micropylar plate considerably shorter, not Y-shaped but bilobed, kidney or heart-shaped and usually covering less than half of the capsule length. Redtenbacher (1908: 448) established Nearchus for two newly described species, N. maximus Redtenbacher, 1908 (= Phobaeticus magnus nom. nov.) from Laos and Thailand and N. grubaueri Redtenbacher, 1908 (= Ph. (?) incertus Brunner v. Wattenwyl, 1907 syn. nov.) from Peninsular Malaysia and Banka Island in the Straight of Malacca. Redtenbacher (1908: 448) stated the new genus to be identical to Pharnacia Stål except for having a conspicuously elongate and lancet-like subgenital plate in && (apparently referring to the more elongate species which actually belong in Phobaeticus). But as discussed above the length of the subgenital plate is an extremely variable feature within Phobaeticus and the closely related Phryganistria Stål, 1875, and hence only worth for distinction at the species level. Detailed examination of N. maximus Redtenbacher, the type species of Nearchus, and comparison with species of Phobaeticus have not revealed any differences that can be regarded as being of generic value. Consequently, Nearchus Redtenbacher is here synonymised with Phobaeticus Brunner v. Wattenwyl (syn. nov.). A synonymy of Phobaeticus and Nearchus was already suggested by Seow-Choen (1998a: 40). Bragg (1995b: 237) designated N. maximus Redtenbacher, and Brock (1995: 94) subsequently selected N. grubaueri Redtenbacher (= Ph. (?) incertus Brunner v. Wattenwyl, 1907 syn. nov.) as the type species of Nearchus. Due to an earlier publication date the type selection of Bragg (1995b) has priority (Bragg, 2001: 388). Lobophasma was established by Günther (1935b: 139) to include Eucarcharus rex Günther, 1928, Eucarcharus lobulatus Carl, 1913 and Eucarcharus inversus Brunner v. Wattenwyl, 1907 of which he designated E. rex Günther as the type species. Careful investigation of E. rex and comparison with species of Phobaeticus have shown this to belong in Phobaeticus, which consequently places Lobophasma as a junior synonym (syn. nov.). This is e.g. seen in the presence of a distinct praeopercular organ, slender body and shape of the eggs


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FIGURES 110-114: Phobaeticus spp. 111. Phobaeticus redtenbacheri (Dohrn, 1910), % Sabah, Mount Kinabalu 1500 m (coll. FH, No. 0264-1) 112. Phobaeticus serratipes (Gray, 1835), & West Malaysia, captive reared (coll. FH, No. 0030-10) 113. Phobaeticus serratipes (Gray, 1835), % West Malaysia, captive reared (coll. FH, No. 0030-6) 114. Phobaeticus kirbyi Brunner v. Wattenwyl , 1907, & NE-Sarawak, Batu Niah (coll. FH, No. 0160-6)



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and micropylar plate. The large, foliaceous lateral appendages of abdominal tergite VII of Lobophasma rex, which Günther (1935b: 139) stated to be the main key feature of Lobophasma, are unique in the entire Pharnaciini, but are a considerably variable feature within Phobaeticus and therefore not of generic value. Furthermore, Lobophasma sensu Günther is seen to be polyphyletic since E. inversus Brunner v. Wattenwyl belongs in Tirachoidea Brunner v. Wattenwyl, 1893 (→ 6.5). Baculolonga was established by Hennemann & Conle (1997a: 347) to contain Ph. serratipes (Gray, 1835), Ph. kirbyi Brunner v. Wattenwyl, 1907, Nearchus redtenbacheri Dohrn, 1910 and the newly described Baculolonga philippinica. The authors designated C. serratipes Gray as the type species and distinguished Baculolonga from Phobaeticus on the basis of egg-morphology and the longer median segment of %%. However, in Brunner v. Wattenwyl’s illustration of the % HT of the type species Phobaeticus sobrinus Brunner v. Wattenwyl (1907: pl. 7: 1a) the median segment appears shorter than it actually is, which was proven by subsequent examination of the specimen. As further investigation and comparison have shown the length of the median segment to be a very variable feature in Phobaeticus, Baculolonga was correctly synonymised with Phobaeticus by Otte & Brock (2005: 268). Although the genus mostly contains very long and often extremely slender forms, the && of certain more robust species (e.g. Ph. sinetyi Brunner v. Wattenwyl, 1907) may at first glance strongly resemble some of the more slender members of Pharnacia Stål, 1877. The presence of a distinct praeopercular organ however readily distinguishes Phobaeticus-species from Pharnacia. Due to numerous single sexes and eggs are still unknown, the definition of species-groups or an intrageneric systematization remain difficult. Distribution (Fig. 84): So far recorded from India, Sri Lanka, Bangladesh, Myanmar, Laos, Thailand, Peninsular Malaysia, Singapore, Sumatra, Banka Id. prope Sumatra, Borneo, Palawan & the Philippines. Species included: 1. Phibalosoma annamallayanum Wood-Mason, 1877a: 161. comb. n. 2. Phobaeticus chani Bragg spec. nov. 3. Nearchus foliatus Bragg, 1995b: 274. comb. nov. 4. Phibalosoma hypharpax Westwood, 1859: 75, pl. 13: 6. = Pharnacia spectabilis Redtenbacher, 1908: 453. syn. nov. 5. Pharnacia ingens Redtenbacher, 1908: 453. 6. Phobaeticus kirbyi Brunner v. Wattenwyl, 1907: 185. = Pharnacia sagitta Redtenbacher, 1908: 454, pl. 22: 2. = Pharnacia pilicornis Redtenbacher, 1908: 455. 7. Eucarcharus lobulatus Carl, 1913: 12. comb. nov. 8. Phobaeticus lumawigi Brock, 1997: 43. 9. Phobaeticus magnus nom. nov.

[S-India (Travancore Province)] [N-Borneo (Sabah)] [Borneo (Sabah & Kalimantan)] [Sri Lanka] [E-India (Malabar), (Tenasserim)] [Borneo]

N-Myanmar

[Sri Lanka] [Philippines (Luzon Id.)] [Laos, Bangladesh, Thailand & Myanmar] = replacement name for Nearchus maximus Redtenbacher, 1908: 448. 10. Nearchus maximus mjöbergi Günther, 1935a: 10. [Borneo (Kalimantan)] 11. Phobaeticus mucrospinosus spec. nov. [E-Sumatra] 12. Phobaeticus palawanensis spec. nov. [Palawan] 13. Baculolonga philippinica Hennemann & Conle, 1997a: 349. [Philippines (Mindoro Id.)] 14. Pharnacia pinnipes Redtenbacher, 1908: 452. [Peninsular Malaysia (Selangor)] 15. Nearchus redtenbacheri Dohrn, 1910: 409. [N-Borneo (Sabah)] 16. Eucarcharus rex Günther, 1928: 218. comb. nov. [Sumatra & W-Borneo (Sarawak)] = Phobaeticus lambirica Seow-Choen, 1998a: 40, fig. 1. syn. nov. 17. Cladoxerus serratipes Gray, 1835: 42. [Peninsular Malaysia, Singapore,


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Sumatra] = Phasma (Bacteria) acanthopus Burmeister, 1838: 565. = Bactridium grande Rehn, 1920: 242, pl. 10: 12. syn. nov. = Phibalosoma maximum Bates, 1865: 341. syn. nov. 18. Phobaeticus sinetyi Brunner v. Wattenwyl, 1907: 184. [SE-India (Madras Province)] 19. Phobaeticus sobrinus Brunner v. Wattenwyl, 1907: 184, pl. 7: 1. [Sumatra] = Phobaeticus beccarianus Brunner v. Wattenwyl, 1907: 185. syn. nov. ? Phobaeticus incertus Brunner v. Wattenwyl, 1907: 185. [Peninsular Malaysia & Banka Id.] = Nearchus grubaueri Redtenbacher, 1908: 448. syn. nov.

Keys to the species of Phobaeticus Brunner v. Wattenwyl , 1907

&&* 1. Mesonotum unarmed; legs without large foliaceous appendages; no lateral lobes on abdominal tergites II– VI ..................................................................................................................................................................2 - Mesonotum spinose; mid and hind legs with numerous large, foliaceous appendages (Fig. 326); tergites II–VI with dentate lateral lobes; Borneo ............................................................................................foliatus 2. Abdominal segments III–VI at best 4.5x longer than wide ..........................................................................3 - Extremely slender and elongate species (body length > 330.0 mm); abdominal segments III–VI 6.5x longer than wide; Northwestern Borneo....................................................................chani Bragg spec. nov. 3. Subgenital plate not or indistinctly projecting over posterior margin of anal segment ................................4 - Subgenital plate elongate and ± distinctly projecting over posterior margin of anal segment ...................12 4. Head distinctly longer than wide, vertex flat................................................................................................5 - Head strongly convex, almost spherical (Fig. 307); Borneo ................................................................ kirbyi 5. Lateral lobes of tergite VII projecting by less than body width ...................................................................6 - Lateral lobes of tergite VII irregularly leaf-like, dentate and projecting by more than the body width (Fig. 270); Sumatra & NW-Borneo....................................................................................................................rex 6. Median segment at least 2/3 the length of metanotum ................................................................................8 - Median segment only about half as long as metanotum; SE-India & Myanmar..........................................7 7. All tibiae with distinct dorsal lobes; abdominal tergite VII with a prominent, rounded lateral lobe (Fig. 271); praeopercular organ formed by a single lobe-like, triangular appendage .................................. ingens - Tibiae without conspicuous lobes; abdominal tergite VII at best with a slight lateral lobe; praeopercular organ formed by two spines or ± enlarged triangular lobes ................................................................ sinetyi 8. Posteroventral carina of mesofemora with three prominent, dagger-like teeth in basal section (Fig. 328); E-Sumatra ............................................................................................................. mucrospinosus spec. nov. - Armature of posteroventral carina of mesofemora different ........................................................................9 9. Lateral lobes of tergite VII indistinct; not Sri Lanka..................................................................................11 - Lateral lobes of tergite VII prominent, rounded or truncate; Sri Lanka .....................................................10 10. Lateral lobes of tergite VII rounded; tergite VIII shorter than IX and X combined; medioventral carina of meso- and metafemora smooth; mesopleurae unarmed; lobes of praeopercular organ acute ...... hypharpax - Lateral lobes of tergite VII truncate; tergite VIII longer than IX and X combined; medioventral carina of meso and metafemora spinose; mesopleurae granulose; lobes of praeopercular organ broad and rounded .. .........................................................................................................................................................lobulatus 11. Very long species (body length 228.0–278.0 mm); median segment longer than metanotum .......serratipes - Smaller species (body length ± 200.0 mm); median segment shorter than metanotum ...................sobrinus 12. Subgenital plate at best projecting over apex of abdomen by length of anal segment; Philippines...........13



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Subgenital plate very elongate, lancet-like and projecting over anal segment by at least the length of tergites IX and X combined; not Philippines....................................................................................................14 13. Tergites II–VII with a posteromedian tubercle; apex of subgenital plate rounded (Fig. 246); Mindoro Id ... ...................................................................................................................................................philippinicus - Tergites II–VII smooth; apex of subgenital plate pointed (Fig. 254); Luzon Id..............................lumawigi 14. Armature of legs ± regular; without conspicuously enlarged lobes or teeth ..............................................15 - Mesofemora with slightly enlarged dorsal teeth and enlarged ventral spines sub-basally; mesotibiae with a prominent triangular dorso-median lobe; very large species (body length (275.0–300.0 mm); N-Borneo ... ................................................................................................................................................. redtenbacheri 15. Medioventral carina of meso- and metafemora spinose (Fig. 329); Laos, Bangladesh, Thailand & Myanmar ....................................................................................................................................magnus nom. nov. - Medioventral carina of meso- and metafemora unarmed; Central Borneo ....................................mjoebergi * && of Ph. palawanensis spec. nov. are not known; body lengths given are including the subgenital plate. Since Ph. incertus Brunner v. Wattenwyl, 1907 is probably not a member of this genus it is excluded from the key above.

%%* 1. Mesonotum not spinose; legs without large foliaceous appendages; tergites II–VI without lateral lobes ...2 - Mesonotum spinose (Fig. 312); legs with numerous large, foliaceous appendages; tergites II–VI with dentate lateral lobes; Borneo ....................................................................................................................foliatus 2. Apterous........................................................................................................................................................3 - Alate..............................................................................................................................................................6 3. Head distinctly longer than wide; median segment shorter than metanotum; not Palawan .........................4 - Head spherical (Fig. 312); median segment 1.5x longer than metanotum; Palawan...................................... ................................................................................................................................ palawanensis spec. nov. 4. Median segment at least 2/3 the length of metanotum; not India.................................................................5 - Median segment half the length of metanotum; SE-India ................................................................... sinetyi 5. Large species (body length 143.0–165.5 mm); tergites III–VI 7x longer than wide; semi-tergites of anal segment tapered and acute apically (Fig. 242); Philippines ......................................................philippinicus - Smaller species (body length 125.0 mm); tergites III–VI 4.5x longer than wide; semi-tergites of anal segment rounded apically (Fig. 264); Sumatra ......................................................................................sobrinus 6. Alae well developed; projecting over posterior margin of abdominal segment IV......................................7 - Brachypterous; alae at best slightly projecting over posterior margin of abdominal tergite III .................10 7. Semi-tergites of anal segment slender, elongated and ± finger-like; poculum with a ± distinct central hump or spine..........................................................................................................................................................8 - Semi-tergites of anal segment broad, short and roundly rectangular in lateral aspect (Figs. 228 & 261); poculum without a central hump or spine.....................................................................................................9 8. Large (body length 132.5 mm); head with a distinct, black longitudinal postocular stripe; semi-tergites of anal segment 2x longer than high (Fig. 261); N-Borneo ...........................................chani Bragg spec. nov. - Smaller (body length 117.0–119.2); head without a distinct postocular stripe; semi-tergites Of anal segment broader, 1.5x longer than high (Fig. 228); SE-India & Myanmar .............................................. ingens 9. Large and very slender species (body length 155.0–176.0 mm); head 2x longer than wide, vertex flat (Fig. 311); Laos, Thailand, Bangladesh & Myanmar................................................................magnus nom. nov. - Smaller, more robust species (body length 113.5–124.5 mm); head 1.5x longer than wide, vertex slightly convex; Sri Lanka ......................................................................................................................... hypharpax 10. Head indistinctly longer than wide, globose to almost spherical (Figs. 308 & 310); Borneo....................11


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-

Head elongate, distinctly longer than wide, vertex flat (Fig. 306); Peninsular Malaysia, Singapore & Sumatra ...........................................................................................................................................serratipes 11. Lobes of anal segment straight (Fig. 233); posteroventral carina of profemora dentate alae short, reaching posterior margin of tergite II (29.0 mm).................................................................................. redtenbacheri - Semi-tergites of anal segment down-curving (Fig. 239); posteroventral carina of profemora unarmed (Fig. 346); alae longer, reaching posterior margin of tergite III.................................................................... kirbyi * %% of Ph. lobulatus (Carl, 1913), Ph. lumawigi Brock, 1998, Ph. mucrospinosus spec. nov., Ph. mjoebergi (Günther, 1935) and Ph. rex (Günther, 1928) are not known

Phobaeticus annamallayanus (Wood-Mason, 1877) comb. nov. Phibalosoma annamallayanum Wood-Mason, 1877a: 161. ST: &: Southern India, Annamallay Forests, leg. Colonel R.C. Beddome (in alcohol – not traced in NZSI, presumed lost); ST, &: Southern India, Travancore Hills, leg. F.Day (valde defectum – not traced in NZSI, presumed lost). Phibalosoma annamalayanum, Wood-Mason, 1877b: 346. [Misspelling] Pharnacia annamallayanum, Kirby, 1904a: 359. Brock, 1996: 27. Otte & Brock, 2005: 264.

Etymology: The specific name refers to the type-locality, the Annamallay Hills in southern India. Comments: Wood-Mason (1877a: 161) originally described Phibalosoma annamallayanum from two && collected in the Annamallay Forests and Travancore Hills in southern India. Wood-Mason’s type specimens are not traced in NZSI (personal communication with Dr. M. S. Shishodia) and inquiries to the museum in New Dehli have received no reply. Consequently, the types could not be examined for this study and must be presumed lost. This species is hence not included in the keys above. Unfortunately, the original diagnosis by Wood-Mason is very brief and does not mention any features that serve for distinguishing Ph. annamallayanum from related taxa. Wood-Mason (1877a: 161) stated it to be a “characteristically Malayan form” which was very close to Ph. acanthopus Burmeister, 1838 (= Phobaeticus serratipes (Gray, 1835) and commented: “Very closely allied to the proceeding [Ph. serratipes], from which it differs in its stouter body, in its shorter and thicker legs, and in the relative proportions of the different parts of the body, particularly the meso- and metathorax.”. Subsequently, Wood-Mason (1877b: 346) discussed the proposed relation to Ph. serratipes in more detail and stated Ph. annamallayanum to exhibit “a lamellar process […] at the hinder end of the sixth ventral segment of the abdomen [referring to the praeopercular organ on sternum VII]” and the metanotum to be conspicuously longer than the median segment. The presence of distinct praeopercular organ on abdominal sternum VII, the length relation of the metanotum and median segment and distribution in Southern India undoubtedly place Ph. annamallayanum in Phobaeticus Brunner v. Wattenwyl, 1907 (comb. nov.). Wood-Mason (1877a: 161) cited a body length of ca. 220.0 mm for the & from Annamallay Forest. Ph. annamallayanum is apparently close to Phobaeticus sinetyi Brunner v. Wattenwyl , 1907 also from South India and might even be conspecific. This is emphazised by both species being sympatric on Annamallay Hills. However, due to the insufficient description by Wood-Mason (1877a & 1877b) and unavailability of the type specimens any decision about the identity of Ph. annamallayanum and possible synonymy with Ph. sobrinus must await availability of newly collected material including the so far unknown %% from the typelocality of Ph. annamallayanum.



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Phobaeticus chani Bragg spec. nov. (Figs. 116–117, 181–183, 257–262, 313, 338, 379) HT: & & eggs (removed from body): Sabah, Penampang district, Ulu Moyog, c. 650 m, Native, 25.II.1989 (BMNH, ex coll. CLC). PT, &: Sabah, Buntui, Sunsuron, J. Kulip, 10.X.1989 (FRSC). PT, %: Sabah, Sipitang, Coll. Yong Ah Lok, 15.IV.1983 (CLC). Further material [3 %%]: 1 %: Sabah, Kinabalu Park HQ, 1600 m; coll. Toru Kikuta 03.IV.1993 (KNP, No. SP/EPH/00021); 1 %: Kinabalu Park H.Q., Sabah E. Malaysia, 1500 m (a.s.l.), 5.IV.1991, Coll. H. Sadanori (KNP, No. SP/EPH/00015); 1 %: Kinabalu Park, Sabah E. Malaysia, 600 m (a.s.l.), 21.7.1994, leg. Ladson (KNP, No. SP/EPH/00020). [Examined from photographs only]

Diagnosis: This is the longest known extant insect of the world && being readily distinguished from all other members of Phobaeticus by the extremely slender body and strongly elongated body segments, the abdominal segments II–VI being up to 6.5x longer than wide (at best 4.5x longer than wide in all other known species). The striking eggs of Ph. chani Bragg spec. nov. differ from all other eggs known from the genus by having the dorsoventral keel of the capsule elevated into large leaf-like lobes, and comparatively smaller oval micropylar plate (Figs. 181–183). Closely related to the Bornean Ph. redtenbacheri (Dohrn, 1910), Ph. mucrospinosus spec. nov. from Sumatra and Ph. serratipes (Gray, 1835) from Peninsular Malaysia, Singapore and Sumatra. From the first && differ by: the larger size; slightly shorter median segment which is about equal in length to the metanotum; conspicuously shorter not lanceolate subgenital plate; lack of distinct spines of the praeopercular organ and lack of a distinct median lobe on the posterodorsal carina of the meso and metatibiae. Ph. mucrospinosus spec. nov. differs in the presence of remarkable dagger-like spines on the ventral carinae of the mesofemora and the much smaller size. From Ph. serratipes && differ by: the larger size and more slender body; lack of distinct triangular lobes of the praeopercular organ; enlarged sub-basal spine on the ventral carinae of the meso and metafemora and less numerous serrations of the profemora. Also related to Ph. kirbyi Brunner v. Wattenwyl, 1907 from Borneo, but the larger size, relatively longer body segments, longer subgenital plate and more elongate and flat head easily distinguished it from this species. Due to having well developed alae which reach about half way along abdominal segment V, broad semi-tergites of the anal-segment and an unarmed poculum (Fig. 261) %% strongly resemble those of Ph. ingens (Redtenbacher, 1908) from SE-India and N-Myanmar. They are however readily distinguished by the larger size, more elongate body, dark postocular stripe of the cheeks and comparatively more slender semi-tergites of the anal-segment. Etymology: This new species, the longest known extant insect of the world, is named in honour of C.L. Chan (Kota Kinabalu, Sabah), who kindly loaned the & HT and % PT from his extensive phasmid collection and agreed to deposit both specimens in The Natural History Museum, London (BMNH). Description: Both && are damaged. The HT has part of left fore tibia, the fifth tarsomere of the right fore leg, and both antennae missing; the abdomen has been damaged by insect pests: five holes have been eaten into the dorsal surface, including one small hole in the anal segment (Fig. 257), and there is a small hole in the left hand edge of the pronotum (Fig. 313). The & PT has the left mid leg and right hind leg missing, tarsomeres 1–4 are missing from the right foreleg, and 5th tarsomere from left fore leg; the antennae are bent and difficult to measure. The % PT lacks the left fore leg and has a broken right antenna; the fore tarsus has been glued back in position. && (Fig. 116): A very long and very slender species (body length incl. subgenital plate 333.5–357.0 mm) with spinose mid and hind femora. The HT is dark green with some whitish blotches on the thorax abdomen and legs; the head is creamy-brown with a brown line running from the eye to the back of the head (Fig. 313).


HENNEMANN & CONLE

The PT is discoloured, yellowish-brown rather than green, but the head is similarly marked and pale markings can be seen on the legs and body. Head, thorax and abdominal segments 1–6 smooth, 7–11 sethose. Head broader than body, narrowing at the posterior. Pronotum narrow, widening towards posterior, twice as long as wide. Mesonotum widening evenly from about 4.0 mm at anterior to 5.0 mm at posterior, rest of body of uniform almost width (5.0–5.5 mm). Metanotum and median segment of equal length. Body without any lobes or spines. Abdominal segments I–VI of increasing length, VII equal to II, combined length of VIII– IX less than VII. Segments X–IX with slight longitudinal carina. Supraanal plate short, triangular (Fig. 257). Apex of sternum VII with a depression. Abdominal tergites VII–XI, sternum VII, and subgenital plate sethose. Subgenital plate straight with a fine longitudinal carina, apex rounded (Figs 258–259), 1.0 mm longer than abdomen in HT, 3.0 mm in PT. Cerci conical, apex slightly upturned. Gonapophyses protruding slightly beyond supraanal plate. All tibiae and femora with five distinct carinae which bear stiff setae. In most cases the carinae are expanded, to some degree, as a serrated lamella, the exceptions are: dorso-posterior of the profemur and protibia, and the medio-ventral of the meso and metafemora. Meso and metafemora spinose. All tarsi with a dorsal lobe on segments 1–3. Profemur and fore tibia triangular in cross section, the two dorsal carinae so close they are almost fused. All carinae except the dorso-posterior carina are lamellate; ventro-posterior and dorso-anterior carinae are serrated: dorso-anterior of fore femur with about 12 particularly notable serrations which decrease in size distally. Base of profemur strongly compressed and curved. The carinae of the mesofemur are armed with spines and large triangular spine-like lobes, particularly notable are a large triangular lobe (2.5–3.0 mm long) near the base of the ventro-anterior carina, and one or two near the middle of the dorso-posterior carina (Fig. 338): HT with two on right leg and one on left, paratype without lobes on dorso-posterior. Dorso-anterior carina with only a few relatively small spines. Mesotibia with medio-ventral carina with triangular serrations, all others strongly spinose; dorso-posterior with a low-lying serrated lobe at the mid point (HT with a second very small serrated lobe on left tibia). Metafemur with two notable spine-like lobes: one near the base of the ventro-anterior carina and one on the apical third of the dorso-posterior. Medio-ventral carinae with few spines. All ventral carina with spines, dorsal carinae serrated. Metatibiae armed as mesotibiae but the dorso-posterior carina has only two small lowlying serrated lobes in the mid section. %% (Fig. 117): The male is brown with small white blotches on the legs, particularly on the hind tibiae. Some areas of the body and legs suggest the specimen may originally have been greenish in places. The head has a very dark brown line running from the eye to the back of the head. Costal margins of elytra and hind wings white. Body without any lobes or spines, mesonotum sparingly granulose, rest of body smooth. Head broader than body, narrowing at the posterior. Pronotum narrow, widening towards posterior, about 1.5 times longer than wide. Body of uniform width (slightly less than 2mm), except abdominal segment VIII which widens to 2.8mm at the posterior. Metanotum and median segment not examined (hidden by wings). Anal segment slightly laterally compressed, rectangular when viewed laterally (Fig. 261). Poculum hemispherical (apex hidden by lateral margins of tergite IX, possibly due upward movement caused by shrinkage of abdomen). Cerci blunt, slightly swollen cylinders. Wings reaching just beyond mid point of abdominal segment V. Legs with most of the carinae serrated with triangular spines. All tibiae with a triangular spine-like lobe on dorso-posterior carina before the mid point; mid and hind femora with similar, but smaller, lobes in this position. Mid and hind tibiae with apical lobe on both posterior carinae. Mid and hind femora with more serrations ventrally than dorsally; medio-ventral carinae less serrate than ventro-anterior and ventro-posterior. Fore femur with dorso-anterior and ventroposterior carinae serrated; dorso-posterior with only one or two serrations; ventro-anterior and medio-ventral not serrated. Mid and hind tibiae with very few spines on dorso-anterior carinae. Fore tibia with ventro-anterior and medio-ventral carinae unarmed; dorso-posterior with only one small spine and one large triangular lobe.



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Eggs (Figs. 181–183): When the HT was originally collected it laid one or more eggs. One of these was illustrated by Chan and the illustration was seen by Bragg in 1990. However attempts to locate the egg(s) or drawing have been unsuccessful. The following description is based on eggs removed from the body of the HT. Capsule and operculum smooth, mid brown with a pale brown line along the ventral extension. The capsule is a complex shape: it appears to be a typical laterally compressed sphere but with very large, curved extensions on midline of the dorsal and ventral surfaces. Viewed from the opercular end these extensions curve in a clockwise direction, they also extend beyond the polar end of the capsule; the extension is not present on the micropylar plate. Micropylar plate oval, positioned at about the mid point of the egg, near the polar end of the main capsule. Operculum oval, conical, capitular stalk quite long, capitulum small (possibly larger when laid). The unusual shape makes useful measurements difficult; there is no clear demarcation between the capsule and the extensions to the capsule. Measurements [mm]: Overall size (including extensions): length 9.4, width 2.4, height 4.5. Main capsule: length c. 6mm, width 2.4, height c. 3.3, capitular stalk c. 0.7. TABLE 13. Measurements [mm] of Phobaeticus chani Bragg spec. nov.

&, HT (BMNH)

&, PT (FRCS)

%, PT (BMNH)

Body

356.0

330.5

132.5

Body (incl. subgen. pl.)

357.0

333.5

-

Head

11.0

10.5

5.5

Pronotum

8.5

9.0

4.0

Mesonotum

69.0

64.0

24.5

Metanotum

24.0

23.0

-

Median segment

24.0

22.0

-

Tegmina

-

-

9.0

Alae

-

-

59.0

Subgenital plate

27.5

27.0

-

Profemora

79.0 / 79.0

80.0 / 81.0

- / 35.0

Mesofemora

71.0 / 70.0

- / 66.0

30.0 / 30.0

Metafemora

76.0 / 75.0

74.0 / -

36.0 / 36.0

Protibiae

> 87.0 / 117.0

114.0 / 115.0

- / 43.5

Mesotibiae

80.0 / 78.0

- / 77.0

31.0 / 31.0

Metatibiae

88.0 / 88.0

92.0 / -

37.0 / 37.0

Protarsi

- / > 24.0

> 25.0 / > 15.0

- / 11.0

Mesotarsi

23.0 / 23.0

- / 23.0

10.0 / 10.0

Metatarsi

24.0 / 23.0

25.0 / -

12.0 / 12.0

Antennae

-

41.0

39.0

Overall length

567.0

549.0

-

* Table shows measurements for both the left and right legs

Comments: This new species is the world’s longest known extant insect. The longest specimen, the & HT in BMNH, has a body length of 357 mm, exceeding the previous record of Phobaeticus kirbyi Brunner v. Wattenwyl by 40 mm. The overall length with the fore legs stretched out straight is 567 mm, but this would have been greater by 7–10 mm if the specimen was complete; this exceeds the previous record of 555 mm for a


HENNEMANN & CONLE

specimen of Phobaeticus serratipes (Gray) recorded by Seow-Choen (1997: 33). The egg of this species is not typical of the genus although it can be considered a modification of the standard Phobaeticus pattern. The main capsule, operculum, and capitulum appear to be more or less typical but the large extensions on the dorsal and ventral surfaces radically alter the appearance. There are several possible evolutionary advantages of this egg shape. Related species are known to drop or flick their eggs from height. The extensions to the capsule undoubtedly increase air resistance and hence slow the fall of the egg which would offer two advantages: impact with the ground is lessened, reducing risk of damage and a slow descent would increase the dispersal by the wind. Other possibilities are that air trapped in the curved extensions would help keep the egg afloat in the event of flooding, and that the extensions would increase the chances of the egg becoming trapped in crevices above the ground. Three %% in the Kinabalu Park collection (KNP) appear to be this species, based on photographs taken in December 2006. However, a more detailed examination is needed to confirm their identity. Distribution (Fig. 379): Northern Borneo, endemic. Sabah (Penampang District: Ulu Moyog, c. 650 m; Sibitang; Kinabalu National Park 600 m; Kinabalu National Park headquarters 1500–1600 m & Buntui: Sunsuron). Number of specimens examined: 6

Phobaeticus foliatus (Bragg, 1995) comb. nov. (Figs. 103–104, 229–231, 276, 312, 326–327, 380) Nearchus foliatus Bragg, 1995b: 274, figs. 1–5. HT, &: H.C. Siebers, Midden O-Borneo, 21. VIII. 1925 (RMNH). Bragg, 2001: 389, fig. 146. [Description of %] Otte & Brock, 2005: 209. Further material [2 %%, 1 &]: BORNEO, KALIMANTAN: 1 %: H.C. Siebers, Midden O-Borneo, 12.X.1925, 160; Nearchus foliatus Bragg, 1995, det. P.E. Bragg, II.1997 (RMNH). N-BORNEO, SABAH [examined from photographs only]: 1 %: Borneo, Sabah, Mt. Kinabalu, Silau Silau Trail c. 5200’, 26th September 1992; Coll. C.L. Chan & Poon (coll. CLC); 1 &: Silau-Silau Trail, 16.VIII.2006, Coll. MLT Bushell (coll. CLC).

Diagnosis: Related to Ph. redtenbacheri (Dohrn, 1910), but well characterized and easily distinguished from this and all other members of the genus by the spinose mesonotum, prominent leaf-like lobes of the legs and lateral lobes of abdominal tergites II–IX of both sexes. Etymology: “Foliatus” (lat. = with leaf-like lobes) refers to the prominent leaf-like lobes of the legs of this spectacular species. Description: Bragg (2001: 389) provided detailed descriptions of both sexes. The & HT has suffered some damage: the right fore leg, parts of the left protarsus and most of the antennae are missing and the apex of the abdomen appears to have been laterally compressed during preservation. The description of the % is based on the specimen in RMNH only. && (Fig. 103): Long (body length 192.0 mm; incl. subgenital plate 214.0 mm) and rather robust species (maximum body width 7.5 mm) with a long, lanceolate subgenital plate (41.0 mm), spinose mesonotum, laterally lobed abdominal tergites and striking leaf-like lobes on the legs. General colouration of body and legs almost uniformly mid-brown (HT) to greenish brown (& in coll. CLC). The latter specimen has the entire body set with irregular darker brown speckles, the bases of all femora pale brown and the legs prettily furnished with irregular pale transverse bands and pale brown mottling on the leaf-like appendages. Lateral appendages of abdominal tergites dark brown to black with conspicuous pale markings along the outer mar-



141

gin. Eyes dark reddish brown. Head: 1.4x longer than wide, globose, vertex convex and smooth. Eyes rather large, circular and projecting hemispherically; their length contained contained about 3x in that of cheek. Antennae rather long and reaching about half way along metanotum. Scapus dorsoventrally flattened, 3x longer than wide and narrowed towards the base. Pedicellus distinctly shorter than scapus, cylindrical. Following antennomeres very elongate and increasing in length. Thorax: Pronotum slightly shorter and narrower than head, indistinctly longer than wide and slightly narrowed in anterior portion; lateral margins gently raised. Median transverse depression shallow, slightly curved and not reaching lateral margins of segment. Mesothorax very slightly widening towards the posterior and about 2.5x longer than head and pronotum combined. Mesonotum smooth dorsally, lateral margins each set with 2–5 distinct, more or less paired and 2–5 smaller irregularly positioned spines. Metanotum about 2.5x longer than wide, rectangular and lateral margins each set with three rather small spines. Meso- and metapleurae and sterna smooth. Abdomen: Median segment ¾ the length of metanotum, parallel-sided, 2x longer than wide and with a small posteromedian tubercle. Segments II–VII increasing in length, II 2x and VII 3x longer than wide, all parallel-sided. Tergites II–IX with a small posteromedian tubercle and II–X each with an irregularly dentate, leaf-like lateral expansion in posterior half; this is large on II–VI, most prominent on VII (Fig. 276), small on VIII–IX and very indistinct on anal segment (X). Sternites II–VI smooth. Praeopercular organ formed by a pair of triangular lobes at posterior margin of sternum VII. Tergite IX slightly more than half the length of VIII. Anal segment with a broadly rounded posteromedian indention, the angles acute (Fig. 230). Supraanal plate small, triangular and keeled. Cerci reaching to apex of anal segment, slender and densely sethose. Subgenital plate strongly keeled, moderately lanceolate, tapered towards an acute apex and projecting by 22.0 mm (HT) over posterior margin of anal segment (Fig. 229); surface rugulose. Legs: All long and slender, mid and hind legs densely armed with large teeth and furnished with large leaf-like lobes. Profemora a little longer than head, pro- and mesonotum combined, mesofemora almost as long as mesothorax, metafemora reaching to posterior margin of abdominal segment V and metatibiae roughly reaching apex of subgenital plate. Anterodorsal carina of profemora densely serrate, posterodorsal carina with a large foliaceous lobe 1/3 the way along femur and posteroventral carina with irregular triangular serrations of different sizes (Fig. 327). Posterodorsal carina of protibiae with three slightly enlarged triangular lobes in great distance and 3–4 lobes in apical portion. Posteroventral carina serrate and with a slightly enlarged triangular tooth about 1/3 the way along tibia. Meso- and metafemora with four large foliaceous lobes on posterodorsal carina, two outer ventral carinae sparsely dentate and with four paired, distinct triangular teeth in great distance to another (Fig. 326). Medioventral and anterodorsal carinae distinct but unarmed. Meso- and metatibiae with three large foliaceous lobes on posterodorsal carina and a small triangular tooth apically. Posteroventral carina dentate and with two prominent foliaceous lobes in basal portion, anteroventral carina serrate, medioventral and anterodorsal carinae unarmed. All basitarsi longer than following three segments combined, dorsal carina slightly raised and tri-dentate, ventral carinae minutely tri-dentate. %% (Fig. 104): Medium sized (body length 124.0 mm) and moderately slender for the genus with long alae (49.0 mm), a spinose mesothorax, laterally lobed abdominal tergites and several acute triangular teeth and sometimes striking leaf-like lobes on the mid and hind legs. General colouration of body and legs greenish brown, mesonotum slightly darker and with a blackish longitudinal band along lateral margins. Tegmina and costal region of alae pale greyish or greenish brown and furnished with a few faint darker markings, veins sepia. Anterior margin of tegmina and basal portion of anterior margin of alae very pale green. Anal region of alae transparent grey with brown veins. Armature of legs mid brown, the leaf-like- lobes of the mid and hind legs to a variable degree mottled with pale cream. Bases of all femora straw to pale greenish brown. Eyes dull reddish brown with blackish mottling. Antennae mid brown and becoming paler towards the base. Head (Fig. 312): Generally as in && and slightly narrowed posteriorly. Eyes very prominent, almost


HENNEMANN & CONLE

spherical and projecting strongly from head capsule; length contained a little less than 3x in that of cheek. Antennae projecting over posterior margin of abdominal tergite II, otherwise as in &&. Thorax: Pronotum distinctly shorter and narrower than head, slightly longer than wide, posterior and anterior margin slightly raised. Transverse median depression shallow and almost straight, not reaching lateral margins of segment. Mesothorax slender and about 2.6x longer than head and pronotum combined. Mesonotum parallel-sided only very slightly widened at posterior margin and armed with 9–12 laterally directed small to medium spines (Fig. 312). Mesosternum with a faint longitudinal median carina. Metasternum and pleurae smooth. Tegmina just covering base of alae, elongate, oval and strongly constricted towards the base; central portion roundly convex. Alae reaching to middle of abdominal tergite V. Abdomen: Median segment 2x longer than metanotum. Tergites II–VI ± 4.5 longer than wide, parallelsided and very slightly broadened at posterior margin. V and VI with a pair of small posteromedial tubercles. VII about 2/3 the length of VI, slightly widening towards posterior margin and 2.5x longer than wide. VIII 2/ 3 the length of previous, strongly convex, IX almost quadrate. Anal segment longer than VIII and IX combined, strongly tectiform and laterally compressed. Semi-tergites long, straight and slender with the apex rounded (Fig. 231). Interior surfaces with the outer margin raised and densely set with minute black teeth; a very few teeth in central portion. Cerci slender, round in cross-section and with the apex slightly in-curving and constricted. Poculum strongly convex, cup-like, almost reaching posterior margin of tergite IX and angular with a conical central hump. Legs: All moderately long and slender, profemora longer than head, pro- and mesothorax combined, mesofemora as long as pro- and mesonotum combined, metafemora slightly projecting over abdominal segment VI and metatibiae slightly exceeding apex of abdomen. Anterodorsal carina of profemora with ± 20 irregular, acutely triangular serrations of variable sizes. Posterodorsal carina with a single tooth or prominent triangular lobe about ¼ off the base. Posteroventral carina serrate and with several enlarged triangular teeth in particular in the basal and apical portions. Posterodorsal carina of protibiae serrate and with 3–4 large teeth or tooth-like lobes, anterodorsal and posteroventral carinae minutely serrate. Mid and hind legs furnished with large leaf-like lobes similar to && (coll. CLC), or the dorsal armature only represented by enlarged triangular teeth, all carinae acutely serrate. Medioventral carina of meso- and metafemora distinct but unarmed. All carinae of meso- and metatibiae minutely but acutely serrate, the posterodorsal carina with three triangular lobelike teeth or prominent lobes; generally as in &&. Basitarsi longer than combined length of remaining segments, as in &&. Variation: Ph. foliatus shows considerable variability in particular concerning to the armature of the legs. The % from Kalimantan in RMNH (Fig. 104) and % from Mount Kinabalu in coll. CLC appear to represent the opposite extremes of the variation range in this species, the armature being remarkably more strongly developed in the specimen from Mount Kinabalu. The posterodorsal carinae of the mid and hind legs of this specimen bear very prominent leaf-like lobes similar in shape and relative size to those seen in &&, whereas these are only represented by enlarged, acutely triangular teeth on the hind legs of the % from Kalimantan in RMNH (this specimen has the mid legs regenerated which probably accounts for the absence of foliaceous lobes, Fig. 104). The % from Mount Kinabalu also has the lateral lobes of the abdominal tergites decidedly larger with the outer margin acutely dentate. These differences are also true but much less striking in the & HT in RMNH (Fig. 103) and & in coll. CLC which are from the same two localities. Comments: Bragg (1995b) described this remarkable species from a single & in RMNH and originally included it in Nearchus Redtenbacher, 1908 which is a junior synonym of Phobaeticus Brunner v. Wattenwyl , 1907 (syn. nov.). This automatically transfers N. foliatus to Phobaeticus (comb. nov.). It is however remarkable for several characters which promptly distinguish it from all other members of the genus and entire tribe Pharnaciini, being the only representative with distinct spines on the mesonotum, large serrate lateral lobes on all abdominal tergites and numerous prominent, leaf-like appendages on the legs. Unfortunately the eggs are as yet unknown. Detailed comparison with other taxa of Phobaeticus have however, apart from the characters



143

mentioned above, not revealed any features which would justify a separate generic placement of Ph. foliatus. The shape and structure of the head, antennae and genitalia are very typical for Phobaeticus and place it close to the sympatric Ph. redtenbacheri (Dohrn, 1910). The % was subsequently described by Bragg (2001: 389) based on a specimen in RMNH from the same locality and source. The appendages of the abdominal tergites and prominent leaf-like expansions of the legs show remarkable convergence to the Neotropical genus Trychopeplus Shelford, 1909 (Type species: Trychopeplus multilobatus Redtenbacher, 1908) a member of Diapheromeridae: Diapheromerinae: Diapheromerini. Eggs unknown. Distribution (Fig. 380): Central and NE-Borneo, endemic. Kalimantan (Middle East-Borneo) and Sabah (Mount Kinabalu National Park 1500 m). Number of specimens examined: 4 TABLE 14. Measurements [mm] of Phobaeticus foliatus (Bragg, 1995) comb. nov.

&, HT (RMNH)

(RMNH)

%

Body

192.0

124.0


214.0

-

Head

12.0

5.0

Pronotum

7.0

5.0

Mesonotum

47.0

24.0

Metanotum

16.0

8.0

Median segment

12.0

10.0

Subgenital plate

41.0

-

Alae

-

49.0

Profemora

70.0

49.0

Mesofemora

45.5

30.0

Metafemora

55.5

39.0

Protibiae

96.0

64.0

Mesotibiae

52.0

34.0

Metatibiae

69.0–70.0

46.0

Antennae

> 9.0

70.0

Phobaeticus hypharpax (Westwood, 1859) (Figs. 77, 124, 177–178, 246–248, 275, 334–335, 339, 381) Phibalosoma hypharpax Westwood, 1859: 75, pl. 13: 6, 6a–b. HT, %: Holotype; Phibalosoma Hypharpax, Westw Ceylon (BMNH). Kaup, 1871a: 19, pl. 1: 11. [Description and illustration of egg] Kaup, 1871b: 37. [Description of &] Pharnacia hypharpax, Redtenbacher, 1908: 453. Phobaeticus hypharpax, Brock, 1996: 28. Otte & Brock, 2005: 268. Tirachoidea hypharpax, Brunner v. Wattenwyl, 1893: 83. Kirby, 1904a: 360. Cladoxerus acanthopus var. (?), de Haan, 1842: 132. Pharnacia spectabilis Redtenbacher, 1908: 453. HT, &: Coll. Br. v. W., Ceylon, Dr. Haly; det. Br. v. W. Pharnacia spec-


HENNEMANN & CONLE

tabilis; 17.226; Ph. spectabilis Redt. (NHMW, No. 863). syn. nov. Phobaeticus spectabilis, Brock, 1996: 30. Otte & Brock, 2005: 270. Further material: [14 &&, 7 %%, 1 & (nymph), 1 egg]: SRI LANKA: 1 %: Coll. Br. v. W., Ceylon, Dr. Haly; det. Br. v. W. Pharnacia hypharpax; 16.664 (NHMW, No. 864); 1 &: Coll. Br. v. W., Ceylon, Dr. Haly; det. Br. v. W. Pharnacia hypharpax; 17.228 (NHMW, No. 864); 1 %: Coll. Br. v. W. Ceylon, Makwood; 16.101 (NHMW, No. 864); 1 %, 1 & + 1 egg (ex ovipositor): Ceylon, Racuapura, III.1912, Pres. by Imp. Pur. Ent. Brit. Mus. 1922-468, leg. E.E. Green (BMNH); 1 &: Ceylon, pres. by Mr. Salvaga c/o Cox & co., Charing Cross, 191524. (BMNH); 1 & (nymph): Kandy, Sept.1914, leg., Whittle Coll., 1922-79 (BMNH); 1 &: Archabald Coll. B. M. 1926391, Karen Hill, THDG, Tracts, June 1923 (BMNH); 1 %: Ceylon (MHNG); 1 &: Ceylon (HLDH); 2 &&, 1 %: Ceylon, Worms (SMFM); 1 & (nymph): Ceylon, Worms; 8 (SMFM); 1 &: Ceylon 1912, A. Hansen (SMFM); 1 &: Ceylon, H.V. Hilprecht; Pharnacia spectabilis Redt. (ANSP); 1 %: Ceylon (ANSP), 1 &: Ceylon (SNMB). SOUTH INDIA: 1 &: S. India: Coorg, Mercara. 9-XI.1934, Maj. F.C. Fraser; Walking stick insect Ballquatrie, Mercara, Coorg, 9.XI.24 (BMNH). NO DATA: 1 &: (BMNH); 1 &, 1 %: (SMFM).

Diagnosis: && are characteristic for the sparse but very broad and prominent serrations of the anterodorsal carina of the profemora and unarmed medioventral carina of the meso- and metafemora. Closely related the second Sri Lankan species Ph. lobulatus (Carl, 1913) comb. nov. and Ph. sinetyi Brunner v. Wattenwyl , 1907 from the Madras Province of southeastern India. && differ from those of Ph. lobulatus by: the larger size; more rounded and less broadened lateral lobes of tergite VII; tergite VIII being shorter than IX and X combined (Fig. 246); unarmed mesopleurae; much longer and filiform antennae and different armature of the legs. From Ph. sinetyi it is distinguished by: the comparatively more slender body; longer median segment and smooth medioventral carina of the meso- and metafemora of both sexes, as well as the more prominent dorsal serrations and distinctly smaller teeth of the posteroventral carina of the profemora (Fig. 339), different armature of the mid and hind legs (Figs. 334–335) and broader lateral lobes of abdominal tergite VII of &&. %% clearly differ by: the dentate posteroventral carina of the profemora; presence of tegmina and well developed alae and more elongate, slender semi-tergites of the anal segment (Fig. 248). Eggs differ from those of Ph. sinetyi by the strongly sculptured, rugulose and wrinkled capsule surface and relatively larger micropylar plate (Figs. 177–178). Etymology: The derivation of the specific name hypharpax is doubtful. It may be a combination of the Latin “harpax” (= rapacious) and the Greek prefix “hypo-” (= under) meaning as much as “not rapacious”. Another possibility may be it was derived from Hypharpax MacLeay, 1825 a genus of Australasian Coleoptera (Carabidae: Harpalinae). Description: && (Fig. 124): Long to very long (body length 185.0–236.0 mm) and slender species (maximum body width ± 6.0 mm) with a short subgenital plate and prominent dorsal serrations on the profemora. Degree of leg armature variable (→ below). General colour of body and legs mid to dark ochracheous or greyish brown, either plain or to a variable degree furnished with numerous irregular paler markings and speckles; slightly shiny dorsally. Head of paler colour than rest of body. Meso- and metafemora with more or less distinct pale transverse bands and with a broad whitish or greyish band at the apex. Armature of legs dark brown to black. Eyes dark reddish brown. Antennae dark greyish brown, scapus often blackish dorsally. Head: Elongate, oval, almost 2x longer than wide, slightly narrowing towards the posterior and with a faint impressed coronal-line; vertex smooth and very gently convex. Posterior portion with two longitudinal lateral impressions and each side of the coronal-line. Between the eyes with a slightly raised, smooth, Cshaped area. Eyes small, moderately convex and with anterior margin slightly angled; length contained about 3.5x in that of cheek. Antennae rather short and reaching about 2/5 the length along mesonotum. Scapus REVISION OF PHARNACIINI


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slightly curved, dorsoventrally flattened and about 2.5x longer than wide. Pedicellus ¼ the length of scapus, cylindrical. Thorax: Pronotum shorter and distinctly narrower than head, 1.3x longer than wide, posterior margin broader than anterior margin. Median transverse depression distinct, widely V-shaped but not reaching lateral margins of segment. Mesothorax constricted anteriorly and 2.3–2.5x longer than head and pronotum combined. Mesonotum with a very faint slightly darker longitudinal median line (in most cases only visible in posterior portion of segment). Metanotum slightly more than 1/3 the length of mesonotum, about 2x longer than wide, rectangular. Meso- and metasternum with a very faint longitudinal median carina. Abdomen: Median segment slightly shorter than metanotum, anterior margin gently rounded. Segment II shorter than median segment and III–VII. III–VI very slightly increasing in length, and 2.6–2.8x longer than wide. Tergites V and VI with scale-like posteromedian tubercle (more prominent on V). VII about 5/6 the length of VI, lateral margins with a rounded posterior lobe of variable size. This lobe with the posterior margin ± acutely angulate to triangular and sometimes slightly projecting over posterior margin of the segment (Fig. 275). Praeopercular organ indistinct and represented by a pair of short, roundly raised carinae at posterior margin of sternum VII. Tergite VIII distinctly narrower than previous, almost 3x longer than wide, strongly convex and constricted medially. IX about half the length of VIII, indistinctly longer than wide, strongly convex. Anal segment 1.5x longer than IX with fine median carina and a widely triangular indention at posterior margin, outer angles acutely triangular (Fig. 247). Supraanal plate very small strongly keeled and rounded. Cerci small, oval in cross section, finely sethose and slightly projecting over anal segment. Gonapophyses yellowish brown, filiform, up-curving but not reaching apex of anal segment. Subgenital plate longitudinally keeled, boat-like and slightly projecting over posterior margin of anal segment (Fig. 246). Legs: All long and slender, profemora a little longer than combined length of pro- and mesonotum, mesofemora slightly shorter than mesothorax, metafemora projecting over posterior margin of abdominal segment IV and metatarsi just not reaching apex of abdomen. Anterodorsal carina of profemora with 8–10 prominent, broad and ± wave-like serrations which decrease in size towards the apex of femur (Fig. 339). Posterodorsal carina occasionally with 1–2 small, triangular teeth. Posteroventral carina with 5–7 small, pointed teeth. Anterodorsal carina of protibiae moderately raised and protruded into two ± well developed broad, triangular lobes in basal half and a smaller lobe-like tooth in apical half of tibia. Posterodorsal carina with a small apical spine, otherwise unarmed. Dorsal carinae of meso- and metafemora unarmed except for 2– 3 rather small teeth on anterior carina (Fig. 335); occasionally with a single more prominent, triangular tooth one third the way along posterior carina (Fig. 334). Medioventral carina smooth except for a single, minute subapical spine. Ventral carinae very sparsely and minutely dentate, the two basal teeth occasionally more prominent than the remaining (Fig. 334). Ventral carinae of meso- and metatibiae minutely dentate, the posteroventral carina sometimes with an enlarged, triangular tooth about 1/3 off the base of tibia. Usually with a very large, roundly triangular prae-medial lobe on posterodorsal carina and a distinct triangular tooth on anterodorsal carina (Fig. 334), but sometimes only represented by slightly enlarged teeth (Fig. 335). Posterodorsal carina forming a rounded, dentate lobe at the apex of tibia. Probasitarsus with a uniformly raised dorsal carina, as long as remaining segments combined. Second tarsomere ¼ the length of basitarsus with the dorsal carina raised and rounded. Meso- and metabasitarsus as long as remaining tarsomeres combined except claw and with an apically dentate dorsal carina, which gradually increases in height towards the apex. %% (Fig. 77): Medium-sized (body length 113.5–124.5 mm) and rather slender for the genus with long alae (46.0–59.0 mm). General colouration of body and legs ochracheous to greyish mid brown often with a slight green or yellow wash on mesonotum. Head with a ± distinct pale yellowish spot between the eyes, three distinct pale orange longitudinal lines on the vertex and a broad blackish postocular stripe along the cheeks. Tegmina greyish mid brown with the central portion darker brown and the anterior margin pale yellowish or white. Costal region of alae greyish bown with sepia veins, the anterior margin pale yellow or white in the basal portion. Anal region transparent grey with brown veins. Mid and hind legs with indistinct pale trans-


HENNEMANN & CONLE

verse bands. Bases of profemora pale orange. Leg armature dark brown. Eyes pale reddish brown. Dorsal surface of scapus dark brown to black, remaining antennomeres yellowish or greyish brown. Head: Generally as in && but eyes more prominent and projecting hemispherically; their length contained only a little more than 2x in that of cheeks. Antennae filiform and projecting over posterior margin of metanotum. Scapus almost 3x longer than wide and dorsoventrally compressed. Length of antennomeres increasing in length towards apex of antennae, all densely sethose. Thorax: Pronotum generally as in &&, distinctly shorter and narrower than head. Mesothorax 2.5x to almost 3x longer than head and pronotum combined. Meso- and metasternum with a very fine longitudinal median carina. Tegmina projecting over posterior margin of metanotum, elongate, narrowed towards the base and with a rounded central hump. Alae ± distinctly projecting over posterior margin of abdominal tergite IV. Abdomen: Median segment 2x longer than metanotum. Segments II–V roughly of equal length and width, about 5.5x longer than wide and shorter than median segment; all gently constricted medially. VI slightly shorter than previous and about 5x longer than wide. VII slightly shorter than IV, about 4x longer than wide and slightly broadened posteriorly. Tergite V with minute posteromedian tubercle. VIII 2/3 the length VII strongly broadening towards posterior margin and convex. IX a little longer than VIII, but distinctly narrower and constricted medially. Anal segment about as long as IX, laterally compressed and tectiform. Semi-tergites very elongate, strongly tapered, slender with a narrow apex and gently down-curving (Fig. 248); lateral surface with a conspicuous longitudinal impression in apical portion. Interior surface with a longitudinal median bulge apically which is densely covered with minute black teeth. Cerci elongate, in-curving and tapered towards the apex. Poculum reaching about 2/3 the way along tergite VIII, strongly convex, cup-like and with a very prominent and acute central hump. Legs: All very long and slender, profemora longer and mesofemora almost as long as head, pro- and mesothorax combined, metafemora reaching about half way along abdominal segment V and metatibiae slightly exceeding apex of abdomen. Armature generally as in && but slightly less prominent and lacking any considerably enlarged teeth or lobes. Dorsal carinae of meso- and metafemora smooth except for a very few minute teeth, two outer ventral carinae minutely serrate. Meso- and metatibiae occasionally with a single enlarged, triangular pre-medial tooth on posterodorsal carina. Basitarsi as in &&, but dorsal carina not considerably raised. Variation: While %% are quite constant, && show a great range of intraspecific variability concerning to the size (→ Table 16) and armature of the legs. A & contained in Brunner v. Wattenwyl’s collection in NHMW (collection No. 17.228) is remarkable for its very large size (body length 236.0 mm) and poorly developed leg armature (Fig. 335). An even less developed leg armature is seen in the HT of Pharnacia spectabilis Redtenbacher, which is considerably shorter (208.5 mm) but was possibly collected from the same locality (see comments below). The && in SMFM measure body lengths of only 185.0–197.5 mm and are a little more robust, which might be due to they were collected from a different locality possibly at higher altitude in the central highland of Sri Lanka. These && furthermore have the leg armature much more strongly developed, bearing prominent teeth on the meso- and metafemora and a large, roundly triangular pre-medial dorsal lobe on the meso- and metatibiae (Fig. 334). These are just represented by small triangular teeth in the two && in NHMW. Eggs (Figs. 177–178): The following description is based on a single fully developed egg extracted from the ovipositor of the & from Racuapura in BMNH (accession No. BM 1922-468). Capsule basically lens-shaped, laterally compressed, longer than high. General colouration of capsule and operculum dark yellowish brown, keel, capitulum and micropylar plate darker. Whole capsule surface minutely punctured and wrinkled, being covered with irregular raised carinae and tubercles. A dorsoventral keel begins and ends at the operculum and is only interrupted at the micropylar plate and notched at polararea. Lateral surfaces of keel with fine impressed radial ridges. Micropylar plate bilobed, forming an oval lobe on each side of the dorsal median line and surrounded by a bold black band. Central region of plate roughly granulose. Micropylar cup small, at polar end of plate and near centre of dorsal surface. Operculum slightly



147

convex, oval and with an irregularly shaped capitulum on a short stalk in the centre. Measurements [mm]: length (including capitulum) 5.5, length 4.9, width 2.9, height 3.8, length of micropylar plate 1.3. Comments: Westwood (1859: 75) originally described Phibalosoma hypharpax based on a single % from “Ceylon” in the BMNH and provided a very accurate illustration of the HT including details of the genitalia (Westwood, 1859: pl. 13: 6, 6a–b). Kaup (1871b: 37) provided a description of the so far unknown & and a description and illustration of the egg (1871a: 19, pl. 1: 11). The & recorded by Kaup (1871b) is present in Kaup’s collection in HLDH, but the appertaining egg is not traced. Examination of the & HT of Pharnacia spectabilis Redtenbacher, 1908 in NHMW and comparison with various && of Ph. hypharpax (Westwood) have proven Redtenbacher’s species to be a synonym (syn. nov.). The specimen is however remarkable for its poorly developed leg armature and lack of any conspicuously enlarged teeth or lobes on the mid and hind legs, thus representing an extreme and showing the wide range of intraspecific variability in this species. Redtenbacher (1908: 453) recorded a body length of 200 mm for the HT but the exact length is 208.5 mm. Furthermore, the specimen was obviously collected along with one % and the giant 236.0 mm & of Ph. hypharpax in NHMW, as it bears exactly the same data and labels stating “Ceylon, Dr. Haly”. Ph. hypharpax (Westwood) appears to be quite widely distributed in the tropical lowland and mountainous regions of southern Sri Lanka but absent in the northern portion of the island. Although this species is represented in several European museum collections almost all specimens lack a more precise locality than “Ceylon”. Only one specimen has so far been recorded from SE-India. && in particular show a considerable range of intraspecific variation, concerning to the size, colouration and armature of the legs (see above). Distribution (Fig. 381): Sri Lanka (Racuapura; Kandy & Karen Hill) and Southeast India (Mercara). Number of specimens examined: 24 TABLE 15. Measurements [mm] of Phobaeticus hypharpax (Westwood, 1859).

%, HT

%%

&, HT of Ph. spectabilis (NHMW)

&&

(BMNH) Body

113.5

119.5–124.5

208.5

185.0–236.0

Pronotum

3.8

3.8–4.0

8.5

6.5–9.0

Mesonotum

20.5

23.5–25.0

43.0

36.5–49.5

Metanotum

6.0

5.9–6.8

18.5

14.0–18.5

Median segment

11.0

10.8–12.0

12.0

11.0–15.0

Alae

46.0

54.0–59.0

-

-

Profemora

35.0

39.5–40.5

-

45.5–60.0

Mesofemora

26.5

31.0–35.5

44.0

33.5–45.5

Metafemora

31.5

37.0–42.5

50.0

40.5–52.0

Protibiae

41.7

-

-

-

Mesotibiae

35.8

-

-

-

Metatibiae

34.6

-

-

-

Antennae

> 28.0

33.5–37.0

> 15.0

32.5


HENNEMANN & CONLE

FIGURES 115-119: Phobaeticus spp. 115. Phobaeticus kirbyi Brunner v. Wattenwyl , 1907, % NE-Sarawak, Batu Niah (coll. FH, No. 0160-8) 116. Phobaeticus chani Bragg spec. nov., & HT, Sabah, Penampang District, Ulu Moyog (BMNH) 117. Phobaeticus chani Bragg spec. nov., % PT, Sabah, Sipitang (BMNH) 118. Phobaeticus rex (Günther, 1928), & HT, Sumatra, Redjang-Lebong (MNHU) 119. Phobaeticus philippinicus (Hennemann & Conle, 1997), & Philippines, Mindoro Island (coll. OC)



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Phobaeticus ingens (Redtenbacher, 1908) (Figs. 101–102, 227–228, 271, 362, 373) Pharnacia ingens Redtenbacher, 1908: 453. LT [by present designation], &: Meppadi, Malabar, Mr. Atzenwiller; Pharnacia ingens Redtb. Type (MHNG, No. 69) [Not: PLT, &: Upper-Tenasserim, Moolai to Moolat, 4000–6000 feet (presumably in NZSI). Recorded by Wood-Mason (1877a: 161) as “Phibalosoma acanthopus Burmeister, 1839”; = Phryganistria grandis Rehn, 1906] Phobaeticus ingens, Brock, 1996: 29. Otte & Brock, 2005: 269. [Not: Phibalosoma acanthopus, Wood-Mason, 1877a: 161. Misidentification = Phryganistria grandis Rehn, 1906] Further material [5 %%]: MYANMAR: 2 %%: N-Myanmar, Tenasserim Highland, 18.3.–19.IV.1996, leg. Lehmann (coll. FH, No’s 0251-1 & 2); 3 mar, Tenasserim, leg. Lehmann V.1995–20.IV.1996 (coll. OC).

%%: Myan-

Diagnosis: This striking species is well characterised by the rather broad and robust body, heart-shaped abdominal tergite VII (Fig. 271), characteristic leg armature, distinct white markings on the complete body and legs and praeopercular organ of && which consists of only one central spine-like appendage (two in all other species in the genus, Fig. 362). %% resemble those of the Bornean Ph. chani Bragg spec. nov. by differ by the smaller size, slightly more robust body, lack of a dark postocular stripe and comparatively broader semi-tergites of the anal segment (Fig. 228). Etymology: The specific name ingens (lat. = huge, gigantic) refers to the impressive size of the && of this beautiful species. Description: && (Fig. 101): Very large (body length 218.0 mm) and moderately robust species (maximum body width 12.0 mm) with very characteristic leg armature. General colouration of body and legs yellowish to greyish mid brown with great portions of dorsal surfaces of head, meso-, metanotum, median segment and single abdominal tergites white. Ventral surface of body and lateral surfaces of head brown with whitish speckles. Vertex with a bold wedge-shaped white marking. Mesonotum brown at anterior margin with a bold lateral brown patch near posterior margin and irregular brown patterns on lateral margins. Metanotum with brown lateral patterns. Median segment with an elongate brown patch along lateral margins. Tergite II with a bold )(-shaped white marking. Great portions of III and VI white with brown speckles laterally. IV and V brown with a pair of bold white patches near posterior margin; these more elongate and indistinct on V. Posterior portion of VII with two bold, white transverse bands, which are connected at their apices. VIII with a bold white, longitudinal median stripe. IX and anal segment white except for a small irregular brown anterolateral marking. Subgenital plate brown with a white lateral marking sub-basally. Legs more or less distinctly marbled with white. Armature of meso- and metafemora dark drab. Eyes reddish brown. Antennae reddish mid brown. Head: Oval, 1.6x longer than wide, vertex very gently rounded, with an impression transverse furrow between the eyes. Eyes of moderate size, circular and slightly convex; their length contained 3.5x in that of cheek. Antennae at least projecting over pronotum (broken in the LT). Scapus dorsoventrally flattened, 2.5x longer than wide and gently narrowed towards the base. Pedicellus distinctly shorter than scapus, round in cross-section. Thorax: Pronotum shorter and slightly narrower than head, about 1.2x longer than wide and with anterior margin narrower than posterior margin. Median transverse depression short, straight and not reaching lateral margins of segment. Entire dorsal surface with an impressed longitudinal median line. Mesothorax decidedly constricted at the anterior and almost 2.5x longer than head and pronotum combined. Mesonotum with a slightly raised longitudinal median line. Metanotum parallel-sided and about 2.6x longer than wide. Mesopleurae set with a longitudinal row of minute but acute granules. Mesosternum with a very faint and


HENNEMANN & CONLE

irregular longitudinal median line, metasternum smooth. Abdomen: Median segment a little more than half the length of metanotum, 1.3x longer than wide, parallel-sided. Segments II–VI slightly increasing in length, in average 1.5x longer than wide, parallel-sided. Tergites IV and VI each with a rounded, black posteromedian tubercle, V with a rounded, scale-like lobe. VII strongly dilated into a large, rounded lateral lobe, which projects by more than half the body width, and in dorsal aspect gives the segment a heart-shaped appearance (Fig. 271). Praeopercular organ formed by a single prominent, dorsoventrally flattened, roughly triangular median appendage at posterior margin of sternum VII (Fig. 362). VIII slightly longer but distinctly narrower than VII, 3.5x longer than wide, strongly convex and gently constricted medially. IX about half the length of VIII, indistinctly longer than wide, strongly convex, rectangular. Anal segment longer than IX, flattened towards the apex and with a fine longitudinal median carina, posterior margin with a very shallow concave excavation, outer angles roundly triangular. Supraanal plate very small, rounded and keeled. Cerci small and oval in cross-section. Subgenital plate strongly keeled, boatshaped and tapered towards a blunt apex, which slightly projects over posterior margin of anal segment (Fig. 227). Legs: All rather long and robust with prominent, characteristic lobes and serrations. Profemora about as long as pro-, and mesonotum combined, mesofemora slightly shorter than mesothorax, metafemora slightly projecting over posterior margin of abdominal segment IV and hind tarsi projecting over apex of abdomen. Meso- and metafemora very gently down-curving. Anterodorsal carina of profemora with 12–13 broad triangular serrations, posteroventral carina with a similar number of long and slender spines. Anterodorsal carina of protibiae distinctly raised, slightly wave-like and protruded into a prominent, triangular median tooth; a further, slightly enlarged tooth near the base and several towards the apex of tibia. Posteroventral carina sparingly serrate. Two outer ventral carinae of mesofemora armed with numerous rather distinct, straight and pointed spines, these considerably enlarged in basal portion of posteroventral carina. Anterodorsal carina smooth except for 3–4 minute teeth. Posterodorsal carina with two slightly enlarged teeth about ¼ off the base and another two slightly enlarged teeth near apex. Metafemora with the two outer ventral carinae minutely and sparingly serrate, dorsal carinae smooth except for a slightly enlarged tooth about ¼ off the base posterodorsal carina. Medioventral carina of meso- and metafemora with 8–12 pointed spines. Posterodorsal carina of mesotibiae protruded into a very prominent, triangular and foliaceous median lobe, a much smaller, triangular lobe about ¾ the way along tibia and a rounded, dentate lobe apically. Medioventral carina very sparingly dentate and with a single enlarged tooth about 1/3 off the base. Posterodorsal carina of metatibiae with a large, triangular median lobe and a rounded, dentate apical lobe. Medioventral carina very sparingly and minutely serrate. Probasitarsus elongate, with the dorsal carina uniformly elevated and unarmed (remaining tarsomeres missing in the LT). Dorsal carina of meso- and metabasitarsus gradually raised towards the apex; as long as following three tarsomeres combined. %% (Fig. 102): Medium-sized (body length 117.0–119.2 mm) and rather slender for the genus with moderately long alae (55.0–57.5 mm). General colouration of body and legs dark brown, abdomen greyish brown. Mesopleurae with a whitish posterior spot and lateral margins of tergite IX with a bold white longitudinal band. Abdominal sternites irregularly set with indistinct whitish speckles. Cerci grey or yellowish brown. Bases of profemora straw, those of meso- and metafemora and basitarsi dull orange. Armature of legs black. Antennae dark reddish brown. Tegmina and costal region of alae greyish brown (tegmina slightly darker), the latter becoming transparent with dark brown veins towards the interior. Anal region of alae transparent grey with blackish veins. Anterior margin of tegmina and basal half of alae with a longitudinal yellowish white band; this followd by a fine, dark brown interior longitudinal line on tegmina. Eyes dark brown. Head: Generally as in &&. Between the eyes with a short, curved transverse impression. Eyes very prominent and projecting hemispherically; their length contained less than 2.5x in that of cheek. Antennae projecting over posterior margin of metanotum, with 27–28 segments. Scapus oval in cross-section, slightly narrowed towards the base and about 2.5x longer than wide. Pedicellus ¼ the length of scapus, round in cross-



151

section. Following antennomeres distinctly increasing in length, the seven apical ones decreasing. Terminal antennomere very elongate, as long as four previous combined. Thorax: Pronotum generally as in && but relatively longer. Mesothorax 2.5x longer than head and pronotum combined, slightly broadened posteriorly. Mesonotum set with a very few indistinct granules and minutely rugulose along the lateral margins. Mesosternum with a faint longitudinal median carina; metasternum smooth. Metanotum about ¼ the length of mesonotum. Tegmina elongate, oval, strongly tapered in basal half and with a well decided, conical central hump. Alae reaching to posterior margin of abdominal tergite V. Abdomen: Median segment about 3.5x longer than metanotum, 4x longer than wide and slightly tapered towards the posterior. Segments II–VI slightly increasing in length, II 4.5x, VI 5.5x longer than wide, all gently constricted medially. Tergite VII shorter than VI, about 5x longer than wide and slightly expanded posteriorly. V–VII each with a small posteromedian tubercle. VIII 2/3 the length of VII, 2.3x longer than wide, broadening towards posterior margin and strongly convex; broadest segment. IX slightly shorter than VIII, strongly convex and constricted medially; anterior margin broader than posterior margin. Anal segment as long as IX. Semi-tergites very broad, indistinctly longer than high with the apex strongly angled and roundly truncate, lower apical angle acute (Fig. 228). Interior surface densely set with small black teeth. Cerci obtuse, rather elongate, oval in cross-section, gently in-curving and slightly projecting over apex of anal segment. Poculum strongly convex, cup-like and reaching about 2/3 the way along tergite IX; posterior portion with a slight longitudinal median carina. Legs: All moderately long and slender, profemora about as long as head, pro-, and mesonotum combined, mesofemora slightly longer than pro-, and mesonotum combined, metafemora reaching about half way along abdominal segment V and hind tarsi slightly projecting over tip of abdomen. Armature similar as in && but considerably less developed. Anterodorsal carina of profemora with 7–14 very prominent black, triangular serrations; posteroventral carina with 8–15 smaller black teeth. Anterodorsal carina of protibiae smooth, except for 1–3 flat, broad lobes in great but equal distance to another and a few minor basal serrations; the median lobe largest and occasionally the proximal and apical ones very indistinct. Posteroventral carina sparingly and minutely serrate. All carinae of meso- and metafemora serrate, more sparsely but distinct on dorsal carinae. Medioventral carina set with 11–14 small spines. Two outer ventral carinae of mesofemora very slightly dilated basally with teeth more prominent in this portion. All carinae of meso- and metatibiae minutely serrate. Posterodorsal carina with a prominent, triangular median lobe and raised into a prominently rounded and dentate lobe at the apex. Probasitarsus with a slightly raised dorsal carina and almost 1.5x longer than remaining tarsomeres combined. Meso- and metabasitarsus slightly longer than combined length of remaining tarsomeres, dorsal carina gently rounded and occasionally with one or two minute teeth. Comments: Redtenbacher (1908: 453) described Pharnacia ingens based on a & from the Malabar Coast, southwestern India in MHNG and a & from Upper Tenasserim, Myanmar. The latter & is a specimen that had erroneously been recorded as “Phibalosoma acanthopus Burmeister, 1839” by Wood-Mason (1877a: 161). Redtenbacher recognized Wood-Mason’s misidentification and decided the & from Myanmar to be conspecific to the & in MHNG, which is however certainly not the case. The whereabouts of Wood-Mason’s specimen have so far remained unknown although it is persumed to be traced in NZSI. Inquiries have however not received any reply. Therefore, the & from India in MHNG is here selected as the LT of Ph. ingens since it is the only specimen which Redtenbacher had examined himself. Wood-Mason’s specimen from Myanmar, the PLT, most certainly belongs in Phryganistria Stål, 1875 (→ Phryganistria grandis Rehn, 1906). Redtenbacher (1908: 453) cited body lengths of 208.0 and 260.0 mm, the latter referring to the PLT. The presence of a praeopercular organ in && confirms the current placement in Phobaeticus Brunner v. Wattenwyl, 1907. The organ itself however is of a structure rather different from other Phobaeticus-species, consisting of only one triangular, lobe-like appendage at posterior margin of abdominal sternum VII. This feature readily distinguishes && from those of all other species, all of which have two appendages. Examination of five %% from Myanmar in the authors collections and comparison with the LT of Ph.


HENNEMANN & CONLE

ingens has shown these widely correspond in leg armature and shape of the head and thorax, thus might represent the opposite sex. However, without seeing large series of both sexes from the same locality, these %% are here assigned with doubt only. The short median segment of && would suppose the corresponding %% to be apterous rather than having fully developed alae. Furthermore, there is a distance of more than 1500 kilometres between the type-locality (Malabar Coast, SE.India) and the Highland of Tenasserim in northern Myanmar. Also, as can be seen in other species of the genus size and armature of the legs are subject to strong intraspecific variability and concerns to both sexes, which is an additional factor that renders the correct assignment of sexes more difficult. Eggs unknown. Distribution (Fig. 373): Southwest India (Malabar Coast: Meppadi) and Northern Myanmar (Highland of Tenasserim). Number of specimens examined: 6 TABLE 16. Measurements [mm] of Phobaeticus ingens (Redtenbacher, 1908)

&, LT (MHNG)

%%

Body

218.0

117.0–119.2

Head

9.4

5.4–5.6

Pronotum

7.0

3.2–3.8

Mesonotum

43.8

20.8–21.1

Metanotum

22.5

5.8–6.1

Median segment

12.0

12.4–12.7

Tegmina

-

9.0–9.2

Alae

-

55.0–57.5

Profemora

50.5

30.0–34.2

Mesofemora

37.3

27.2–27.7

Metafemora

47.7

33.6–33.8

Protibiae

56.0

35.9–43.1

Mesotibiae

49.0

25.7–29.0

Metatibiae

54.0

32.5–35.4

Antennae

> 15.0

41.0–45.0

(coll. FH)

Phobaeticus kirbyi Brunner v. Wattenwyl , 1907 (Figs. 114–115, 171–172, 237–239, 345–346, 361, 382, 418) Phobaeticus kirbyi Brunner v. Wattenwyl , 1907: 185. HT, &: Borneo (BMNH). Brock, 1996: 29. Seow-Choen, 1998a: 41. Seow-Choen, 1998b: 89. Otte & Brock, 2005: 269. Pharnacia kirbyi, Bragg, 1995a: 27, fig. 1. [Description and illustration of egg] Baculolonga kirbyi, Hennemann & Conle, 1997: 348. Bragg, 2001: 379, figs. 142 (%, &) & 143 (egg). Pharnacia pilicornis Redtenbacher, 1908: 455. HT, %: Brunei, Borneo, Staudinger; Collectio Br. v. W.; det. Br. v. W. Pharnacia pilicornis; 20.854 (NHMW, No. 865). [Synonymised with Pharnacia sagitta by Günther, 1932: 315] Phobaeticus pilicornis, Brock, 1998a: 50. [As synonym of Ph. kirbyi] Otte & Brock, 2005: 269. [As synonym of Ph. kirbyi]



153

Pharnacia sagitta Redtenbacher, 1908: 454, pl. 22: 2. HT, & (subadult nymph): Coll. Br. v. W., Borneo, Doben; det. Br. v. W. Pharnacia sagitta; 18.444; 65 (NHMW, No. 866). [Synonymised by Bragg, 1995a: 27] Günther, 1932a: 315. Günther, 1943: 155. Phobaeticus sagitta, Brock, 1998a: 55. [As synonym of Ph. kirbyi] Otte & Brock, 2005: 269. [As synonym of Ph. kirbyi] Pharnacia maxima, Kirby, 1904a: 359. [Misidentification relating to the HT of Ph. kirbyi Brunner von Wattenwyl]] Pharnacia serratipes, Kirby, 1896: 448 & 450. [Misidentification relating to the HT of Ph. kirbyi Brunner von Wattenwyl] Redtenbacher, 1908: 455 (in part - only the specimen of Kirby, 1896: 448, the HT of Ph. kirbyi Brunner v. Wattenwyl). Hypermergethes herberti Brunner v. Wattenwyl, in litt. %: Borneo; Hypermergethes herberti Br. det. (ETHZ). Further material: [22 &&, 20 %%, 8 nymphs, 3 eggs]: BORNEO, BRUNEI: 1 % (penultimate instar nymph), 1 & (penultimate instar nymph): Brunei, Seria, Badas, leg. Hennemann & Herbert VIII. 1994 (coll. FH, No’s 0160-1 & 2); 1 %, 1 &: Brunei, Badas, leg. M. Herbert IV.1995 (coll. MH, No’s 10-4 & 10-5); 1 &, 1 %: N-Borneo, Brunei, Capt. Waterstraat, Van der Poll, Coll. Bought, Janson 1909-463 & 464 (OXUM). BORNEO, SARAWAK: 4 %%, 4 &&, 3 eggs: NO-Sarawak, Niah National Park, leg. Hennemann & Conle 2.-3. VIII. 1996 (coll. FH, No’s 0160-4 to 11 & E); 1 & (penultimate instar nymph): NW-Sarawak, Mt. Serapi, ca. 200 m, leg. Hennemann & Conle VII.1996 (coll. FH, No’s 0160-3); 1 &, 3 %%: Sarawak, Batu Niah, XII.1980, A. Harman leg., BM 1982-387 (BMNH). BORNEO, SABAH: 4 &&, 3 && (nymphs): Brit. Nord-Borneo, Kinabalu-Gebirge, 16. XII. 1912 (SMNS); 1 %, 1 &: Sandakan, C.V.Creagh, 96-197; 1 & (nymph): B.N. Borneo, nr. Kinabalu, Kabayau, 600m, 9.V.1929 from H.M. Pendlebury collection (RMNH); 1 & (nymph): Murmet-Territory, N-Borneo, Higgins leg. 1871 (RMNH); 1 &: Borneo Exped. Mr. Morvet, Boeleit, Dadop. (RMNH), 1 %: No. 193 (k / 52), XL.VIII. (RMNH); 1 &: Flalek Papan, Kamp., Juli 1912 (RMNH), 1 %: B.N. Borneo, Mt. Kinabalu, Kian, 3000 ft., April 1929 (RMNH); 1 &: NO-Borneo, S.-Peleben, VII.-VIII.1935, Baron Plessen leg. (RMNH); 3 %%: Brit. N-Borneo, Darvel-Bay, Pagel S.V. leg., det. K. Günther „Phasgania pilicornis“ (MNHU); 1 %: N. Borneo, Kina Balu 1500 m, leg. Rolle 25. VI.1904 (ZMUH); 1 &, 1 %: Nord-Borneo, Waterstraat (ETHZ); 2 %%: Labuan, Brit. N. Borneo, Pharnacia pilicornis Redt. %, Det. Hebard 1917, Hebard Cln. (ANSP); 1 &: Labuan, Brit. N. Borneo; Phobaeticus n. sp. I, Det. Hebard 19; Hebard Cln. (ANSP); 1 &: Labuan, Brit. N. Borneo; Pharnacia pilicornis Redt. &, Det. Hebard 1917; Hebard Cln. (ANSP). BORNEO, KALIMANTAN: 1 &: Borneo, Exped. Dr. Nieuwenhuis, Long-Bloe, XII.1898 (RMNH); 1 &: Borneo Exped. Mt. Morvet, Bukit Dadap [data difficult to read] (RMNH); 1 &: E-Borneo, 125m, Tabang, 21.VIII.1956; det. P.E.Bragg (RMNH). BORNEO: 1 %: Borneo, Hypermergethes herberti Br. det. (ETHZ); 1 %, 1 &: Borneo, Phryganistria serratipes (MNHN, coll. Finot, Box-No. 280); 1 %: Wattan-Borneo, Purchased and pres. ´73 by Mrs. F.W. Hope (OXUM); 1 &: Borneo (ZSMC). NO DATA: 2 &&: (MNCN).

Diagnosis: Well characterised and distinguished from other members of the genus by the strongly globose head of both sexes (Fig. 307–308), very short subgenital plate of && which stays clearly beyond the apex of the abdomen (Fig. 237) and characteristic colouration especially of the head of %% (Fig. 308) which have the posteroventral carina of the profemora entirely unarmed (Fig. 346). The eggs clearly distinguish Ph. kirbyi from all remaining members of the genus by the distinct radial ridges of the dorsoventral keel of the capsule (Figs. 171–172). Closely related to Ph. serratipes (Gray, 1835) from Peninsular Malaysia, Singapore and Sumatra, and the three other Bornean species Ph. redtenbacheri (Dohrn, 1910), Ph. mjoebergi (Günther, 1935) and Ph. chani Bragg spec. nov.. From Ph. chani Bragg spec. nov. it furthermore differs by: the smaller size; relatively more robust body; less elongate body segments and lack of any enlarged teeth on the mesofemora of &&, as well as the considerably shorter alae and slender, finger-like semi-tergites of the %% (Fig. 239). && of Ph. redten-


HENNEMANN & CONLE

bacheri (Dohrn) and Ph. mjoebergi (Günther) at once differ from Ph. kirbyi by the strongly elongated, lanceolate subgenital plate, which considerably extends over the apex of the abdomen. %% of Ph. kirbyi are similar to those of Ph. redtenbacheri (Dohrn) but can in addition to the characteristic features mentioned above be distinguished by: the longer alae and apically tapered and acute semi-tergites of the anal segment. Etymology: Dedicated to William Forsell Kirby (1844–1912), a well known English entomologist and folklorist, who first recorded this species as “Pharnacia serratipes”(Kirby, 1896: 448). Description: The colouration is described from colour photos of live adult insects taken at Batu Niah (NE-Sarawak) in August 1996. && (Fig. 114, 418): Long to very long (body length 187.0–317.0 mm) and slender species (average body width 4.0–5.0 mm) with a very short subgenital plate. Variable in size, colouration and leg armature. General colouration of body and legs mid to dull green or mid to dark brown, either uniformly coloured or with darker and paler speckles. Occasionally brown specimens occur which have a prominent white marking on the vertex, a bold white longitudinal median stripe on the pronotum, bold white patches and markings on the entire body, or a white longitudinal median line running along the full length of the dorsal body surface; in such specimens the meso- and metafemora with a broad whitish transverse sub-basal band. Armature of the legs dull red with black points. Eyes reddish brown. Antennae dark reddish brown and black ventrally, two basal segments of paler colour. Head (Fig. 307): Strongly globose, indistinctly longer than wide and conspicuously broader than thorax; vertex convex. A slightly depressed area between the bases of the antennae. Eyes rather small, cylindrical and strongly convex; their length contained about 2.3x in that of cheeks. Antennae filiform and reaching to posterior margin of mesonotum, consisting of about 35 antennomeres. Scapus slender, 3x longer than wide, dorsoventrally flattened and narrowed towards the base. Pedicellus 1/3 the length of scapus, cylindrical. Following antennomeres first increasing then decreasing in length towards apex of antennae. Thorax: Pronotum 1.7x longer than wide, shorter and distinctly narrower than head, the posterior portion somewhat broader than the anterior. Median transverse depression distinct very gently curved, but not reaching lateral margins of segment. Mesothorax about 3.6x longer than head and pronotum combined, mesonotum parallel-sided. Metanotum 4.5x longer than wide, rectangular and about 1/3 the length of mesonotum. Meso and metasternum simple. Abdomen: Median segment slightly shorter than metanotum and slightly widened in posterior portion. Segments II–VI slightly increasing in length, II less than 4x, VI almost 5x longer than wide. VII slightly shorter than VI and very gently widened posteriorly. Tergite V with a scale-like posteromedian tubercle. Praeopercular organ formed by two prominent, sickle-shaped lobes near posterior margin of sternum VII (Fig. 361). Tergites VIII–X narrower than previous, VIII about half as long as VII and slightly narrowed in anterior half, IX half as long as VIII and indistinctly longer than wide. Anal segment conspicuously longer than IX, with a fine longitudinal median carina and a deep triangular posteromedian excavation; outer angles considerably deflexed and roughly triangular (Fig. 238). Supraanal plate very short, rounded. Subgenital plate longitudinally keeled, boat-shaped but short and only reaching half way along anal segment (Fig. 237). Legs: All very long and slender but very variable in length. Profemora about as long as head, pro- and mesothorax combined, mesofemora slightly shorter than mesonotum, metafemora reaching about 1/3 the way along abdominal segment V and metatibiae very slightly projecting over apex of abdomen. Anterodorsal carina of profemora with 16–28 prominent and acutely triangular serrations (Fig. 345), posterodorsal carina very indistinct, unarmed. Armature of the posteroventral carina variable, either completely smooth, set with 3.4 minute teeth, or sub-basally with a ± prominent triangular lobe-like tooth and 3–5 smaller teeth along the length of femur. All carinae of mid and hind legs quite regularly set with small, pointed teeth (smaller on dorsal carinae). Medioventral carina very indistinct and set with a few minute spines in the median section of femur. Posterodorsal carina of meso- and metatibiae forming a slight rounded lobe at the apex; occasionally with a slightly enlarged tooth some 2/5 off the base, and rarely with another enlarged tooth 1/5 off the apex.



155

Probasitarsus very elongate and longer than remaining tarsomeres combined, dorsal carina slightly elevated, all carinae smooth. Meso- and metabasitarsi a little longer than following three tarsomeres combined, the dorsal carina slightly but gradually raised towards the apex, all carinae minutely dentate. %% (Fig. 115, 418): Moderately sized to large (body length 132.0–166.0 mm) and very slender for the genus (2.0–3.0 mm) with shortened alae (43.0–49.5 mm) and a very characteristic colouration. Body and legs mid green (often greenish brown or pale brown when preserved). Head bright yellow with a distinct, broad black lateral marking on the cheeks. Tegmina and costal region of alae pale greenish brown, anterior margin broadly white. Anal region of alae transparent grey with brown veins. Dorsal serrations of the profemora orange or red with the posterior edges black; other leg armature black. Eyes yellow to ochre. Antennae black, except for a green to greenish brown scapus and pedicellus. Head (Fig. 308): Strongly globose and generally as in &&, but eyes much more prominent and projecting hemispherically; their length contained only about 1.5x in that of cheeks. Antennae projecting over posterior margin of median segment. Scapus dorsoventrally flattened, gently narrowed towards the base and almost 4x longer than wide. Scapus cylindrical and less than 1/3 the length of scapus. Thorax: Pronotum 1.8x longer than wide, with a distinct median transverse depression and the anterior 1/ 3 of segment conspicuously constricted. Mesothorax about 3.5x longer than head and pronotum combined. Mesosternum with a distinct longitudinal median keel (very faint on metasternum). Metanotum short, only half the length of median segment. Tegmina slender, strongly narrowed in basal half and with a faint central hump. Alae slightly projecting over posterior margin of abdominal tergite III Abdomen: Median segment 6x longer than wide. Abdominal segments II–VI 8x and VII 6x longer than wide. Posterior margins of tergites VI–V slightly raised, on V forming a transverse, scale-like posteromedian hump. VIII about 2x longer than wide, about half the length of VII and slightly widened towards the posterior. IX slightly shorter than VIII and constricted medially. Anal segment almost as long as VII, strongly tectiform and laterally compressed. Semi-tergites elongate, tapered towards a slender apex and slightly down-curving (Fig. 239). Interior surfaces apically with a blunt longitudinal keel set with several dark brown teeth. Cerci long, slender and in-curving, the apex distinctly tapered. Poculum strongly convex, cup-like and with a blunt, conical central hump. Legs: All very long and slender, profemora longer than head, pro- and mesothorax combined, mesofemora a little longer than pro- and mesonotum combined, metafemora reaching to posterior margin of abdominal segment V and metatibiae projecting considerably over apex of abdomen. Anterodorsal carina of profemora armed with ± 20 prominent, acutely triangular teeth (Fig. 346), posteroventral carina unarmed (rarely with 2– 3 very minute teeth). All carinae of mid and hind legs densely armed with minute pointed teeth (more sparsely on dorsal carinae). Ventral surfaces of meso- and metafemora smooth. Probasitarsus very elongate, unarmed and conspicuously longer than remaining tarsomeres combined. Meso- and metabasitarsus a little longer than remaining tarsomeres and with all carinae minutely serrate. Eggs (Figs. 171–172): The following description is based on three eggs laid by wild specimens from Batu Niah, NE-Sarawak. Due to Bragg (1995a & 2001) had a sample of 28 eggs at hand for examination, his measurements are given in addition to the ones taken from the three examples in the first author’s collection (coll. FH, No. 0160-E). Capsule basically lens-shaped, laterally compressed, slightly longer than high. General colouration of capsule and operculum creamish pale to mid brown. Whole capsule surrounded by a prominent dorsoventral keel, beginning and ending at the operculum and only interrupted at polar-area and the micropylar plate. Polar impression distinct. Keel with an irregular outer margin and laterally furnished with distinct, impressed radial ridges, each basing in a deep circular impression where they join with the main capsule. Micropylar plate bilobed, strongly constricted medially and forming an oval lobe on each side of the dorsal median line. Outer margin with a fine reddish brown line. Micropylar cup small and placed at polar end of plate. Operculum slightly convex, oval and with an irregularly knob-shaped, black capitulum in the centre.


HENNEMANN & CONLE

Measurements [mm]: Length (including capitulum) 6.0–6.4, length 5.3–5.6, height 4.4–4.8, width 3.1– 3.2, length of micropylar plate 1.9–2.1. Measurements according to Bragg (1995b: 29 & 2001: 383) [mm]: Length 5.26–5.84, height 4.68–5.06, width 3.12–3.31, length of capitulum 0.71–0.84 (dehydrated). Comments: Kirby (1896: 450) described the presumed & of Ph. serratipes (Gray, 1835) based on a tentatively identified specimen from Borneo in BMNH. For this specimen Kirby quoted a body length of “1 foot 10 lines “ which converts to 328 mm. This was given as 330 mm by Brunner v. Wattenwyl (1907: 185) and subsequent authors (Bragg, 1995a: 28, 2001: 383). Kirby (1896: 448) referred to a second specimens and stated “Another specimen, from a doubtful locality, measures only 10 ½ inches in total length”. This latter specimen is not traced in BMNH. Kirby (1896: 448) already commented that he was not certain whether his Bornean specimens really represented Gray’s species and subsequently (Kirby, 1904a: 359) decided them to be Pharnacia maxima (Bates, 1865) which however is a junior synonym of Ph. serratipes (Gray) (syn. nov.). Brunner v. Wattenwyl (1907: 185) recognised that Kirby’s Bornean specimens were a distinct species and based on these described Phobaeticus kirbyi. Consequently, and as the second specimen is not traced the unique & in BMNH must be considered the HT of Brunner v. Wattenwyl’s species (Bragg 1995a: 28). Redtenbacher (1908) described two species in the genus Pharnacia Stål, 1877 which are junior synonyms of Ph. kirbyi. Pharnacia pilicornis Redtenbacher, 1908 was described from a unique % from Brunei in NHMW and synonymised with Pharnacia sagitta Redtenbacher, 1908 by Günther (1932: 315). Pharnacia sagitta Redtenbacher, 1908 was described from a single subadult & nymph and synonymised with Ph. kirbyi by Bragg (1995a: 27). Bragg (1995a, 2001: 379) provided detailed descriptions of both sexes and the eggs, information on the synonymy and distribution of Ph. kirbyi as well as illustrations of the %, genitalia and egg. Bragg stated the length of the body and extremities to be considerably variable in specimens from different localities in Borneo. The && from Mt. Kinabalu in SMNS and specimens in ANSP are remarkably shorter than all other recorded material. For 95 years Ph. kirbyi was regarded “the longest insect in the world”, based on Kirby’s specimen in BMNH which had erroneously been identified as Pharnacia serratipes (Gray, 1835) with a quoted body length of 330 mm. Re-measuring of the specimen has shown the actual body length to be only 317.0 mm. The overall length is 499 mm if the legs are included. Bragg (1995a: 28, 2001: 383) listed an even longer specimen with an overall length of 546 mm (body length 283 mm) from Kuala Belalong (Brunei: Temburong District). As can be seen from the numerous records, Ph. kirbyi is widely distributed in Borneo and quite abundant in certain localities. Bragg (1995a: 27 & 2001: 384) reported the species to be exceptionally common along the path leading from the Park Head Quarters to the Great Cave in Niah National Park (NE-Sarawak), with more than 40 adult specimens and numerous mating pairs encountered in only one night during August 1992. The authors encountered similar numbers at the same locality in August 1996. Distribution (Fig. 382): Borneo, endemic. Sabah (Sandakan [Bragg, 2001: 379]; Sepilok [Bragg, 2001: 379]; Ulu Dusun [Bragg, 2001: 379]; Labuan; Sepitang; Kabayau 600 m [Bragg, 2001: 379]; Boeleit; Murmet Territory; Flalek Papan & Mount Kinabalu up to 1500 m), Brunei (Badas, swamp forest & Temburong District: Kuala Belalong 60 m), Sarawak (Niah National Park (= Batu Niah); Kuching [Bragg, 2001: 379]; 3km NE of Sungai Arip, 20 km SW of Tatau [Bragg, 2001: 379]; Baram River [Bragg, 2001: 379] & Mount Serapi 200 m), Kalimantan (Balekpapan; Mount Muret: Bukit Dadap; Long Bloe Oe; Bengen River: Tabang 125 m & Wattan). Number of specimens examined: 53



157

TABLE 17. Measurements [mm] of Phobaeticus kirbyi Brunner v. Wattenwyl, 1907.

&, HT

&&

%%

%, LT of pilicornis (NHMW)

(BMNH) Body

317.0

187.0–283.0

134.5–166.0

132.0

Pronotum

9.5

6.4–9.0

3.4–5.0

3.5

Mesonotum

68.0

41.0–60.5

30.5–37.5

30.0

Metanotum

23.0

14.7–20.0

7.0–8.3

-

Median segment

23.5

13.0–19.5

13.0–15.2

-

Tegmina

-

-

8.8–11.0

9.0

Alae

-

-

49.0–49.5

43.0

Profemora

71.0

50.0–74.0

54.0–59.5

52.5

Mesofemora

61.5

40.0–56.5

39.5–43.5

38.5

Metafemora

72.0

48.2–67.0

47.3–52.0

–

Protibiae

103.0

64.0–100.0

75.0–76.0

73.6

Mesotibiae

68.5

43.0–69.0

47.0–49.0

45.8

Metatibiae

94.0

55.0–90.0

61.0–65.0

-

Antennae

> 83.0

58.0–68.0

71.0–79.0

-

Phobaeticus lobulatus (Carl, 1913) comb. nov. (Figs. 125, 256, 274, 383) Eucarcharus lobulatus Carl, 1913: 12. HT, &: Eucarcharus lobulatus Carl, Ceylan (MHNG, No. 24). Zompro & Brock, 2003: 15. Otte & Brock, 2005: 136. Lobophasma lobulatus, Günther, 1935b: 138.

Diagnosis: Closely related and very similar to the second Sri Lankan species Ph. hypharpax (Westwood, 1859) but distinguished by: the smaller size; shape of the head; granules of the mesopleurae; more broadened and posteriorly rounded lateral lobes of tergite VII; tergite VIII being longer than the combined length of IX and X; much shorter and broader antennae and armature of the legs. Etymology: “Lobulatus” (lat. = lobed, with lobes) refers to the prominent rounded lateral lobes of abdominal tergite VII of &&. Description: && (Fig. 125): Medium-sized (body length 162.0 mm) and moderately slender for the genus (maximum body width 7.0 mm). General colouration of body more or less uniformly mid to dark brown with slightly lighter areas on the head, pronotum and near posterior margins of the meso- and metanotum. Great portions of sternum VI, almost entire sternum VII, the basal portion of the subgenital plate and bases of basitarsi whitish. Legs with irregular pale greyish transverse bands, all femora with a weakly defined black transverse band sub-basally. Leg armature black, the dorsal serrations of the profemora dull orange. Eyes dark reddish brown. Antennae dull reddish brown. Head: Suboval, 1.5x longer than wide, vertex very slightly convex, broadest at eyes and slightly narrowed towards the posterior. Between the bases of the antennae with a small transverse depression. Between the eyes with a very slightly impressed area and posterior portion of vertex very gently angled down-ward. Eyes rather small, circular and slightly convex; their length contained about 3x in that of cheeks. Antennae short, at least projecting over posterior margin of pronotum and consisting of 22 rather short and broad joints. Scapus dors-


HENNEMANN & CONLE

oventrally flattened, oval in dorsal aspect and 2.3x longer than wide. Pedicellus distinctly shorter than scapus, cylindrical. Thorax: Pronotum about 2/3 the length of head, about 1.5x longer than wide, almost parallel-sided. Median transverse depression indistinct. Anterior half of segment with an impressed median line. Mesothorax 2.2x longer than head and pronotum combined, slightly constricted at anterior margin and broadened posteriorly. Mesonotum with a very faint pale longitudinal median line. Mesopleurae with a longitudinal row of rounded granules. Metanotum less than half the length of mesonotum, parallel-sided, 2.5x longer than wide. Meso- and metasternum smooth. Abdomen: Median segment a little more than 2/3 the length of metanotum, parallel-sided, slightly more than 2x longer than wide. Segments II–VI parallel-sided and increasing in length, II 1.5x longer than wide and shorter than median segment, VI 2.5x longer than wide. Tergite VII slightly shorter than previous, lateral margins posteriorly dilated into an almost semi-circular lobe which projects by about half of body width (Fig. 274). Praeopercular organ formed by a pair of rounded, sub-cordiformous lobes at posterior margin of sternum VII. Tergite VIII slightly longer but distinctly narrower than VII, almost 4x longer than wide, constricted medially and strongly convex. IX half the length of VIII but of equal width, 1.5x longer than wide, rectangular. Anal segment shorter than IX and with a fine median carina, the posterior margin with a very broad but shallow triangular median excavation, outer angles narrowly rounded; posterior portion of anal segment strongly angled downward (Fig. 256). Supraanal plate very small, rounded and keeled. Cerci small, oval in cross-section, tapered towards apex and finely bristled. Gonapophyses elongate, filiform, slightly up-curving and ± reaching apex of anal segment. Subgenital plate slightly longitudinally keeled, boat-like and just reaching to posterior margin of anal segment, anterior margin down-curving at the apex. Legs: All long and slender with all carinae, except the protibiae, dentate. Profemora a little longer than pro- and mesonotum combined, mesofemora slightly shorter than mesothorax, metafemora reaching about half way along abdominal segment V and metatibiae roughly reaching apex of abdomen. Anterodorsal carina of profemora with 14–17 prominent, acutely triangular serrations; posteroventral carina with a similar number of small teeth. Five very faint, roundly triangular teeth in equal distance are present on the anterodorsal carina of the protibiae, this carina gently deflexed. Meso- and metafemora with the two outer ventral carinae gently widened sub-basally, teeth relatively larger in this section than on rest of femur. Posterodorsal carina of mesofemora with three indistinctly enlarged teeth near apex (the apical one largest) and a broad triangular tooth about ¼ away from the base. Dorsal carinae of metafemora sparsely dentate. Medioventral carina of meso- and metafemora set with 5–8 minute spines. Posterodorsal carina of meso- and metatibiae with a very broad but flat lobe about 1/3 off the base and forming a rounded, dentate lobe apically. Medioventral carina gently raised and rounded sub-basally and in this section set with 2–3 distinct, triangular teeth. Basitarsi shorter than combined length of remaining tarsomeres, all carinae gently elevated and the ventral carinae of the meso- and metabasitarsi with a few minute teeth. Comments: Carl (1913: 13) described E. lobulatus from a unique & in MHNG and placed his new species in the genus Eucarcharus Brunner v. Wattenwyl, 1907. The author stated it to be closely related to Eucarcharus inversus Brunner v. Wattenwyl, 1907 which however is generically distinct and belongs in Tirachoidea Brunner v. Wattenwyl, 1893. Eucarcharus Brunner v. Wattenwyl, 1907 (Type species: Lonchodes feruloides Westwood, 1859) is here shown to belong in the tribe Stephanacridini and is hence not closely related (→ 4.7.2). The not conically raised head, distinct praeopercular organ and genitalia clearly place E. lobulatus in Phobaeticus Brunner v. Wattenwyl, 1907. Due to the prominent lateral lobes of tergite VII Günther (1935b: 138) transferred E. lobulatus Carl to his Lobophasma Günther, 1935 (Type species: Eucarcharus rex Günther, 1928) but this genus is here shown to be a junior synonym of Phobaeticus (syn. nov.). Only known from the unique & HT. % and egg unknown. Distribution (Fig. 383): Sri Lanka. Number of specimens examined: 1



159

TABLE 18. Measurements [mm] of Phobaeticus lobulatus (Carl, 1913) comb. nov.

&, HT (MHNG) Body

162.0

Head

9.0

Pronotum

6.0

Mesonotum

34.0

Metanotum

12.5

Median segment

9.0

Profemora

41.3

Mesofemora

32.0

Metafemora

34.0

Protibiae

43.3

Mesotibiae

31.5

Metatibia

40.5

Antennae

18.5

Phobaeticus lumawigi Brock, 1997 (Figs. 120, 253–254, 330, 384) Phobaeticus lumawigi Brock, 1997: 43, figs. 1–2 (&). HT, &: Philippines, Mountain Province, North Luzon, VII.1996, leg. I. Lumawig; Holotype Phobaeticus lumawigi Brock (BMNH). Otte & Brock, 2005: 269.

Diagnosis: Related to the second Philippine species of the genus, Ph. philippinicus (Hennemann & Conle, 1997), and the Bornean Ph. redtenbacheri (Dohrn, 1910). Concerning to the length of the subgenital plate of && (30 mm) which is longer than in the first (23.2 mm) but only about half the length of that in the latter species (56.0–62.5 mm) it represents an intermediate (Fig. 254). It is easily distinguished from both species by: the smaller size; more slender and deeply notched anal segment (Fig. 253) and characteristic armature of the mid legs (Fig. 330). The shape of the head resembles Ph. kirbyi (Brunner v. Wattenwyl, 1907) from Borneo and Ph. palawanensis spec. nov. from Palawan, but the && of the latter species are not known. Etymology: Dedicated to Ismael O. Lumawig (Philippines) the collector of the HT of this distinctive species. Description: The unique HT was obviously discoloured by provisional conservation in ethanol and has the abdomen considerably shrunken. It is complete except for lacking the left fore leg and apices of the antennae. The left hind leg appears to have been regenerated. Taking typical discolouration of Phasmatodeans caused by ethanol into account, supposes the HT was either pale brown or dull green in live. && (Fig. 120). Long (body length 202.0 mm; including subgenital plate 211.0 mm) and slender species with a rather long and pointed subgenital plate (30.0 mm). General colouration of body and plain yellowish brown, head and pronotum slightly darker. Vertex with indistinct dark longitudinal lines on the vertex. Mesonotum with a fine dark brown longitudinal median line. Armature of the legs brown, the enlarged teeth of the mesofemora black towards the tip. Eyes dull reddish brown. Antennae pale greyish brown and becoming blackish basally; scapus and pedicellus pale yellowish brown. Head: Globose, slightly longer than wide and narrowed in posterior portion, vertex gently rounded. Between the bases of the antennae with a shallow transverse depression. Eyes circular, of moderate size and strongly convex; their length contained almost 3x in that of cheeks. Antennae slightly projecting over posterior margin of mesonotum (apical antennomeres broken


HENNEMANN & CONLE

in the HT). Scapus dorsoventrally flattened, almost 3x longer than wide, 2.5x longer than pedicellus and gently narrowed towards the base. Pedicellus round in cross-section and constricted towards apex. Thorax: Pronotum slightly shorter but decidedly narrower than head, 1.5x longer than wide and almost parallel-sided. Median transverse depression distinct, curved and almost reaching to lateral margin of segment. Mesothorax about 3x longer than head and pronotum combined, mesonotum parallel-sided. Metanotum less than 1/3 the length of mesonotum, parallel-sided and about 2x longer than wide. Meso and metasternum simple. Abdomen: Median segment slightly shorter than metanotum, anterior margin widely rounded. Abdominal segments II–VII roughly of equal length and almost 3x longer than wide. Praeopercular organ formed by two slender but pointed backward pointing spines at posterior margin of sternum VII. Tergite VIII 3/5 the length of VII and considerably narrower than II–VII. IX half the length of VIII, indistinctly longer than wide and rectangular. Anal segment slender, slightly longer than IX and the posterior margin with a broad and deep triangular excavation; posterolateral angles slender and acute (Fig. 254). Supraanal plate very small and triangular. Cerci slender, elongate and narrowing towards a pointed apex. Subgenital plate long, lanceolate, longitudinally keeled, projecting over abdomen by almost the combined length of tergites XI and X and tapered to a narrow apex (Fig. 253). TABLE 19. Measurements [mm] of Phobaeticus lumawigi Brock, 1997.

&, HT (BMNH) Body

202.0

Body (incl. subgenital plate)

211.0

Pronotum

7.5

Mesonotum

48.5

Metanotum

14.0

Median segment

11.0

Subgenital plate

30.0

Profemora

62.0

Mesofemora

46.0

Metafemora

55.0

Protibiae

83.0

Mesotibiae

50.0

Metatibiae

65.0

Antennae

62.0

Legs: All very long and slender, profemora a little longer than pro- and mesothorax combined, mesofemora slightly shorter than mesonotum, metafemora reaching about half way along abdominal segment V and metatibiae projecting over tip of subgenital plate. Anterodorsal carina of profemora with about 20 acute, moderate serrations which decrease in size towards apex of femur. Posteroventral carina with a similar number of small teeth and a considerably more prominent, triangular tooth about one third away from the base. Protibiae unarmed except for a very few rather small dorsal teeth. Posterodorsal carina of mesofemora with two prominent, elongate and pointed triangular sub-basal teeth followd by a slightly smaller tooth in a distance of about 6 mm. Two outer ventral carinae each with a prominent, triangular spine almost beneath the pair of teeth on posterodorsal carina (Fig. 330). All carinae otherwise sparsely and minutely dentate. Medioventral carina set with a few very minute spines. Posterodorsal carina of meso- and metatibiae dorsally set with 3 slightly



161

enlarged, triangular teeth (very faint on metatibiae) and apically protruded into a small, rounded and dentate lobe. Probasitarsus about as long as remaining tarsomeres combined, all carinae unarmed. Meso- and metabasitarsi as following three tarsomeres combined; the ventral carinae with a few minute teeth. Comments: Only known from the unique & HT. % and egg unknown. Distribution (Fig. 384): Philippines (Northern Luzon Island: Mountain Province). Number of specimens examined: 1

Phobaeticus magnus nom. nov. (Figs. 108–109, 184–185, 234–236, 310, 329, 385, 421–422) Phobaeticus magnus nom. nov. [Replacement name for Nearchus maximus Redtenbacher, 1908, a junior homonym of Phibalosoma maximum Bates, 1865: 341. HT = NT of N. maximus, see below]. HT, &: Thailand, Lomgao, Petchabun 21.X.1985, Phimpisarn; BMNH(E) 2005-98, Nearchus maximus Redt, 1908 &, det. P. Brock (BMNH). Nearchus maximus Redtenbacher, 1908: 448, pl. 21: 7a–b (apex of & abdomen). HT, &: Laos, Siam (MNHN – lost). NT [by present designation], &: Thailand, Lomgao, Petchabun 21.X.1985, Phimpisarn; BMNH(E) 2005-98, Nearchus maximus Redt, 1908 &, det. P. Brock (BMNH). [Secondary homonym of Phibalosoma maximum Bates, 1865: 341] Bragg, 1995b: 274. Nearchus maximus maximus, Bragg, 2001: 393. Otte & Brock, 2005: 209. Clitumnus operculatus Brunner v. Wattenwyl, 1907: 192 (in part ). PLT, & (nymph): Coll. Br. v. W., Assam, Staudinger; det. Br. v. W. Clitumnus operculatus (NHMW, No. 323). Baculum operculatus Brock, 1998a: 48. [Designation of LT] Ramulus operculatus, Otte & Brock, 2005: 305. [Not: Phobaeticus maximus (Bates, 1865), Otte & Brock, 2005: 269; = Phobaeticus serratipes Gray, 1853 syn. nov.] Further material: 6 &&, 2 %%, 1 & (nymph), eggs]: N-THAILAND: 1 &: N-Thailand, Lomgao, Petchabun, via Lehmann 1985 (coll. FH, No. 0130-1); Eggs: NE-Thailand, Nong Khai Province, Ampur Muang, Tumbol Pochai, Bang Rong mex village, laid by & collected I.2006, leg. L. Kwantale (coll. FH, No. 0130-E); 2 &&: N-Thailand, Lomgao, Petchabun, leg. Lehmann 21.10.1995 (coll. OC, No’s 00185 & 00186); 1 &, 1 %, 1 & (nymph): Thailand, Nakhon Ratchasima, S Khao Lak Chang, 400–475 m, 101°21É, 14°31´50 N, 23.IX.1998, leg. I. Fritzsche (coll. OZ, No’s 0448-1 to 3). 1 &: N-Thailand, via T. Thron (coll. FH, No. 0130-3). MYANMAR: 1 %: Myanmar, Tenasserim, leg. Lehmann 20.III.1994 (coll. FH, No. 0130-2). NO DATA: 1 & + 3 eggs on card: no data (MNCN).

Diagnosis: && are the only ones of the genus from continental Asia which have a long and lanceolate subgenital plate (Fig. 234) and are furthermore characteristic for the very prominent leg armature (Fig. 329), which includes strong spines on the medioventral carina of the meso- and metafemora. %% are characteristic for the distinctive and pretty colouration, very slender semi-tergites of the anal segment (Fig. 236) and comparatively long alae, which project well over abdominal segment IV. The eggs are characteristic for the comparatively elongate micropylar plate, which is almost 2/3 the length of the capsule and longer than wide (Figs. 184–185). Etymology: “Magnus” (lat. = large) refers to the very large size of this striking species and is meant to replace the original “maximus” (lat. = the largest) in a similar sense. For reasoning see comments below. Description: The description of the colouration is based on various colour photographs of live insects taken in the Nong Khai Province, N-Thailand and live captive reared specimens. && (Fig. 108, 421): Very long (body length 218.0–292.0 mm, including subgenital plate 234.0–315.0 mm) and slender species (maximum body width 6.0–7.0 mm) with a very long and lancet-like subgenital plate (39.0–56.0 mm); body surface slightly glabrous. General colouration of body and legs yellowish or greyish


HENNEMANN & CONLE

pale to mid brown with several slightly pale creamish to whitish areas. The HT has the entire terminal three abdominal segments creamish white. Complete body to a various degree furnished with paler and darker speckles and spots. Head usually with a large ± decided whitish or white marking on the vertex, the cheeks and posterior margin mid to dark brown. Mesonotum occasionally with a faint and washed pale green longitudinal median stripe. Meso- and metapleurae often whitish in the posterior portion. Meso- and metasternum greyish mid to dark brown and darker than rest of body, both irregularly set with elongate-oval pale yellow or cream spots. Abdominal tergite VII often with a white transverse marking posteriorly and tergites VIII and IX with a white patch posterolaterally. Bases of basitarsi and apices of femora white. Complete leg armature orange to dull red with black points. Eyes cream with sepia mottling (reddish brown in dried specimens). Antennae mid brown, scapus and pedicellus dark brown. Head: Suboval, 1.5x longer than wide, broadest at eyes and slightly tapering towards the posterior, vertex flat and angled posteriorly. Eyes prominent but moderately convex with the anterior margin gently angular; length contained almost 3x in that of cheeks. Between the bases of the antennae with a small but deep semicircular impression. Antennae at least reaching to posterior margin of mesonotum (broken in all examined specimens). Scapus almost 2.5x longer than wide, almost parallel-sided and dorsoventrally flattened. Pedicellus distinctly shorter than scapus, round in cross-section and slightly club-like. Following antennomeres first increasing, then decreasing in length towards apices of antennae. Thorax: Pronotum shorter and narrower than head, 1.3x longer than wide, the anterior half conspicuously constricted. Median transverse depression distinct, almost straight but short and not reaching lateral margins of segment. Mesothorax about 2.5x longer than head and pronotum combined, mesonotum parallel-sided except for being slightly widened at posterior margin and with a very indistinct median line. Tegmina represented by very small, kidney-shaped scales (c. 1.0 mm). Metanotum less than 1/3 the length of mesonotum, 2.5x longer than wide, parallel-sided and with a faint median line. Meso- and metasternum with very shallow longitudinal median keel. Abdomen: Median segment very slightly shorter than metanotum, about 2.5x longer than wide and gently constricted towards the anterior. Segments II–VI gradually increasing in length II 3x, V 3.5x and VI 4x longer than wide. VII about as long as III and 3.5x longer than wide. Praeopercular organ formed by two distinct, acutely triangular and carinate lobes on posterior margin of sternum VII. Tergite VIII narrower than previous, strongly convex, gently constricted medially and slightly more than half the length of VII. IX half the length of VIII, rectangular and indistinctly longer than wide. Anal segment 2/3 the length of VIII, parallel-sided with a distinct longitudinal median carina and the posterior margin broadly triangularly excavated, the outer angles short but acute. Supraanal plate rather prominent, triangular, strongly keeled and slightly projecting over posterior margin of anal segment (Fig. 235). Cerci small, oval in cross-section, tapered towards apex and slightly projecting over anal segment; finely sethose. Subgenital plate longitudinally keeled, very long, lancet-like and gradually tapering towards a pointed tip; projecting over apex of anal segment by ± the combined length of tergites VIII–X (Fig. 234). Legs: All long and slender with all carinae except protibiae more or less prominently spinose. Profemora a little longer than pro- and mesothorax combined, mesofemora slightly shorter than mesothorax, metafemora projecting over posterior margin of abdominal segment IV and metatarsi reaching to apex of anal segment. Anterodorsal carina of profemora with 16–20 moderately sized but very acute serrations, these decreasing in size towards apex of femur. Posteroventral carina armed with 10–14 prominent and long, pointed teeth; posterodorsal carina set with 6–10 distinct acute spines. All carinae of protibiae smooth except for a few very indistinct serrations on the posteroventral carina. Two outer ventral carinae of meso- and metafemora armed with several, irregularly set, prominent triangular spines; these straight on the posterior carina and slightly hooked on anterior carina. Medioventral carina armed with 8–10 strong and prominent, pointed spines which decrease in size towards apex of femur (Fig. 329). Dorsal carinae with a similar number but slightly smaller teeth than on ventral carinae. All ventral carinae of meso- and metatibiae densely set with rather long and



163

pointed spines, dorsal carinae sparsely dentate. Probasitarsus as long as, meso- and metabasitarsus slightly shorter than remaining tarsomeres combined, all carinae smooth, dorsal carina elevated and gradually raised towards the apex. %% (Fig. 109, 422): Large (body length 155.0–176.0 mm) and very slender for the genus (maximum width of abdomen 2.0–2.5 mm) with long alae (57.5–71.0 mm); body surface slightly glabrous. General colouration of body yellowish or greenish mid brown. Head with a large white marking on the vertex, which covers almost the complete dorsal head surface and narrows towards the posterior. Cheeks with a faint, washed blackish postocular stripe. Mesonotum mid to dull green. Great portions of mesopleurae whitish, metapleurae grey with the lower margin white and the dorsal portion black. Mesosternum dark brown with distinct straw elongate-oval patches and the lateral margins cream. Metasternum with a white patch posteromedially, otherwise mid brown and with two parallel, dark brown to black longitudinal median lines. Tegmina and costal region of alae greyish or yellowish brown. Anterior margin and posterior half of posterior margin of tegmina broadly white and interiorly bordered by a fine black line; central portion with a bold oval white patch and an elongate black marking on the central hump which terminates towards the apex of the tegmen. Costal region of alae with a bold white band along anterior margin. Anal region transparent grey with brown veins. Abdominal tergites and sternites VIII and IX as well as the poculum with conspicuous white markings. Lateral margins of abdominal tergites with a very fine white longitudinal line. Abdominal sternites II–VII dark brown with a white line along lateral margins. Legs yellowish or reddish mid brown, the complete armature (except dorsal serrations of profemora) dull red with black points. Each coxa with a white lateral patch and all femora with a small, white lateral spot near the apex. Basal portion of profemora straw or pale grey. Bases of basitarsi white. Eyes cream with brown mottling. Antennae very dark reddish brown. Head (Fig. 310): Oval, 2x longer than wide and distinctly narrowed towards the posterior, vertex flat, otherwise as in &&. Between bases of antennae with a small but deep impression. Eyes large and projecting hemispherically from head capsule; their length contained about 2x in that of cheek. Antennae projecting over posterior margin of metanotum, otherwise as in &&. Thorax: Pronotum 2x longer than wide, shorter and narrower than head and generally as in &&. Mesothorax about 2.6x longer than head and pronotum combined, mesonotum parallel-sided. Meso- and metasternum with very fine longitudinal median carina. Tegmina oval, constricted basally and with a distinct, roundly conical central hump. Alae projecting over posterior margin of abdominal tergite IV. Abdomen: Median segment considerably longer than metanotum. Segments III–VI 6.5x, II and VII shorter, only 5.5x longer than wide. Tergite V with small scale-like posteromedian tubercle. VIII slightly more than 2/3 the length of VII, 3x longer than wide, strongly convex and constricted medially. IX slightly shorter than previous, anterior half slightly swollen. Anal segment longer than VIII laterally compressed, distinctly keeled and tectiform. Semi-tergites very elongate, slender and tapered towards a narrow apex; gently downcurving (Fig. 236). Interior surface apically armed with several minute black teeth. Cerci elongate, oval in cross-section, gradually tapered towards a pointed apex, gently in-curving and almost reaching tip of anal segment. Poculum convex, cup-like, longitudinally keeled, reaching about half way along tergite IX and with a very prominent and acute, backward pointing central spine. Legs: All very long and slender, profemora longer than head, pro- and mesothorax combined, about as long as pro- and mesonotum, metafemora slightly projecting over posterior margin of abdominal segment IV and tarsi very slightly extending over apex of abdomen. Anterodorsal carina of profemora with 20–24 acutely triangular serrations, posteroventral carina with a similar number of medium sized, pointed spines. Medioventral carina of meso- and metafemora distinct and armed with 6–10 distinct, needle-like spines. Spination otherwise as in && but much less distinct. Probasitarsus very elongate, 1.5x longer, meso- and metabasitarsus as long as remaining tarsomeres combined, all carinae smooth, the dorsal carina slightly gradually raised towards the apex. Eggs (Figs. 184–185): Capsule basically lens-shaped, laterally compressed, longer than high. General


HENNEMANN & CONLE

colouration of capsule and micropylar plate greyish mid brown, operculum slightly darker brown. Capitulum black with a dark brown stalk. Whole capsule surrounded by dorsoventral keel, beginning and ending at the operculum and only interrupted near the polar-area. Keel very faint on micropylar plate. Capsule surface slightly rugulose. Micropylar plate elongate, tapered towards the anterior and broadest 1/3 of posterior end; almost 2/3 the length of capsule. Outer margin marked by a dark sepia brown line. Micropylar cup small, positioned at polar end of plate; black. Operculum flat, oval and in its centre with a prominent, irregularly shaped, roughly cone-shaped capitulum on a short stalk. Measurements [mm]: length (including capitulum) 5.1–5.6, length 4.3–4.8, height 3.4–4.1, width 2.7–2.9, length of micropylar plate 2.7–3.1. Comments: Redtenbacher (1908: 448) originally described Nearchus maximus from a single & from “Laos, Siam”, showing that Redtenbacher appears to have been in doubt about the exact collecting locality. Extensive search in MNHN has not traced Redtenbacher’s HT why it must be presumed lost. The original description, measurements and illustration of the apical abdominal segments clearly identify Redtenbacher’s species and leave no doubt about the conspecifity of the material at hand from N-Thailand and Myanmar. In order to fix Redtenbacher’s species, the & from Lomgao Petchabun, N-Thailand in BMNH is here selected as the Neotype of Redtenbacher’s Nearchus maximus. This designation comfors the Article 75.3. of the Code (ICZN, 1999) on the conditions of neotype designations. Phibalosoma maximum Bates, 1865 was transferred to Phobaeticus by Otte & Brock (2005: 269) ans is here shown to be a junior synonym of Phobaeticus serratipes (Gray, 1835). As the genus Nearchus Redtenbacher, 1908 is a junior synonym of Phobaeticus Brunner v. Wattenwyl, 1907 (syn. nov.) its type species N. maximus Redtenbacher, 1908 automatically becomes a member of Phobaeticus. Followingly, and in accordance to the Article 53.3 of the Code (ICZN, 1999 ) this act makes Nearchus maximus Redtenbacher a secondary homonym of Phibalosoma maximum Bates with the latter still being an available synonym. Therefore, Phobaeticus magnus nom. nov. is here established as a replacement name for Nearchus maximus Redtenbacher, which is in accordance to Article 23.3.6. of the Code (ICZN, 1999) on the principles of priority. The here selected NT of N. maximus Redtenbacher automatically becomes the HT of Ph. magnus nom. nov. Examination of the & PLT of Clitumnus operculatus Brunner v. Wattenwyl , 1907 in NHMW has shown this to represent a penultimate instar nymph of Ph. magnus. The specimens from Nakhon Ratchasima (Thailand) in coll. OZ are considerably shorter than specimens from other localities but do not show any other significant differences. An adult & and several nymphs were collected by L. Kwantalae (Nong Khai) at Bang Rong mex village, Ampur Muang in the Nong Khai Province, NE-Thailand. These were found during the day-time in dry secondary growth and kept alive in large cages on guava (Psidium guajava, Myrtaceae), mango (Mangifera indica, Anacardiaceaee) and Tamarindus indica (Fagaceae). Numerous eggs were obtained and distributed to European breeders, some were kindly given to the authors for examination. Breeding Ph. magnus in captivity in Europe is now being attempted by several enthusiasts but has as yet proven rather difficult. In captivity it accepts oak (Quercus robur & Q. ilex, Fagaceae) and bramble (Rubus fruticosus, Rosaceae) as alternative foodplants. Distribution (Fig. 385): Northern Thailand (Lomgao Petchabun; Nong Khai Province: Ampur Muang, Tumbol Pochai “Bang Rong mex village” & Nakhon Ratchasima: S Khao Lak Chang 400–475 m), N-Myanmar (Highland of Tenasserim), NE-India (Assam) and Laos [Redtenbacher, 1908: 448]. Number of specimens examined: 10



165

TABLE 20. Measurements [mm] of Phobaeticus magnus nom. nov.

&, HT (MNHN)*

&, NT (BMNH)

&&

%%

Body (incl. subgen. plate)

276.0

297.0

234.0–315.0

-

Body

-

263.0

218.0–292.0

155.0–176.0

Head

-

12.2

9.3–12.8

6.3–7.2

Pronotum

-

8.7

6.7–9.6

4.0–5.0

Mesonotum

56.5

53.0

45.0–62.6

30.5–34.5

Metanotum

34.5**

18.2

13.8–19.0

8.9–10.0

Median segment

-

14.0

12.6–17.2

13.1–15.0

Subgenital plate

43.0

55.0

39.0–56.0

-

Tegmina

-

-

-

10.1–12.0

Alae

-

-

-

57.5–71.0

Profemora

65.0

68.0

54.5–70.0

43.7–51.0

Mesofemora

-

55.0

44.0–56.0

34.5–41.5

Metafemora

59.0

63.0

50.0–65.8

43.0–47.8

Antennae

-

> 50.0

> 40.0–72.0

68.0–76.5

* according to Redtenbacher (1908: 448) ** including median segment

Phobaeticus mjoebergi (Günther, 1935) comb. nov., stat. nov. (Fig. 386) Nearchus maximus mjöbergi Günther, 1935a: 10. HT, &: Zentral Borneo, Mt. Tibang, 1700m (NHRS). Nearchus maximus mjobergi, Bragg, 1995b: 274. Bragg, 2001: 394. Otte & Brock, 2005: 209.

Diagnosis: Closely related to the Bornean Ph. redtenbacheri (Dohrn, 1910) but distinguished by: the smaller size; relatively longer mesonotum; lacking any larger teeth or lobes on the legs and the smooth medioventral carina of the meso and metafemora. Etymology: Dedicated to Dr. E. Mjöberg a Swedish naturalist who predominantly worked in Indonesia and collected the HT of this species on an expedition to Central Borneo during 1925–1926. Description: The following characterization is based on Günther’s original description of the HT in NHRS (1935a: 10). &&: Very long (body length 206.0 mm, including subgenital plate 249.0 mm) and slender species (width of metanotum 5.5 mm) with a very long lancet-like subgenital plate, which prominently projects over the posterior margin of the anal segment (by 43.0 mm). Anal segment keeled, posterior margin truncate and straight. Supraanal plate keeled and rounded apically. Cerci short, small and tapering towards the apex. Praeopercular organ formed by two blunt spines at posterior margin of abdominal sternum VII. Anterodorsal carina of profemora strongly serrate, posteroventral carina with numerous long and slender teeth. Protibiae with minute teeth at wide intervals on all carinae. Meso and metafemora with all carinae regularly dentate, the dorsal teeth are small and those on the ventral carinae long and slender. Medioventral carina of meso and metafemora unarmed. Meso- and metatibiae densely but rather minutely serrate, medioventral carina with a few small teeth at wide intervals. Any larger teeth or lobes are lacking.


HENNEMANN & CONLE

Comments: Günther (1935a: 10) originally described P. mjoebergi as a subspecies of Nearchus maximus Redtenbacher, 1908 (= Phobaeticus magnus nom. nov.) which however is restricted to continental Asia. Although the HT in NHRS could not be examined its distribution in Borneo and some of the features mentioned by Günther clearly prove it to be a valid species. It is undoubtedly more closely related to Ph. redtenbacheri (Dohrn) from Borneo rather than to Ph. magnus nom. nov. Bragg (2001: 394) provided an English translation of Günther’s original description. Only known from the unique & HT. % and egg unknown. Distribution (Fig. 386): Central Borneo (Kalimantan: Mount Tibang 1700 m). Number of specimens examined: None TABLE 21. Measurements [mm] of Phobaeticus mjoebergi (Günther, 1935) comb. nov., stat. nov.

&, HT (NHRS)* Body (incl. subgenital plate)

249.0

Body

206.0

Mesonotum

51.0

Metanotum

18.0

Median segment

13.0

Subgenital plate (projecting part)

43.0

Profemora

75.0

Mesofemora

56.0

Metafemora

79.0

Protibiae

117.0

Metatibiae

85.0

* according to Günther (1935a: 10)

Phobaeticus mucrospinosus spec. nov. (Figs. 105, 255, 328, 387) HT, &: Mus. Leiden, P.t. Stolz, Solok Borenl., Febr. 1914; Pharnacia spec. ignota K. Günther det. 1956 (RMNH). PT, 3 eggs (removed from abdomen of HT): same data as HT (RMNH).

Diagnosis: This new species is related to Ph. serratipes (Gray, 1835) from Peninsular Malaysia and Sumatra, the Sumatran Ph. sobrinus Brunner v. Wattenwyl, 1907 and perhaps Ph. chani Bragg spec. nov. from Borneo. && (the only sex known) however easily distinguished from these and all other members of the genus by the three distinct and conspicuously dagger-like spines in the basal quarter of the posteroventral carina of the mesofemora. Etymology: The name mucrospinosus refers to the characteristic ventral armature of the mesofemora and is a combination the two Latin words “mucro” (= dagger) and “spinus” (= spine). Description: && (Fig. 105): The unique & in RMNH is in poor condition and incomplete. Apart from showing damage caused by parasites, the complete antennae, both fore legs, and last four tarsomeres of the meso- and metatarsi are broken off and missing. Very long (body length 233.0 mm) and slender species (maximum body width 5.0 mm) with a short subgenital plate. General colour of the body creamish mid brown with a few white spots on metanotum, median segment and abdominal tergites II–III. Head more yellowish brown than rest of body and with several faint REVISION OF PHARNACIINI


167

dark longitudinal lines. Legs pale brown to straw and becoming lighter towards the bases of the femora. Dagger-like ventral spines of the mesofemora and enlarged teeth of the tibiae and metafemora dark brown. Eyes reddish mid brown. Head: Oval, 1.3x longer than wide, broadest at the eyes and distinctly narrowed towards the posterior; vertex almost flat. Posterior margin slightly angulate and convex medially. Vertex with a dark faint coronal line in front which terminates near the centre of head capsule and two dark longitudinal slightly converging lateral lines; the outer ones shorter. Between the eyes with a large transversely triangular, gently raised area. A transverse impression between the bases of the antennae. Eyes small and projecting hemispherically from head capsule; their length contained 2.4 in that of cheeks. Thorax: Pronotum narrower but almost as long as head, slightly widening towards the posterior; 1.6x longer than wide. Anterior margin raised and follow by a distinct transverse furrow. Median transverse depression deep, gently curved but just not reaching lateral margins of segment. Mesothorax more than 3x longer than head and pronotum combined, mesonotum parallel-sided except for being very slightly broadened at the posterior. Median line dark brown, very faint. Metanotum about half the length of mesonotum, parallelsided, 4.4x longer than wide. Meso- and metasternum with a very fine and slightly raised longitudinal median line. Abdomen: Median segment slightly shorter than metanotum, posterior margin broader than anterior margin, slightly constricted medially. Segments II–V slightly increasing in length, VI about as long as V, II less than 2.5x, VI almost 4x longer than wide. VII slightly shorter and narrower than previous, about 3x longer than wide. Tergites II–IX with a blunt posteromedian tubercle; this most prominent on V and VI. Praeopercular organ moderately developed and formed by two small, blunt spiniform tubercles at posterior margin of sternum VII. Tergite VIII considerably narrower but almost as long as VII, 5x longer than wide, strongly convex and parallel-sided. IX a little more than 1/3 the length of VIII, strongly convex and broadened in posterior half. Anal segment slightly shorter than IX, parallel-sided, with a fine longitudinal median carina and a small roughly semi-circular posteromedian notch; the posterolateral angles truncate. Supraanal plate very small, tapered towards apex and with a fine median keel. Cerci very small, slightly laterally compressed and gently up-curving. Subgenital plate longitudinally keeled and roughly reaching to posterior margin of anal segment (apex broken in the HT, Fig. 255). Legs: Mid and hind legs long and slender with all carinae minutely spinose. Mesofemora about as long as mesothorax, metafemora reaching to posterior margin of abdominal segment V and metatibiae slightly projecting over apex of abdomen. Medioventral carina of meso- and metafemora indistinct and set with 10–15 minute spines. Posteroventral carina of mesofemora with three prominent, long (3.3 mm, 4.0 mm and 2.6 mm), basally constricted and conspicuously dagger-like spines in basal quarter of femur (Fig. 328). Metafemora with two needle-like sub-basal spines on same carina. Posterodorsal carina of meso- and metatibiae with a broad, triangular tooth near the middle (very indistinct on metatibiae) and raised into a rounded, dentate lobe at the apex. Medioventral carina of the meso and metatibiae with a flat, triangular spine about one third of the base of tibia (very indistinct on metatibiae). Metabasitarsus elongate, all carinae minutely dentate. Eggs: Three eggs (PT) could be extracted from the abdomen of the HT. They are however in a rather early stage of development and do not show any specific structures which would allow a confirmed differentiation from the eggs of other species in the genus. Hence only a very brief characterization and measurements are provided. Of moderate size, globose, almost as wide as long and oval in cross-section. Micropylar plate oval, longer than wide, about half the length of capsule and with a median notch posteriorly. Operculum oval, slightly convex and with an irregularly cone-shaped capitulum. Greyish mid brown. Measurements [mm]: length (including capitulum) 4.7, length 3.5, width 3.0, height 3.4, length of micropylar plate 2.2. Comments: % unknown.


HENNEMANN & CONLE

Distribution (Fig. 387): Eastern Sumatra (Solok). Number of specimens examined: 1 TABLE 22. Measurements [mm] of Phobaeticus mucrospinosus spec. nov.

&, HT (RMNH) Body

233.0

Head

9.6

Pronotum

8.0

Mesonotum

54.0

Metanotum

22.5

Median segment

17.3

Mesofemora

53.5

Metafemora

63.8

Mesotibiae

61.2

Metatibiae

79.0

Phobaeticus palawanensis spec. nov. (Figs. 122, 263, 311, 347, 388) HT, %: Philippinen, Palawan, Napsan, Salakot Pass, 350 m, 7. IX. 1997, ex coll. FH, No. 0410-1 (ZSMC). PT, 2 %%: Philippinen, Palawan, Napsan, Salakot Pass, 350 m, 7.IX.1997 (coll. OC).

Diagnosis: This new species is related to Ph. philippinicus (Hennemann & Conle, 1997) from the Philippines and the Bornean Ph. redtenbacheri (Dohrn, 1910), but %% (the only sex known) differ from those of both species by the strongly globose, almost spherical head (Fig. 311). From the first it additionally differs by: the long median segment, which is longer than the metanotum; presence of small, vestigial tegmina and alae; shape of the semi-tergites of the anal segment (Fig. 263) and the more spine-like dentations of mid and hind legs. Ph. redtenbacheri is easily distinguished from Ph. palawanensis spec. nov. by the well developed tegmina and alae. The globose head also resembles the Bornean Ph. kirbyi Brunner v. Wattenwyl , 1907 but this species differs in a number of features such as the presence of well developed tegmina and alae and a conspicuous black ventrolateral marking on the cheeks. Etymology: The specific name refers to the geographic distribution of this new species, which is the so far only known representative of Phobaeticus Brunner v. Wattenwyl, 1907 in Palawan. Description: %% (Fig. 122): Medium sized (body length 140.0–162.0 mm) and slender species (maximum body width 2.5 mm), with a long median segment and small, vestigial tegmina (2.5–3.0 mm) and alae (1.5–2.1 mm). General colouration greenish to yellowish brown (possibly dull green when alive), legs pale greenish. Armature of the legs dull orange, the dorsal serrations of the profemora dark brown. Eyes creamish orange brown with irregular sepia brown mottling. Scapus and pedicellus pale brown, remaining parts of antennae black. Head (Fig. 311): Strongly globose and almost spherical, indistinctly longer than wide; vertex with two impressed longitudinal lines. Eyes large, circular and projecting hemispherically from head capsule; their length contained about 2.3x in that of cheeks. Antennae consisting of about 28 segments, projecting over posterior margin of median segment; all antennomeres densely covered with long curly hairs. Scapus dorsoventrally flattened and almost 3x longer than wide, parallel-sided. Pedicellus slightly longer than wide, 1/3 the length of scapus, round in cross-section. REVISION OF PHARNACIINI


169

Thorax: Pronotum distinctly narrower than head, slightly less than 2x longer than wide, anterior half narrowed. Lateral margins slightly raised, median transverse depression faint, straight and short, not reaching lateral margins of segment. Mesothorax almost 4x longer than head and pronotum combined, mesonotum parallel-sided. Metanotum distinctly shorter than mesonotum and slightly shorter than median segment, parallel-sided and almost 4x longer than wide. Meso- and metasternum with a fine longitudinal median carina. Tegmina very small and vestigial, forming a small (3.0 mm) elongate lobe which is strongly constricted towards the base and rounded at the apex. Alae reduced to a pair of very minute elongate lobes (2.1 mm). Abdomen: Median segment slightly longer than metanotum, parallel-sided, 5x longer than wide. Segment II a little longer than median segment and about 5x longer than wide. III–VI about 6x longer than wide, indistinctly constricted medially. All tergites with a very faint longitudinal median line, IV–VI with a minute scalelike posteromedian tubercle. Segment VII shorter than previous, about 4.5x longer than wide. Tergite VIII 2/3 the length of VII and slightly widening towards the posterior. IX about 2/3 the length of VIII, strongly convex, constricted medially and with a blunt median keel; anterior portion widened. Anal segment 1/3 longer than VIII, laterally compressed, strongly keeled and tectiform. Semi-tergites moderately long, straight and tapered towards a narrow apex (Fig. 263). The interior surface apically set with small, hooked spines. Cerci small, round in cross-section, tapering towards a pointed apex and slightly in-curving. Poculum strongly convex, cup-like, with a blunt, conical hump at the angle and reaching about half way along tergite IX. Legs: All very long and slender, profemora longer than head, pro- and mesothorax combined, mesofemora a little longer than mesothorax, metafemora slightly projecting over posterior margin of abdominal segment V and metatibiae extending considerably over apex of abdomen. Anterodorsal carina of profemora with 17–20 prominent, acutely triangular serrations, posteroventral carina with a few minute teeth (Fig. 347). Protibiae unarmed, but finely sethose. Two outer ventral carinae of meso- and metafemora densely set with pointed spines of moderate size. Dorsal carinae very minutely and sparsely dentate. Medioventral carina indistinct and unarmed. Meso and metatibiae with very minute teeth on all carinae (more prominent ventrally). Probasitarsus very elongate, 1.5x longer than remaining segments combined, all carinae smooth. Meso- and metabasitarsi indistinctly longer than remaining tarsomeres combined, all carinae minutely dentate. Comments: & and egg unknown. Distribution (Fig. 388): Palawan (Salakot Pass: Napsan 350 m). Number of specimens examined: 3 TABLE 23. Measurements [mm] of Phobaeticus palawanensis spec. nov.

%, HT (ZSMC) Body Head Pronotum Mesonotum Metanotum Median segment Tegmina Alae Profemora Mesofemora Metafemora Protibiae Mesotibiae Metatibiae Antennae


148.5 4.9 4.5 39.3 8.2 9.7 3.0 2.1 53.5 40.5 48.0 72.4 45.0 61.1 66.0

%%, PT

(coll. OC)

140.0–162.0 4.6–5.1 4.0–4.1 32.2–38.2 8.1–8.9 12.0–14.9 2.5–2.7 1.5–1.7 51.0–61.0 38.3–43.0 42.9–53.9 74.5–87.0 44.0–49.1 60.1–66.0 > 50.0-82.0

HENNEMANN & CONLE

Phobaeticus philippinicus (Hennemann & Conle, 1997) (Figs. 119, 123, 175–176, 240–242, 348–350, 389) Baculolonga philippinica Hennemann & Conle, 1997a: 349, figs. 10–16. HT, %: Philippinen, Mindoro Id., Mt. Halcon 2000 m, leg. Mohagan 15.–18.V.1996 (ZSMC, ex coll. FH 0284-1); AT, &: Philippinen, Mindoro Id., Mt. Halcon 2000 m, leg. Mohagan 2.VI.1996 (ZSMC, coll. FH 0284-3); PT, %: same data as AT (coll. FH 0284-4); PT: 1 %, 1 & (nymph): same data as HT (coll. FH 0284-2 & 5); PT, %: Philippines, Mindoro Id., Barrio Luyang, Mt. Halcon 360m, 500 m, 14.–17.V.1981, leg. R.A. Müller (coll. OZ). Phobaeticus philippinicus, Otte & Brock, 2005: 269. Further material: 2 &&, 2 %%, 1 egg]: PHILIPPINES: 2 %%: Philippinen, Mindoro Isl., Mt. Halcon, leg. Mohagan 15.–26.V.1996 (coll. OC); 2 &&, 1 egg: Philippinen, Mindoro Isl., 08.2006 (coll. OC).

Diagnosis: && are very similar to Ph. serratipes (Gray, 1835) from Peninsular Malaysia and Sumatra and the Philippine Ph. lumawigi Brock, 1997, and %% in particular resemble those of Ph. redtenbacheri (Dohrn, 1910) from Borneo and Ph. palawanensis spec. nov. from Palawan. && are characteristic for the apical lobe of the protibiae which otherwise only found in the Bornean Ph. redtenbacheri (Dohrn). The eggs resemble those of Ph. redtenbacheri (Dohrn) but differ by the relatively larger micropylar plate and lateral holes instead of swellings of the dorsoventral keel of the capsule (Figs. 175–176). From Ph. serratipes it is distinguished by: the longer subgenital plate of &&, which slightly extends over the posterior margin of the anal segment (Fig. 240); the shorter, more globose and posteriorly narrowed head and shorter median segment of both sexes as well as the lack of tegmina and alae and more slender, apically tapered semi-tergites of the anal segment of %% (Fig. 242). The occasional presence of prominent sub-basal spine on the posteroventral carina of the profemora resembles Ph. lumawigi from which && however clearly differ by: the much longer and more slender body; shorter apically rounded subgenital plate; flat and more rounded posteromedian incision of the anal segment (Fig. 241) and different armature of legs. %% differ from Ph. palawanensis spec. nov. by: the larger size and more slender body; posteriorly narrowed and less globose head; considerably shorter median segment; lack of vestigial tegmina and alae; less tooth-like dentations of the meso- and metatibiae and more slender semi-tergites of the anal segment (Fig. 242). From Ph. redtenbacheri it at once differs by: the much shorter, not lanceolate and apically rounded subgenital plate of && (Fig. 240); shorter median segment and less distinct leg armature of both sexes, as well as the lack of tegmina and alae; apically tapered and acute semi-tergites of the anal segment and longer cerci of %%. Concerning to the length of the subgenital plate, && represents a transitory from Ph. serratipes which has the subgenital plate just reaching the posterior margin of the anal segment to Ph. lumawigi and Ph. redtenbacheri in which it is increasingly elongated, and ± lanceolate. Etymology: The name refers to the distribution of this very long species, the Philippine Islands. Description: && (Fig. 119): Very long (body length 236.0–247.5 mm) and slender species (maximum body width 5.0–6.0 mm), subgenital plate slightly projecting over posterior margin of anal segment. General colouration of body and legs pale ochracheous to dark reddish brown, to a various degree all over furnished with numerous irregular lighter brown and blackish speckles. Head in the AT with distinct straw and dark brown patches especially along the coronal line. Eyes dark reddish brown. Legs with very faint pale transverse bands, armature very dark orange. Antennae mid brown and becoming darker towards the apex. Head: Oval slightly globose, 1.4x longer than wide, broadest at eyes and narrowed towards the posterior. Vertex gently rounded and with a fine impressed coronal line. Eyes rather small, circular and projecting hemispherically; their length contained more than 2.5x in that of cheeks. Antennae consisting of about 28 segments and reaching almost 2/3 the way along mesonotum, all antennomeres finely sethose. Scapus about 2.5x longer than wide, dorsoventrally compressed and slightly narrowed towards the base. Pedicellus slightly longer than REVISION OF PHARNACIINI


171

wide, cylindrical and about 1/3 the length of scapus. 3rd Segment indistinctly longer than pedicellus and tapered towards apex. Following segments first increasing then decreasing in length towards apex of antennae. Thorax: Pronotum 2x longer than wide, slightly shorter and narrower than head, the anterior half strongly constricted. Just behind anterior margin with a transverse furrow, which is bended backwards at the outer angles of the segment. Median transverse depression a short but prominent semicircle, opened anteriorly. Mesothorax about 3.6x longer than head and pronotum combined, mesonotum parallel-sided and with a faint, slightly impressed longitudinal median line. Metanotum almost 4x longer than wide, parallel-sided. Mesoand metasternum smooth. Abdomen: Median segment about 3/5 the length of the metanotum, almost 3x longer than wide and gradually widened towards the posterior. Segments II–V increasing in length, II a little longer than median segment and about 3.5x longer than wide, V almost 5x longer than wide. VI a little shorter than V, VII considerably shorter than VI. Tergite VII shorter than previous, about 2/3 the length of VI and 2.5x longer than wide, lateral margins very gently deflexed posteriorly. Tergites II–VI with a blunt posteromedian tubercle, this most distinct on III–V. Praeopercular organ formed by two distinct and slender, pointed spines at posterior margin of sternum VII. Tergite VIII distinctly narrower than previous, constricted medially, strongly convex and about 3.5x longer than wide. IX indistinctly longer than wide, strongly convex. Anal segment with a fine longitudinal median carina, posterior margin with wide and rounded median excavation, the posterolateral angles roundly triangular (Fig. 241). Supraanal plate short, rounded posteriorly, keeled and with a minute posteromedian notch. Cerci long, slender, round in cross-section, gently in-curving and tapering towards a narrow apex. Subgenital plate longitudinally keeled and slightly (2.0–4.0 mm) projecting over posterior margin of anal segment, apex rounded (Fig. 240). Legs: All very long and slender, profemora about as long as head, pro- and mesothorax combined, mesofemora roughly equal in length to mesothorax, metafemora projecting over posterior margin of abdominal segment IV and metatibiae exceeding apex of abdomen. Anterodorsal carina of profemora with 18–25 prominent, sharp serrations which decrease in size towards apex of femur. Posteroventral carina very sparsely set with minute teeth and occasionally with a single ± enlarged tooth sub-basally (& nymph PT, Fig. 350). Posterodorsal carina of protibiae set with one or two triangular teeth of moderate size and forming a conspicuous, rounded and dentate lobe at the apex of tibia. Meso- and metafemora with all carinae dentate, more sparsely although on dorsal carinae. Posterodorsal carina occasionally with two slightly enlarged teeth in basal quarter, the posteroventral carina often with a conspicuous needle-like spine sub-basally. Medioventral carina very fine and with a few minute spines. Posterodorsal carina of mesotibiae (indistinct on metatibiae) with three ± enlarged, broadly triangular teeth in great distance to another (Fig. 348) and with the apex forming a rounded, dentate lobe. Otherwise all carinae of meso and metafemora minutely dentate, although more sparsely on dorsal carinae. Probasitarsus very elongate and about as long as remaining tarsomeres combined, dorsal carina gently deflexed towards the apex, all carinae unarmed. Meso- and metabasitarsi a little shorter than combined length of remaining tarsomeres, ventral carinae minutely dentate. & (nymph): The PT nymph in coll. FH (No. 0284-5) has the leg armature decidedly more developed than the adult && at hand. The posteroventral carina of the profemora bears a very prominent hook-like tooth subbasally (Fig. 350). Both ventral carinae and the posterodorsal carina of meso- and metafemora are armed with a prominent triangular sub-basal tooth (Fig. 349). The three dorsal lobes of the meso- and metatibiae are considerably larger and lobe-like (Fig. 348). %% (Fig. 123): Medium sized to large (body length 143.0–165.5) and very slender (maximum body width 2.0–3.0 mm) for the genus, apterous. General colouration of body and legs plain green or mid to dark brown, occasionally with slightly lighter areas on head and thorax. Green specimens to a variable degree furnished with brownish speckles. Leg armature dull yellow with black points. Eyes creamish brown with darker mottling. Antennae pale greenish brown in green specimens, mid brown in brown specimens.


HENNEMANN & CONLE

Head: Indistinctly longer than wide and almost trapezoidal in dorsal aspect, being distinctly narrowed from the eyes towards the posterior; vertex flat. Posterior margin with three slightly impressed, short longitudinal lines. Eyes large, very prominent and projecting hemispherically; their length contained a little more than 2x in that of cheeks. Antennae projecting over posterior margin of median segment, all antennomeres densely sethose. Otherwise generally as in &&. Thorax: Pronotum narrower but slightly longer than head, almost 2x longer than wide and slightly constricted in anterior half; otherwise as in &&. Mesothorax about 3.7x longer than head and pronotum combined, mesonotum parallel-sided. Metanotum parallel sided, about 2/5 the length of mesonotum and roughly 7x longer than wide. Meso- and metasternum with a fine longitudinal median carina (less decided on metasternum). Abdomen: Median segment slightly shorter and narrower than metanotum, about 5x longer than wide and slightly constricted medially. Segment II 6x, III–VI 7x longer than wide, all gently constricted medially. Tergites IV–VI with a minute posteromedian tubercle, which is most distinct on V. VII shorter than VI, 4.5x longer than wide and gently widened at the posterior. Tergite VIII ¾ the length of VII, strongly widening towards the posterior. IX half the length of VIII, convex and strongly constricted in posterior portion. Anal segment as long as VII, strongly keeled, tectiform and laterally compressed. Semi-tergites long, straight and with the apex narrow (Fig. 242). Interior surface apically with a prominent longitudinal median bulge which is covered with minute teeth. Cerci elongate, tapering towards a pointed apex and distinctly in-curving. Poculum strongly convex, cup-like, angular and with a blunt central spine; reaching about half way along tergite IX. Legs: All very long and slender, profemora longer than head, pro- and mesothorax combined, mesofemora about as long as combined length of pro- and mesothorax, metafemora slightly projecting over posterior margin of abdominal segment V and metatibiae considerably exceeding apex of abdomen. Anterodorsal carina of profemora with ± 20 moderately prominent and sharply triangular serrations. Posteroventral carina set with only 6–12 small teeth. Protibiae unarmed, except for a few minute teeth on posteroventral carina. All carinae of mid and hind legs regularly set with small but pointed spines, more prominent on ventral carinae. Medioventral carina faint and unarmed. Probasitarsus 1.5x longer than remaining tarsomeres combined. Meso- and metabasitarsi slightly longer than combined length of remaining tarsomeres, all carinae with a few minute teeth. Eggs (Figs. 175–176): The following description is based on an egg taken from the ovipositor of one of the && in the second author’s collection (OC). Capsule basically lens-shaped, laterally compressed and about 1.5x longer than high. General colouration of capsule and operculum greyish mid brown, surface very minutely granulose and slightly rugulose. Whole capsule surrounded by prominent, rugulose dorsoventral keel, beginning and ending at the operculum and only interrupted near the polar-area and the micropylar plate. Polar impression distinct. Ventral and dorsal surfaces of keel each and on each side with a rounded swelling near operculum, and a second swelling near polar end. This swellings dorsally with a rather well defined, impressed dark brown area. Micropylar plate slightly darker brown than capsule, decidedly bilobed and distinctly constricted medially. Micropylar cup small, rounded and placed at polar end of plate. Operculum surrounded by raised rim, oval and very slightly convex. In its centre with a moderately sized, irregularly cone-shaped, dark brown capitulum. Measurements [mm]: length (incl. capitulum) 7.2, length 6.2, height 4.4, width 3.1, length of micropylar plate 1.8. Comments: A comparatively small % PT in the first author’s collection (No. FH 0284-2) was mistaken for a penultimate instar nymph. The two primary types have subsequently been deposited in ZSMC. Distribution (Fig. 389): Philippines (Mindoro Island: Mount Halcon, Barrio Luyang). Number of specimens examined: 10



173

TABLE 24. Measurements [mm] of Phobaeticus philippinicus (Hennemann & Conle, 1997).

%, HT (ZSMC)

&, AT (ZSMC)

%%, PT (coll. FH)

& (nymph), PT

&&

(coll. FH)

(coll. OC)

Body

165.5

247.5

143.0–153.5

134.7

236.0–247.0

Head

4.9

10.1

4.7–4.8

4.7

9.3–9.7

Pronotum

5.0

8.3

4.5–4.8

4.5

7.3–7.8

Mesonotum

36.8

57.0

33.0–34.0

30.0

56.5–56.9

Metanotum

15.4

21.0

13.4–14.0

13.2

22.1–23.3

Median segment

11.3

14.8

9.5–10.2

8.0

13.1–14.2

Profemora

60.5

72.3

55.8–56.5

40.5

70.1–70.9

Mesofemora

41.8

-

40.7–43.5

29.0

54.2–55.3

Metafemora

52.0

-

47.0–52.5

34.8

61.5–63.5

Protibiae

76.0

98.7

76.0–76.2

50.6

104.1–105.0

Mesotibiae

47.9

-

46.7–50.0

33.0

66.2–69.6

Metatibiae

63.0

-

59.8–62.9

42.2

82.0–82.4

Antennae

> 24.5

55.0

> 55.5–71.0

> 18.0

>45.0

Phobaeticus pinnipes (Redtenbacher, 1908) Pharnacia pinnipes Redtenbacher, 1908: 452. Type(s), %(%): Malacca, Selangor (MNHN—not traced, presumed lost) Pharnacia pinnipes, Brock, 1995: 95. Phobaeticus pinnipes, Brock, 1996: 29. Brock, 1999: 136. Seow-Choen, 2000: 39. Otte & Brock, 2005: 269.

Comments: The % type(s) of this species which Redtenbacher (1908: 452) stated to be in MNHN is most certainly lost. Extensive research could not trace any specimens in the collection of MNHN that match with the original description and measurements given by Redtenbacher. Ph. pinnipes is hence not included in the keys above. Redtenbacher originally placed this species in Pharnacia Stål and his distinguishing key to the %% postulates close relation to three species now contained in Phobaeticus Brunner v. Wattenwyl, 1907 (Ph. hypharpax (Westwood, 1859), Ph. serratipes (Gray, 1835) and Ph. kirbyi Brunner v. Wattenwyl, 1907). Indeed, the very elongate and slender semi-tergites of the anal segment and in-curving apices of the cerci place Ph. pinnipes in Phobaeticus Brunner v. Wattenwyl. It is apparently very similar to Ph. magnus nom. nov., but without availability of the type no decision on its systematic position can be made with confirmation. The original description of the % of Ph. pinnipes serves the following characters to distinguish it from %% of Ph. magnus nom. nov.: colouration of the head plain; tegmina lacking white interior markings; meso- and metasternum with three black longitudinal lines; anterodorsal carina of the profemora with only 7–9 serrations; meso- and metafemora with a conspicuously enlarged sub-basal tooth; dorsal carina of the basitarsi strongly raised; probasitarsus white; meso- and metatibiae with a large dorso-median tooth. Etymology: The specific name pinnipes (lat. = lobed feet) refers to the strongly raised dorsal carina of the meso- and metabasitarsi. Description: The following is a roughly translated version of the original Latin description provided by Redtenbacher (1908: 452): “Reddish and yellowish brown. Antennae dark reddish. Vertex smooth, with a pair


HENNEMANN & CONLE

of transverse depressions between the eyes. Tegmina dark red, anterior and posterior margin broadly white. Costal region of alae pale reddish, irregularly coloured, anal region plain greyish. Tergites V–VI with a posteromedial tubercle (less prominent on VII) and slightly posterolaterally elevated. Anal segment keeled and formig two acute, triangular lobes. Apices of cerci incurving. Poculum anteriorly with a raised transverse hump, posterior margin rounded. Meso- and metasternum with a black longitudinal median line and lateral margins with a black longitudinal carina. Complete armature of legs and tarsi black. Anterodorsal carina of profemora with 7–9 large serrations, posteroventral carina with a much larger number of minute teeth. All carinae of meso- and metafemora densely serrate, dorsally with an enlarged tooth near base of each femur. Protibiae dorsally with several large teeth which decrease in size towards the apex of tibia. Meso- and metatibiae with all carinae dentate, dorsally with a distinctly enlarged tooth just before the middle. Basitarsi, but meso- and metabasitarsus in particular with strongly raised dorsal carina. Probasitarsus white, meso- and metabasitarsus with reddish mottling.” Redtenbacher cited the following measurements for the HT [mm]: body 163.0, mesonotum 31.0, metanotum + median segment 22.0, tegmina 10.6, alae 74.0, profemora 42.5, metafemora 48.5.

Phobaeticus redtenbacheri (Dohrn, 1910) (Figs. 110–111, 173–174, 232–233, 309, 390) Nearchus redtenbacheri Dohrn, 1910: 409. LT (by present designation), &: N-Borneo, leg. Waterstradt [larger specimen] (ZMPA); PLT, &: same data as HT (ZMPA). Bragg, 1995b: 276, figs. 6–8 (%), 9–11 (egg). [Description of % and egg] Liana, 1996: 5. Bragg, 2001: 394, figs. 147 & 148, figs. 147 (%), 148 (egg). Baculolonga redtenbacheri, Hennemann & Conle, 1997: 348. Phobaeticus redtenbacheri, Otte & Brock, 2005: 269. Nearchus maximus redtenbacheri, Günther, 1935a: 11. Further material: [2 &&, 1 %, 1 egg]: N-BORNEO, SABAH: 1 %: Sabah, Mt. Kinabalu Park, near Park Headquarters, Silau-Silau trail, 1550 m, leg. Hennemann & Conle 4.– 8.VIII.1996 (coll. FH, No. 0264-1); 1 egg: Sabah, Mt. Kinabalu NP, near HQ, 1550 m, laid by PEB-1719 (coll. PEB, No. 1720); 1 &: Borneo, Kinabalu ca. 1500 m, leg. Waterstradt, H. Rolle vend. 25.11.1904 (ZMUH); 1 &: Sabah, Mt. Kinabalu, Mesilau, 6.II.1964, J. Smart leg., Royal Soc. Exped. BMNH, 1964-250 (BMNH).

Diagnosis: Closely related to the Bornean Ph. mjoebergi (Günther, 1935), Ph. kirbyi Brunner v. Wattenwyl , 1907 and Ph. chani Bragg spec. nov. and Ph. philippinicus (Hennemann & Conle, 1997) from the Philippines. With Ph. mjoebergi && share the long and lanceolate subgenital plate but differ by: the larger size; relatively shorter mesonotum; characteristic armature of the legs and dentate medioventral carina of the meso- and metafemora. The long and lanceolate subgenital plate (Fig. 232) and long median segment clearly distinguishes && from those of the latter three species. From Ph. kirbyi it is furthermore readily distinguished by: the less globose and slightly more elongate head of both sexes; less distinct praeopercular organ and characteristic armature of the legs of &&, as well as the serrate posteroventral carina of the profemora; lack of a black ventrolateral marking on the cheeks; shorter alae and longer, more elongate semi-tergites of the anal segment of %% (Fig. 233). The eggs differ by lacking radial ridges on the dorsoventral keel of the egg capsule, but instead have rounded swellings, and relatively smaller micropylar plate (Figs. 173–174). From Ph. chani Bragg spec. nov. it clearly differs by: the shorter, more globose head and different leg armature of both sexes; considerably smaller size; more robust body and relatively shorter body segments of &&; lack of a dark postocular stripe; much shorter alae; more elongate



175

legs and tarsomeres, and long, slender semi-tergites of the anal segment of %%. The eggs at once differ by lacking the conspicuous leaf-like appendages of the dorsoventral keel seen in Ph. chani Bragg spec. nov.. With Ph. philippinicus && have the apical dorsal lobe of the protibiae in common and eggs are rather similar in general appearance. Features which readily distinguish Ph. redtenbacheri from this species are the clearly different leg armature of both sexes and presence of tegmina and alae in the %%. Etymology: Dedicated to the well known Austrian entomologist Josef Redtenbacher (1856–1926), who described several hundreds of new species of Phasmatodea in the well known monograph “Die Insektenfamilie der Phasmiden” (Redtenbacher, 1906 & 1908), amongst them several of the tribe Pharnaciini. Description: && (Fig. 110): Very long (body length 249.0–256.0 mm, including subgenital plate 275.0– 300.0 mm) and slender for the genus (maximum body width ± 6.0 mm) with an extremely long, lanceolate subgenital plate (56.0–62.5 mm). General colouration of body and legs more or less uniformly dull green or mid to dark brown. Leg armature dull red to reddish brown. Eyes dark ochracheous. Antennae dark reddish brown, the two basal segments dull green or brown. Head: Slightly longer than wide, broadest at eyes and distinctly narrowing towards the posterior, vertex very gently rounded and with a faintly impressed coronal line. Eyes small, circular and projecting hemispherically; their length contained about 3x in that of cheeks. Antennae slightly projecting over posterior margin of mesonotum. Scapus 3x longer than wide, dorsoventrally flattened, rectangular. Pedicellus cylindrical and distinctly shorter than scapus. Thorax: Pronotum almost 2x longer than wide, as long but narrower than head and with a small rounded posteromedian tubercle. Median transverse depression moderately distinct, slightly curved and almost reaching to lateral margins of segment. Mesothorax almost 3.8x longer than head and pronotum combined, mesonotum parallel-sided and very slightly constricted at anterior margin. Metanotum almost 3x longer than wide, parallel-sided. Meso- and metasternum simple. Abdomen: Median segment a little longer than metanotum, about 3x longer than wide and slightly widened towards the posterior. Abdominal segments II–VII increasing in length, II 2x, III 2.5x, IV 3x, V–VI 3.5x longer than wide. VII slightly narrower than previous, as long as IV and about 4x longer than wide. Tergites II–VI with a small posteromedian tubercle. Praeopercular organ formed by two short, blunt spines at posterior margin of sternum VII. Tergite VIII narrower than previous, about 2.5x long than wide, about 3/5 the length of VII. IX almost quadrate, strongly convex. Anal segment as long as IX, with a blunt median carina and slightly constricted towards the apex; posterior margin with a rounded median indention. Supraanal plate bilobed, projecting slightly over posterior margin of anal segment. Cerci small, conical. Subgenital plate very long, lanceolate, tapered towards a pointed tip and projecting over anal segment by at least the combined length of tergites VII–X (Fig. 233). Legs: All very long and slender, profemora as long as head, pro- and mesothorax combined, mesofemora about as long as pro- and mesothorax combined, metafemora almost reaching to posterior margin of abdominal segment V and metatarsi roughly reaching tip of subgenital plate. Profemora on anterodorsal carina with 15–20 acute triangular serrations and a similar number of smaller serrations on posteroventral carina. Protibiae occasionally with one ± enlarged, triangular median tooth on the posterodorsal carina and with a small, rounded lobe at the apex. Medioventral carina of meso- and metafemora set with a few minute spines. All other carinae armed with numerous pointed teeth; considerably larger and spine-like on ventral carinae. Posteroventral carina usually with one or two considerably enlarged spines sub-basally. Posterodorsal carina with 2–4 slightly larger, triangular teeth. Meso- and metatibiae with all carinae minutely serrate. Mid point of posterodorsal carina with a broad, triangular tooth and a small, rounded apical lobe. Probasitarsus slightly longer than remaining tarsomeres combined, unarmed. Meso- and metabasitarsi with dorsal carina slightly raised and all carinae minutely dentate, a little longer than following three tarsomeres combined. %% (Fig. 111): Medium sized to rather long (body length 146.0–161.5 mm) and very slender for the genus with considerably shortened alae (28.5–29.0 mm). General colouration of body and legs mid green (often


HENNEMANN & CONLE

brown when preserved), dorsal surface of abdomen, tegmina and costal region of the alae pale greenish brown. Anterior margin of tegmina slightly whitish. Anal region of alae transparent grey with brown veins. Eyes reddish brown. Armature of the legs dark yellow. Antennae dark brown with the ventral surface black. Head (Fig. 309): Oval, broadest at eyes and strongly narrowed towards posterior margin, vertex very gently rounded. Eyes large and projecting hemispherically from head capsule; their length contained less than 2x in that of cheeks. Antennae reaching to posterior margin of abdominal segment II and consisting of about 20 segments. Scapus dorsoventrally flattened, 3x longer than wide, rectangular. Pedicellus less than half the length of scapus, cylindrical. Segments in median portion of antennae extremely elongate. Thorax: Pronotum slightly shorter and narrower than head, constricted in anterior portion, anterior margin and lateral margins slightly raised; otherwise as in &&. Mesothorax about 3.7x longer than head and pronotum combined, mesonotum parallel-sided except for being gently widened at posterior margin. Meso- and metasternum with a distinct longitudinal median keel. Tegmina slender and elongate, spatulate, strongly constricted basally and with a very faint central hump, just covering base of alae. Alae ± reaching to posterior margin of abdominal segment II. Abdomen: Median segment about 2x longer than metanotum. Abdominal segments II–V very slightly increasing in length, II about 5.5x, V almost 7x longer than wide. VI as long as V, VII 3/4 the length of VI. Tergites IV–VI with a small pointed, scale-like posteromedian tubercle (most prominent on V). Tergite VIII 2/ 3 the length of VII and gradually widening towards the posterior. IX slightly shorter than VIII and constricted medially, anterior margin broader than posterior margin. Anal segment very long, longer than VII, strongly keeled, laterally compressed and tectiform. Semi-tergites very elongate, slender, straight and with the apex slightly pointed (Fig. 232). Interior surfaces apically densely covered with minute hooked spines. Cerci of moderate length, very slender, strongly in-curving and tapered towards the apex. Poculum strongly convex, cup-like, angular with an acute, transversely compressed backward pointing central spine. Legs: All very long and slender with all carinae more or less distinctly serrate, except dorsal carinae or protibiae. Profemora distinctly, mesofemora slightly longer than head, pro- and mesothorax combined, metafemora slightly projecting over posterior margin of abdominal segment V and metatibiae considerably exceeding apex of abdomen. Anterodorsal carina of profemora with 18–24 distinct and acutely triangular serrations, posteroventral carina with a similar number of very small teeth. Two outer ventral carinae of mesoand metafemora distinctly armed with triangular teeth of rather uniform size, dorsal carinae very minutely dentate. Medioventral carina unarmed. All carinae of meso- and metatibiae minutely serrate, although very minutely and sparsely on dorsal carinae. Probasitarsus very long and slender, considerably longer than remaining tarsomeres combined, unarmed. Meso- and metabasitarsi about as long as remaining tarsomeres combined, all carinae with 2–4 minute teeth. Eggs (Figs. 173–174): The following description is based on an egg kindly provided for examination by P.E. Bragg (Nottinghamshire, coll. PEB, No. 1720). Capsule basically lens-shaped, laterally compressed, slightly longer than high. General colouration of capsule and operculum mid brown, surface slightly rugulose. Whole capsule surrounded by dorsoventral keel, beginning and ending at the operculum and only interrupted near the polar-area and the micropylar plate. Polar impression wide. Ventral and dorsal surfaces of keel each with a rounded swelling near operculum, and a second smaller swelling near polar end. Micropylar plate lighter brown than capsule, slightly bilobed and constricted medially. Micropylar cup small, rounded and placed at polar end of plate. Operculum surrounded by raised rim, oval and slightly convex. In its centre with a small, roughly cone-shaped, dark brown capitulum. Measurements [mm]: length (incl. capitulum) 6.0, length 5.5, height 4.5, width 3.0, length of micropylar plate 1.7. Comments: Dohrn (1910: 409) originally described Nearchus redtenbacheri based on a & specimen with a body length of 300.0 mm and referred to a second, smaller specimen (275.0 mm) which he stated to be iden-



177

tical except for being considerably smaller. Bragg (2001: 397) respectively treated these as HT and PT. This kind of treatment however is not in accordance to the Code (ICZN 1999: Articles 73–74.), and from the specimen treated as HT and PT by Bragg (2001) the first larger specimen is here designated as the LT, the other smaller specimen being the PLT. Although Dohrn’s type specimens in ZMPA could not be examined for this study the designation of a LT appears necessary to guarantee stability of the specific name. The original description and examination of a & in ZMUH from the same locality and collector (Fig. 110) leave no doubt about its identity. Bragg (1995b & 2001) provided a redescription of the & along with descriptions and illustrations of the % and egg. The author mentioned this species to be not rare near the headquarters of Mount Kinabalu National Park (N-Sabah) with three adult && encountered at this locality during only eight nights (Bragg, 2001: 397). In addition to the material listed above there are a further two %% and && from Mount Kinabalu in the collection of P. E. Bragg (Nottinghamshire, coll. PEB) and several couples from the same locality in the collection of C. L. Chan (Kota Kinabalu, coll. CLC). Günther (1935: 10–11) erroneously interpreted the two Bornean N. redtenbacheri Dohrn, 1910 and N. mjoebergi Günther, 1935 as subspecies of Nearchus maximus Redtenbacher, 1908 (= Phobaeticus magnus nom. nov.) which is however obviously a distinct species and restricted to continental Asia (Thailand, Laos, NE-India and N-Myanmar). Distribution (Fig. 390): Northern Borneo: Sabah (Mount Kinabalu: National Park HQ 1500–1580 m & Mesilau). Number of specimens examined: 3 TABLE 25. Measurements [mm] of Phobaeticus redtenbacheri (Dohrn, 1910).

& HT (ZMPA)*

&&

%%


300.0

275.0–284.0

-

Body

-

249.0–256.0

146.0–161.5

Head

-

9.2–9.5

4.5–5.0

Pronotum

-

7.7–8.0

4.0–4.9

Mesonotum

58.0

52.5–56.5

36.0–38.7

Metanotum

35.0**

16.7–18.0

7.0–8.5

Median segment

-

18.3–19.5

15.0–16.5

Tegmina

-

-

9.0–10.0

Alae

-

-

28.5–29.0

Subgenital plate

56.0

56.0–62.0

-

Profemora

82.0

75.0–78.0

51.0–60.5

Mesofemora

-

57.0–57.0

38.0–45.5

Metafemora

74.0

66.5–68.0

47.0–54.0

Protibiae

118.0

105.5–107.0

69.0–87.5

Mesotibiae

-

65.0–66.0

42.0–51.5

Metatibiae

88.0

84.0–85.0

55.0–70.0

Antennae

-

> 67.0

> 78.0–83.5

* according to Dohrn (1910: 410) ** including median segment


HENNEMANN & CONLE

FIGURES 120–125: Phobaeticus spp. 120. Phobaeticus lumawigi Brock, 1997, & HT, Philippines, N-Luzon Id., Mountain Province (BMNH) 121. Phobaeticus sobrinus Brunner v. Wattenwyl , 1907, % LT, Sumatra, Si-Rambé (MCSN) 122. Phobaeticus palawanensis spec. nov., % HT, Palawan, Salakot Pass, Napsan (ZSMC) 123. Phobaeticus philippinicus (Hennemann & Conle, 1997), % HT, Philippines, Mindoro Id., Mount Halcon (ZSMC) 124. Phobaeticus hypharpax (Westwood, 1859), & Sri Lanka (SMFM) 125. Phobaeticus lobulatus (Carl, 1913), & HT, Sri Lanka (MHNG) REVISION OF PHARNACIINI


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Phobaeticus rex (Günther, 1928) comb. nov. (Figs. 118, 265, 270, 331, 391) Eucarcharus rex Günther, 1928: 218. HT, &: Sumatra, Redjang-Lebong, leg. H. Kubale S.; Eucarcharus rex Günther, 1928 (MNHU). Lobophasma rex, Günther, 1935b: 138. Günther, 1953: 555. Zompro, 2005b: 279. Otte & Brock, 2005: 178. Phobaeticus lambirica Seow-Choen, 1998a: 40, fig. 1 (& and egg). HT, &: Malaysia, Sarawak, Lambir National Park, Dec. 1996, I. Abercrombie (SFDK) syn. nov. Otte & Brock, 2005: 269. Baculolonga lambirica, Bragg, 2001: 384.

Diagnosis: Easily distinguished from all other members of the genus by the remarkable long and leaf-like, posteriorly dentate lateral lobes of abdominal tergite VII, which project by distinctly more than the body width and long supraanal plate. Etymology: “Rex” (lat. = king) refers to the great size of this striking species. Description: && (Fig. 118): Very long (body length 247.0–260.0 mm) and moderately slender species for the genus (7.0–8.0 mm). General colouration of body light to mid brown with large areas of white on head, thorax, abdominal tergites and all legs, occasionally greenish areas near anterior and posterior margin of the mesonotum and lateral regions of the metanotum. Head generally whitish with a bold transverse dark brown band along posterior margin and a pair of black, roughly C-shaped spots between the eyes. Most parts of meso-, metanotum, median segment, tergites III and V–X whitish. Tergite II with two bold black converging stripes and a large, white and wedge-shaped marking at the posterior. Tergite IV with a pair of similar but smaller black markings. Ventral surface of body generally orange brown with irregular greenish and whitish speckles. Meso- and metasternum with a green longitudinal median line, the latter with two elongate greenish white markings. Abdominal sternites more or less distinctly mottled with white. Legs white and distinctly marbled with grey or brown. Tarsi yellowish brown with darker mottling. Eyes sepia with darker brown speckles. Antennae reddish brown, except for two basal segments. Head: Elongate, oval, almost 2x longer than wide, cheeks almost parallel-sided, vertex gently convex in posterior portion. A distinct transverse depression between the bases of the antennae. Eyes small, circular and moderately convex; their length contained about 3.5x in that of cheeks. Antennae at least with 29 joints and reaching 1/3 the way along mesonotum. Scapus dorsoventrally flattened, parallel-sided, 3x longer than wide and almost 3x longer than pedicellus; the latter round in cross-section. Thorax: Pronotum shorter and narrower than head, almost rectangular and parallel-sided, the anterior portion very indistinctly constricted; about 1.5x longer than wide. Median transverse depression indistinct, straight and not reaching lateral margins of segment. Mesothorax about 2.8x longer than head and pronotum combined, mesonotum parallel-sided except for being very gently widened at posterior margin. Metanotum about 2/5 the length of mesonotum, 3x longer than wide. Meso- and metapleurae with a very fine longitudinal median carina. Meso- and metasternum simple. Abdomen: Median segment ¾ the length of metanotum, parallel-sided, about2.4x longer than wide. Abdominal segments II–VI increasing in length, II about equal in length to median segment and 2x longer than wide, IV 3.5x longer than wide. V as long as IV, VI as long as III. Tergites II–VII with posterior margin roundly arcuate and forming a blunt posteromedian tubercle. VI and VII with an indistinct longitudinal median carina. Lateral margins of VII posteriorly protruded into a very long, leaf-like lobe which projects by considerably more than the body width and reaches as far back as to the middle of VIII (Fig. 270); posterior margin roughly dentate. Sternites II–VI with an indistinct blunt median carina. Praeopercular organ formed by two small spiniform tubercles at posterior margin of sternum VII. Tergite VIII distinctly narrower than previ-


HENNEMANN & CONLE

ous, about ¾ the length of VII, strongly convex, constricted medially and about 3.5x longer than wide. IX indistinctly longer than wide, half the length of VIII, rectangular. Anal segment slightly longer than IX and with a fine median carina, posterior margin with a broad, trapezoidal excavation. Supraanal plate elongate, keeled and projecting distinctly over apex of anal segment. Subgenital plate strongly convex, boat-like, longitudinally keeled and not reaching to apex of supraanal plate (Fig. 265). Legs: All long and moderately robust, meso- and metatibiae gently down-curving. Profemora about as long as mesothorax, mesofemora distinctly shorter than mesothorax, metafemora slightly projecting over posterior margin of abdominal segment IV and metatarsi slightly projecting over apex of abdomen. Anterodorsal carina of profemora with 10–12 very prominent, lamellate serrations of variable size, some of which may be distinctly enlarged and foliaceous posteroventral carina armed with ± 15 moderately distinct, acute teeth and elevated into a dentate lobe apically. Protibiae with anterodorsal carina distinctly but irregularly elevated and broadly, acutely wave-like. Posteroventral carina densely but rather minutely serrate. All carinae of mid and hind legs rather regularly dentate, less distinctly on dorsal carinae. Meso- and metafemora with a few minute spines on medioventral carina. Mesofemora with one or two prominent, long sub-basal spines on posteroventral carina and a broadened triangular tooth at same height on posterodorsal carina (Fig. 331). Metafemora without any larger spines or teeth. Posterodorsal carina of meso- and metatibiae with a prominent, broad triangular lobe medially and a deeply dentate lobe apically. Probasitarsus as long as remaining tarsomeres combined except claw, dorsal carina distinctly raised but unarmed, ventral carinae dentate. Meso- and metabasitarsi as long as following three tarsomeres combined, dorsal carina forming a considerably raised and rounded lobe; all carinae dentate. TABLE 26. Measurements [mm] of Phobaeticus rex (Günther, 1928) comb. nov.

&, HT (MNHU)

&, HT of P. lambirica (SFDK)*

Body

247.0

260.0

Head

13.0

8.0

Pronotum

9.0

8.0

Mesonotum

53.0

55.0

Metanotum

20.5

20.0

Median segment

16.5

15.0

Profemora

54.5

54.0

Mesofemora

48.0

45.0

Metafemora

58.0

54.0

Protibiae

65.0

70.0

Mesotibiae

48.0

46.0

Metatibiae

68.0

67.0

Antennae

> 30.0

40.0

* according to Seow-Choen (1998a: 41)

Eggs: Seow-Choen (1998a: 41) briefly mentioned and illustrated the eggs in lateral view and provided the following brief characterization: “The eggs of the new species is in the mould of a typical egg of other Phobaeticus species. The egg itself is very dark grey almost black and measures 6 mm by 3.8 mm. It is rather more globular than the egg of P. serratipes (Gray) and P. kirbyi Brunner and is 3.8 mm thick.” The author mentioned the egg to be typical for the genus and although only illustrated in lateral view the capsule is seen to be



181

surrounded by a distinct dorso-ventral keel and notched near the polar-area. The capitulum is small and knoblike, the operculum oval. Unfortunately Seow-Choen did not probvide a description of the micropylar plate. Measurements according to Seow-Choen (1998a: 41) [mm]: length 6.0, width 3.8, height 3.8. Comments: Günther (1928: 218) presented a very detailed description and illustration of the HT of his Eucarcharus rex from SE-Sumatra in MNHU. Subsequently, Günther (1935b: 139) designated E. rex as the type species of his new genus Lobophasma, which is a junior synonym of Phobaeticus Brunner v. Wattenwyl , 1907 (syn. nov.). The HT of the Bornean Phobaeticus lambirica Seow-Choen, 1998a was only examined from colour photographs of the live insect. Comparison with the HT of Eucarcharus rex in MNHU however has clearly shown these two taxa to represent the same striking and characteristic species (syn. nov.). The disjunct distribution in SE-Sumatra and W-Borneo is remarkable. % unknown. Distribution (Fig. 391): SE-Sumatra (Redjang-Lebong) and W-Borneo (Sarawak: Lambir National Park). Number of specimens examined: 1

Phobaeticus serratipes (Gray, 1835) (Figs. 76, 112–113, 169–170, 243–245, 305–306, 332–333, 343–344, 354, 360, 392, 420) Cladoxerus serratipes Gray, 1835: 42. HT, %: 40-42-1316, Mal.; Type; serratipes G.R. Gray; Malaysia, Cladoxerus serratipes Gray, Cladoxerus serratipes Malabar (BMNH). Phibalosoma serratipes, Westwood, 1859: 75. Stål, 1875: 63. Pharnacia serratipes, Kirby, 1904a: 359. Hennemann, 1993: 45, figs. 1–6. Brock, 1995: 94. Seow-Choen, 1995: 12. Lipinski et al., 1999: 80, fig. 36a–h (REM-study of egg). Phobaeticus serratipes, Brock, 1996: 29. Seow-Choen, 1996: 32. Seow-Choen, 1997: 120, figs. 102–103. Seow-Choen, 1998a: 41. Seow-Choen, 1998b: 89. Brock, 1999: 133, 182, figs. 89a–b (%), 90a–b (&), 90c (egg), pl. 25 (egg), 27 (&). Seow-Choen, 2000: 38, pl. 96 (%, &, egg). Otte & Brock, 2005: 269. Baculolonga serratipes, Hennemann & Conle, 1997a : 347, figs. 8–9 (egg). Bragg, 2001: 385, fig. 145 (%, egg) Bacteria acanthopus Burmeister, 1838: 565. HT, &: 749; acanthophus Westw.*, Bacteria acanthopus Burm.*, Singapore (MNHU). [Synonymised by Kirby, 1904: 359] Zompro, 2005b: 253. Phibalosoma acanthopus, Westwood, 1859: 74. Stål, 1875: 63. Pharnacia acanthopus, Redtenbacher, 1908: 454, pl. 21: 8 (%). Giglio-Tos, 1910: 43. Werner, 1934: 2. Herbert, 1988: 9. Carlberg, 1988: 17. Carlberg, 1989a: 9. Carlberg, 1989b: 159. Bactridium grande Rehn, 1920: 242, pl. 10: 12 (&). HT, &: Sta. Catharina, Brazil; Bactridium grande Rehn Holotype; No. 401 (ANSP). [locality erroneous] syn. nov. Heteronemia grande, Otte & Brock, 2005: 158. Phibalosoma maximum Bates, 1865: 341. HT, &: Sumatra, Wallace; Phibalosoma maximum Bates; Type, Bates, Phiba-


HENNEMANN & CONLE

losoma maximum; E. coll. (1830-73), W.W. Saunders, Purchased and pres.´73 by Mrs. F.W. Hope (OXUM, Nr. 622) syn. nov. Pharnacia maxima, Kirby, 1904a: 359 (in part). Redtenbacher, 1908: 450. [Only &&; % is Tirachoidea jiandenglingenis (Bi, 1994)] Phobaeticus maximus, Otte & Brock, 2005: 269. Tirachoidea maxima, Karny, 1923: 240. Tirachoidea ajax Redtenbacher, in litt. [Specimens in ZMUH] [Not: Pharnacia serratipes, Kirby, 1896: 450. Misidentification relating to Phobaeticus kirbyi Brunner v. Wattenwyl, 1907] [Not: Phasma (Cladoxerus) acanthopus, de Haan, 1842: 131. Misidentification relating to Pharnacia biceps Redtenbacher, 1908: 451] [Not: Phibalosoma acanthopus, Wood-Mason, 1877a: 161. Misidentification. Described as Pharnacia ingens Redtenbacher, 1908: 453 but & PLT from Myanmar (Upper Tenasserim: Moolai to Moolat) is Phryganistria grandis Rehn, 1906] [Not: Pharnacia maxima, Kirby, 1904a: 359. Misidentification relating to Phobaeticus kirbyi Brunner v. Wattenwyl, 1907] Further material: [40 &&, 34 %%, 10 nymphs, eggs]: PENINSULAR MALAYSIA: 5 &&, 4 %%: Grubauer 1902, Kwala Kangsar, Perak, Mus. Caes. Vindobon ; 24.459 & 23.811 (NHMW, No. 454); 2 &&: Coll. Br. v. W., Malacca, Perak ; 25.136 (NHMW, No. 454); 1 % (nymph): Coll. Br. v. W., Malacca, Perak (NHMW, No. 454); 2 &&, 2 %%: W-Malaysia, Perak, leg. Grubauer, B. Jachau vend. 15.12.1901 (ZMUH); 1 &, 1 %: W-Malaysia, Perak, leg. Kinstler 31.I.1887 (ZMUH); 1 &, 1 %, 2 && (penultimate instar), 1 % (penultimate instar): Perak, Kwala Kangsar, B. Jachau vend. 1.4.1901 (ZMUH); 1 %: Camp Jor, Wasserscheide zw. Perak und Pahang (Malakka-Gebiet), Alb. Grubauer leg.; vend. 30.12.1902 (ZMUH); 1 %: W-Malaysia, Perak, Tapah Hills, ca. 400 m, via Wong T.F. 10. 1993 (coll. FH, No. 0030-1); 3 &&, 5 %%: ex Zucht: F: Hennemann 1994, urspr. W-Malaysia, Perak (coll. FH, No’s 0030-2 to 9); 1 &, 1 %: ex Zucht. F. Hennemann 1997, urspr.: W-Malaysia (coll. FH, No’s 0030-10 & 11); 2 &&, 1 %, 1 % (nymph): ex. Zucht O. Conle 1994, urspr. W-Malaysia (coll. OC, No’s 00201-204); 1 %: Penang, 96-85, "Hind wings used for drawings (Ragge)" (BMNH); 1 %: Perak 1904-117 (BMNH); 1 &: Perak 1904-117, Pharnacia acanthopus, Kuala Kangsar, Perak (BMNH); 1 &: Penang (BMNH); 3 &&, 3 %%: W-Malaysia, Cameron Highlands 2000ft., 20.–30.5.1972, leg. C.C. Chua, BM 1975-604 (BMNH); 1 %: Perak, F.M.S., Batang-Padang, Kuala Woh, "at light", ex. F.M.S. Mus., BM 1955-354 (BMNH); 3 &&, 1 % (nymphs): Cameron Highlands 2000ft, 3.4.1972–23.2.1973 (BMNH); 1 %: Malay, Penin. Jos. Camp 2000ft., 19.9.1922 (BMNH); 1 %: Malay, Penin, Kedah Peak 3300ft., 26.3.1928, ex. F.M.S. Mus., BM 1955354 (BMNH); 1 %: Penang, leg. Cantor (OXUM); 1 &: Perak, Kwala-Kangsar (MHNG); 2 &&: Cameron Highlands, Perak, Malaysia, 1200–1600 m, achat Pfanner 1970 (MHNG); 1 &: Malaisie, Perak, Cameron Highl., 1200–1600 m, leg. Pfanner (MHNG); 2 %%: Malaisie, Elevage 1986, Don N. Mal (ISNB); 1 &: Malaya, Kuala Lumpur, Perak, Taikong, Aug. 1931, H.M. Pendlebury (RMNH); 1 & (nymphs): Malay Penin., Kedah, Perak, 3.300 ft., 25.3.1928; 1 & (nymph): Malay Penin., Kedah, Perak, 3.300–3.978 ft., 29.3.1928 (RMNH); 1 &: Malaysia (Penins. Malacca), Perak, Gopeng, II. 1983, (acqistato indeterminato da G. Russo, Bologna, nel 1985); Pharnacia acanthopus (Burmeister), det. R. Poggi 1985 (MCSN); 1 &: Perak, Malacca, coll. Giglio-Tos (MRSN); 1 %: Malacca, Künstler leg. (MNHU); 2 &&: Perak, Malakka; Collection A. Finot, Phryganistria sarmentosa Westwood (MNHN, coll. Finot, Box-No. 279); 1 &: Sungei-Sipoet, Perak (ZSMC). SUMATRA: 1 %: Sumatra, Medan, Siboelangit, Batak Plateau, leg. Grubauer; B. Jachau vend. 1899 15.7.1900 (ZMUH); 1 %: Sumatra, Indragiri, leg. W. Burchard 18.XII.1905; Tirachoidea ajax Redt., Type. Männchen Type in Hauptsammlung, nomen nudum ! (ZMUH); 1 &: Sumatra, Indragiri, Somgai-Lulah, leg. W. Burchard 20.7.1904; Tirachoidea ajax Redt., Type. Männchen Type in Hauptsammlung, nomen nudum ! (ZMUH); 1 %: NO-Sumatra, Banda-Kwala, leg. P. Puttfurchen 5. 8.1898 (ZMUH); 1 &: Sumatra, Padang distr., leg. Van der Poll, Coll. Brought of Janson 1909, 1909-695, Buy. Proepoe, Pad. Bovenland, ± 1600, 5.-8.´98 (Collector D.) (OXUM); 1 & (nymph): Indrapoera, Sumatra, leg. Weyers (ISNB); 1 %: Tamjang, Sumatra, V. Nill leg., det. W. Potvin (ISNB); 1 %: Palambay, Sumatra; "Cladoxerus acanthopus" K. Günther det. (RMNH); 1 &: Sumatra, Modigliani; Best. Verz. Nr. 76 (MCSN); 1 % (nymph): Sumatra, D. Martin, (95/ 6) (ZSMC); 1 %: Sumatra, ex coll. Le Moult (MNCN). NO DATA: 1 %: aus Zucht (MNHU); 1 %: ex Zucht: O. Zompro, det. O. Zompro VI.1993 (MNHU); 1 %: no data (MNHN); 1 &: Ent.-Club 44-12 (BMNH).



183

Diagnosis: Related to Ph. kirbyi Brunner v. Wattenwyl , 1907 and Ph. chani Bragg spec. nov. from Borneo and the Sumatran Ph. sobrinus Brunner v. Wattenwyl , 1907. From the first species it is readily distinguished by: the more elongate and flat head (Figs. 305–306); densely serrate posteroventral carina of the profemora of both sexes (Figs. 343–344); longer subgenital plate which roughly reaches the apex of the anal segment (Fig. 243) and longer, more slender lobes of the praeopercular organ of &&; more slender and parallel-sided semitergites of the anal segment (Fig. 245) and lack of a black ventrolateral marking on the cheeks of %% (Fig. 306); as well as the kidney-shaped micropylar plate and lack of radial ridges on the posteroventral keel of the egg capsule (Figs. 169–170). From Ph. chani Bragg spec. nov. it at once differs by: the smaller size and less elongate, more robust body; relatively shorter body segments; much more prominent triangular lobes of the praeopercular organ; lack of a conspicuous sub-basal spine on the two outer ventral carinae of the meso- and metafemora (Figs. 332–333) and more numerous serrations of the profemora of &&, as well as the considerably shorter alae, relatively longer legs and much more elongate and slender semi-tergites of the anal segment of %% (Fig. 245). Eggs clearly differ by lacking the leaf-like appendages of the dorsoventral keel of the eggcapsule seen in Ph. chani Bragg spec. nov.. From the smaller Ph. sobrinus it can be distinguished by: the longer median segment of both sexes; lack of the prominent lobes on the dorsal and ventral carinae of the protibiae, mesofemora as well as the meso- and metatibiae of &&; presence of tegmina and alae, more distinct leg spination; more acutely pointed hump of the poculum and roundly angulate anterodorsal crest of the anal segment of %% (Fig. 245). Etymology: The specific name serratipes is a combination of the two Latin words “serratus” (= serrate) and “pes” (= foot) to refer to the distinct black serrations of the extremities of %%, the original sex described. Description: && (Fig. 76, 112): Very long (body length 228.0–278.0 mm) and slender species (maximum body width 5.0–6.0 mm), subgenital plate not extending over tip of abdomen. Variable in size, colouration and armature of the mid legs. General colouration of body and legs pale to dark green or brown, either uniformly coloured or with darker speckles. Occasionally green specimens may have the anterior portion of abdominal VIII brown to dark brown. Sometimes specimens occur which have distinct white dorsal markings or most of the dorsal body surface white. Along lateral margins of the mesonotum with a fine, orange longitudinal line (Fig. 420). Armature of legs orange or pale brown in green specimens and dull reddish brown in brown specimens. Eyes pale ochracheous with sepia brown mottling. Antennae mid to dark brown and blackish ventrally. Scapus and pedicellus mid brown. Head (Fig. 305): Elongate, oval in cross-section, 1.5x as longer than wide and slightly narrowing towards the posterior. Vertex flat and in posterior half with a slightly impressed longitudinal lines. Between the bases of the antennae with an almost straight, but decided transverse depression. Eyes of moderate size, circular and projecting strongly from head capsule; their length contained about 2.3x in that of cheeks. Antennae reaching to posterior margin of metanotum. Scapus dorsoventrally flattened, more than 3x as longer as wide and very gently constricted towards the base, 2.5x longer than pedicellus. Pedicellus round in cross-section and indistinctly longer than wide. III about as long as pedicellus, following antennomeres first increasing then decreasing in length towards apices of antennae. Thorax: Pronotum slightly narrower and indistinctly shorter than head, gently constricted pre-medially and posterior margin a little broader than anterior margin. Transverse median depression distinct, slightly curved but not reaching lateral margins of segment. Mesothorax 3.1–3.2x longer than head and pronotum combined, mesonotum parallel-sided. Metanotum about 1/3 the length of mesonotum, 3.5x longer than wide and rectangular. Meso- and metasternum simple. Abdomen: Median segment slightly shorter than metanotum, 3x longer than wide and gently constricted medially. Segments II–VI slightly increasing in length, II 2.3x, IV about 3x and VI 4x longer than wide; VII as long as III. Tergite VII with lateral margins gently rounded in posterior portion. Praeopercular organ formed by two prominent triangular, carinate and slightly sickle-shaped lobes at posterior margin of sternum VII (Fig. 360). Tergite VIII distinctly narrower than previous, ¾ the length of VII, slightly broadened before


HENNEMANN & CONLE

apex and roughly 3x longer than wide. IX rectangular, half the length of VIII and about as long as wide. Anal segment longer than IX, with a fine longitudinal median carina and a widely triangular median excavation at posterior margin, posterolateral angles rounded (Fig. 244). Subgenital plate longitudinally keeled, boatshaped and roughly reaching to apex of anal segment (Fig. 243). Legs: All very long and slender, profemora about as long as head, pro- and mesothorax combined, mesofemora a little longer than mesothorax, metafemora reaching about ¾ the way along abdominal segment V and metatibiae ± reaching apex of abdomen. Anterodorsal carina of profemora distinctly raised and armed with 20–30 prominent and acutely triangular serrations (Fig. 343). Posteroventral carina with a similar number of considerably smaller but pointed triangular teeth. Dorsal and posteroventral carinae of protibiae very indistinctly and sparsely serrate. Dorsal carinae of meso-. And metafemora only with 2–5 teeth each; single or all of the teeth on the posterior carina of the mesofemora often ± enlarged or sometimes forming triangular, lobe-like teeth (Figs. 332–333). Two outer ventral carinae of meso- and metafemora densely and regularly set with pointed teeth of moderate size. Medioventral carina indistinct and armed with a few small spines. All carinae of meso- and metatibiae densely serrate, the dorsal carinae less distinctly although. Posterodorsal carina forming a rounded, dentate lobe at the apex of each tibiae; on mesotibiae occasionally with a ± prominent, triangular tooth medially. Probasitarsus about twice the length of remaining tarsomeres combined, with the dorsal carina distinct but uniformly elevated, a few very minute teeth may be present on ventral carinae. Mesobasitarsi a little shorter, metabasitarsi slightly longer than remaining tarsomeres combined, dorsal carina raised and gradually raised towards the apex, all carinae minutely serrate. %% (Fig. 113): Medium-sized to large (body length 133.5–176.5 mm) and very slender for the genus with shortened alae (36.5–42.0 mm) and characteristic black leg armature. Body greenish pale to mid brown, legs greyish-green, tegmina and costal region of alae greyish brown. Lateral margins of mesonotum with a black longitudinal line and a washed, slightly broader blue longitudinal stripe interiorly. Anterior margin of tegmina and alae broadly white and interiorly bounded by a fine longitudinal black stripe. Anal region of alae transparent pale brown with brown veins. Leg armature distinctly black. Eyes reddish ochracheous with brown mottling. Antennae greyish mid to dark brown dorsally and dark brown to black ventrally. Head (Fig. 306): Generally as in && but more distinctly narrowed towards the posterior. Eyes considerably more prominent, relatively larger and projecting hemispherically; their length contained only about 1.8x in that of cheeks. Antennae projecting over abdominal segment III, otherwise as in &&. Thorax: Pronotum distinctly narrower than head, 1.5x longer than wide and trapezoidal, being narrowed towards the anterior. Median transverse depression distinct, gently curved but not reaching to lateral margins of segment. Mesothorax about 3.2x longer than head and pronotum combined, mesonotum parallel-sided except for being very gently widened posteriorly. Metanotum considerably shorter than mesonotum and ony half the length of median segment. Meso- and metasternum with a fine longitudinal median carina. Tegmina elongate, spatulate, strongly narrowed in basal half and with a small, conical central hump. Alae reaching ± half way along abdominal segment III. Abdomen: Median segment 5x longer than wide and narrowed towards the posterior. Abdominal segments II–V slightly increasing in length, II less than 4x, V 5x longer than wide. VI as long as IV, VII about ¾ the length of VI; all gently constricted medially. Posterior margin of tergite V slightly transversely raised and forming a faint, rounded tubercle medially. Tergite VIII about 2x longer than wide, strongly swollen and gradually widened towards posterior margin. IX a little shorter than VIII, slightly narrowing towards the posterior and constricted medially. Anal segment as long as VII, strongly keeled, laterally compressed and tectiform. Semi-tergites long, slender, straight and with a rounded apex (Fig. 245). Interior surface apically set with numerous slightly back-curving spines. Dorsal margin strongly raised and slightly angular in anterior portion. Cerci elongate, slender, cylindrical in cross-section and in-curving, apex constricted. Poculum strongly convex, cup-like, reaching about 2/3 the way along tergite IX and with a prominent, conical central spine. Legs: All very long and slender, profemora longer and mesofemora almost as long as head, pro- and



185

mesothorax combined, metafemora projecting over posterior margin of abdominal segment V and metatibiae considerably exceeding tip of abdomen. Anterodorsal carina of profemora with 12–20 very prominent and sharp, black triangular serrations; posteroventral carina with a similar number of small teeth (Fig. 344). Mid and hind legs with all carinae distinctly dentate, although much more sparsely on dorsal carinae. Medioventral carina of femora indistinct and with a few minute spines in median section of femur. Meso- and metatibiae occasionally with a slightly enlarged median tooth on posterodorsal carina. Probasitarsus very elongate and slender, 2.5x longer than remaining tarsomeres combined, dorsal carina gently raised, all carinae unarmed. Meso- and metabasitarsi 2x the length of remaining tarsomeres combined, ventral carinae minutely serrate, dorsal carina unarmed except for a few minute teeth at the apex. Eggs (Figs. 169–170, 354): Capsule basically lens-shaped, laterally compressed, longer than high. General colouration of capsule and operculum creamish pale brown, keel, capitulum and micropylar plate a darker brown. Capsule surface almost smooth and slightly shiny. Whole capsule surrounded by an acute dorsoventral keel, beginning and ending at the operculum and only interrupted near the polar-area and the micropylar plate. Polar impression widely rounded and slightly displaced towards the micropylar plate. Micropylar plate slightly bilobed and roughly kidney-shaped, with the anterior margin decidedly impressed medially. Outer margin with a dark brown line. Micropylar cup small, rounded and placed in an oval impression at polar end of plate. Median line very short. Operculum flat, oval and in its centre with a roughly cone-shaped capitulum on a short stalk. Measurements [mm]: length (including capitulum) 5.1–5.7, length 4.2–4.9, width 2.9–3.3, height 3.8–4.2, length of micropylar plate 1.0–1.4, width of micropylar plate 2.3–2.5. Comments: Gray (1835: 42) originally described Cladoxerus serratipes based on a single % from “Malabar” in BMNH. Bragg (2001: 388) correctly stated the type locality, Malabar in S.W. India, given by Gray (1835: 42) to be rather suspect as there were no subsequent records from this region. The % HT of Ph. serratipes (Gray) in BMNH was labelled “Mal” actually referring to Malacca (Peninsular Malaysia) and perhaps misinterpreted as “Malabar” by Gray himself. Burmeister (1838: 565) described the corresponding & as Bacteria acanthopus based on a specimen from Singapore in MNHU, which was synonymised with C. serratipes Gray by Kirby (1904a: 359). Westwood (1859) and Stål (1875) had not identified these two species were identical and listed both as valid. Nor did Redtenbacher (1908: 454) who illustrated the supposedly unknown % of Pharnacia acanthopus (Burmeister, 1838), but stated this species was likely to be a synonym of Ph. serratipes (Gray). The 330 mm & that Redtenbacher erroneously attributed to Pharnacia serratipes (Gray) was the misidentified Bornean specimen of Kirby (1896: 450) in BMNH, which had already been described as Phobaeticus kirbyi by Brunner v. Wattenwyl (1907: 185). In Kirby (1904a: 359) decided this tentatively identified Bornean & to be Pharnacia maxima (Bates, 1865) a species described from Sumatra. Examination of the HT of Phibalosoma maximum Bates, 1865 in OXUM has shown this to be a fairly typical, brown & of Phobaeticus serratipes (Gray) and hence a junior synonym (syn. nov.). Redtenbacher (1908: 450) listed Pharnacia maxima (Bates, 1865) as a valid species and described the supposed % based on a specimen from Tonkin (= N-Vietnam) in NHMW, which however is a typical % Tirachoidea jianfenglingensis (Bi, 1994). Examination of the & HT of Bactridium grande Rehn, 1920 in ANSP has proven this to be a synonym of Ph. serratipes (Gray, 1835) and the type locality “Sta. Catharina, Brazil” to be obviously wrong (syn. nov.). Wood-Mason (1877a: 161) recorded Phibalosoma acanthopus (Burmeister) from N-Myanmar (Upper Tenasserim, from Moolai to Moolat, 4000–6000 feet) based on a misidentified & which most certainly represented Phryganistria grandis Rehn, 1906. Redtenbacher (1908: 453) apparently misinterpreted WoodMason’s & which made it become the PLT of Pharnacia ingens Redtenbacher, 1908, a clearly distinct not even congeneric species from the Malabar Coast of SE-India. A typical % and & from Sumatra (Indragiri, Somgai Lulah) and a & collected by Kinstler in Perak in ZMUH bear the unpublished and therefore unavailable manuscript name “Tirachoidea ajax Redt.”.


HENNEMANN & CONLE

Seow-Choen (1995: 12) described a & from Tasek Chini, Peninsular Malaysia as being the longest insect in the world. The author recorded a body length of 278 mm and an overall length of 555 mm. This is only exceeded by Ph. chani Bragg spec. nov. measuring a maximum body length of 357 mm and 567 mm in overall length. In view of the overall length Ph. serratipes can thus be regarded the second longest extant insect in the world. Brock (1999: 133) listed numerous records of Ph. serratipes in Peninsular Malaysia, stating it to be quite common in parts of Kedah (Gunung Jerai), Pahang (Genting Highlands, Pulau Tioman & Tasek Chini) and Perak (Kuala Kangsar, Tapah Hills & Ulu Piah). This species is widely distributed in Peninsular Malaysia and Sumatra and found at both low and high altitudes. A 3rd instar nymph was observed in Tanah Rata (Cameron Highlands) at 1500 m by the authors during July 1996. The record “Java” goes back to de Haan (1842: 131) and concerned to misidentified specimens of Pharnacia biceps (Redtenbacher, 1908). Hence, and since there are no subsequent records from Java, it can be regarded wrong. Redtenbacher (1908: 454) listed material of Pharnacia acanthopus (Burmeister) from Borneo to be in the collection of NHMW, which has so far remained the only record of Ph. serratipes from Borneo. However, detailed research of the NHMW-collection has not traced any Bornean specimens of Ph. serratipes, why this records as well can be regarded erroneous. TABLE 27. Measurements [mm] of Phobaeticus serratipes (Gray, 1835).

&&

&, HT of acanthopus (MNHU)

%, HT (BMNH)

%%

Body

228.0–274.5 (278.0*)

262.0

174.5

133.5–176.5

Head

9.0–10.7

9.9

–

5.2–5.8

Pronotum

6.5–8.5

8.0

5.0

4.2–5.5

Mesonotum

49.0–60.0

58.5

40.0

32.5–42.5

Metanotum

17.0–20.0

23.0

8.7

6.8–8.2

Median segment

13.7–16.0

20.0

18.6

13.5–17.0

Tegmina

-

-

10.7

9.7–11.3

Alae

-

-

38.5

36.5–42.5

Profemora

63.5–83.0

77.2

65.5

49.0–61.0

Mesofemora

49.5–64.5

59.9

48.5

40.5–49.0

Metafemora

60.0–71.5

68.1

60.5

45.0–60.0

Protibiae

81.0–101.0

109.1

87.0

63.0–88.0

Mesotibiae

51.0–59.5

64.9

51.2

38.5–54.0

Metatibiae

65.5–78.0

78.5

70.3

49.0–72.0

Antennae

85.0–94.0

> 60.0

> 96.0

> 83.0–102.0

* according to Seow-Choen (1995: 12) based on a specimen from Peninsular Malaysia (Tasek Chini).

Seow-Choen (2000: 35) listed the following foodplants to be accepted in captivity in Singapore and Malaysia: Macaranga conifera, Macaranga trilobata (Euphorbiaceae), Mangifera indica (Anacardiaceae), Psidium guajava (Myrtaceae), Rubus fruticosus and Rubus moluccanus (Rosaceae). Ph. serratipes (Gray) is successfully cultured in Europe since the early 1980’s and was included on the Phasmid Study Group culturelist as culture No. 25. It is easily reared in large cages and readily accepts bramble (Rubus spp., Rosaceae), oak (Quercus spp., Fagaceae) and various other Rosaceae and Fagaceae as alternative foodplants. Numerous papers have since dealt with the captive breeding of Ph. serratipes and have provided illustrations of both sexes, the nymphs and eggs, e.g. Carlberg (1988), Hennemann (1993) and Herbert (1988). Descriptions and / REVISION OF PHARNACIINI


187

or illustrations of the adults were also presented by Bragg (2001), Brock (1999) and Seow-Choen (2000). A detailed REM-study of the egg was provided by Lipinski et al. (1999: 80, figs. 36a–h). Distribution (Fig. 392): Peninsular Malaysia (Pulau Penang; Malacca; Perak (Kuala Kangsar; Ulu Piah [Brock, 1999: 134]; Camp Jor; Sungei-Sipoet; Gopeng; Kuala Lumpur: Taikong; Tapah Hills; BatangPadang: Kuala Who & Cameron Highlands: Tanah Rata); Pahang (Tasek Chini [Brock, 1999: 134]; Pulau Tioman [Brock, 1999: 134] & Genting Highlands [Brock, 1999: 134]; Selangor (Kepong; Kedah: Kedah Peak 3300 ft. & Gunung Jerai [Brock, 1999: 134])), Sumatra (Palambay; Tamjang; Indragiri: Somgai-Lulah; Banda Kuala; Medan: Siboelangit, Batak Plateau & Padang District) and Singapore. Java & Northern Myanmar (Upper Tenasserim: Moolat to Moolat) [in error]. Number of specimens examined: 88

Phobaeticus sinetyi Brunner v. Wattenwyl, 1907 (Figs. 106–107, 179–180, 249–251, 340–342, 393) Phobaeticus sinetyi Brunner v. Wattenwyl , 1907: 184. LT (by present designation), %: Coll. Br. v. W. Shembaganor, Prov. Madras, Pantel; det. Br. v. W. Phobaeticus sinetyi; 24.762; PLT, & + 1 egg (ex ovipositor): Coll. Br. v. W., Shembagonor, Prov. Madras, Pantel; det. Br. v. W. Phobaeticus sinetyi (NHMW, No. 308); PLT, 1 &: Kodikanel, P. Décoly; Type (MNHN, coll. Pantel); PLT, %: Kodikanel, P. Décoly (MNHN, coll. Pantel). Cappe de Baillon et al., 1937. Brock, 1996: 30. Brock, 1998a: 58. Brock & Delfosse, 2005: 55. Otte & Brock 2005: 270. ? Phibalosoma annamallayanum Wood-Mason, 1877a: 161. ST: &: Southern India, Annamallay forests, leg. Colonel R.C. Beddome (in alcohol – not traced in NZSI, presumed lost); ST, &: Southern India, Travancore Hills, leg. F.Day (valde defectum – not traced in NZSI, presumed lost). [Possible synonymy with Ph. sinetyi Brunner v, Wattenwyl, 1907; see above] Further material: [14 &&, 23 %%, eggs]: S-INDIA: 1 %: Shembagonor, Madura, S. India (F. Anglade); Phryganistria sinetyi Brunn, Det. R & Rehn 1934; Hebard Collection (ANSP); 1 %: Shembaganur, Madura, S. India, F. Anglade; Hebard Collection (ANSP); 1 %: ex Zucht: K. Rabaey (Belgien), 2005, Indien (coll. OC); 1 %: ex Zucht F. Hennemann, urspr.: SO-Indien, 2005 (coll. FH 0584-1); 1 %, eggs: ex Zucht. F. Hennemann, 2005, urspr.: SO-Indien (coll. FH, No. 0584-1 & E); 1 %: Shembagonor, Madras, Pantel (MNHN); 1 %: S-India, Animalai Hills, Sinchona, 3500 ft., May-1965, P.S. Nathan, F3224.128 (MUME). INDIA: 1 %: Phryganistria sarmentosa India Pantel (MNCN). INDONESIA [in error]: 3 %%: Indonesien, 1963 (RMNH). NO DATA: 1 &: no data (MNHN, coll. Pantel – pinned next to the type specimens); 1 %: Phobaeticus sinetyi Br., ab ove, Geinert I.1918 (MNHN, coll. Pantel); 6 %%, 3 &&: Phobaeticus sinetyi, délevage III.1916 (MNHN, coll. Pantel); 6 eggs: Phobaeticus sinetyi, 1911 (MNHN, coll. Pantel); 5 &&, 1 %: no data [apparently captive reared] (MNHN); 6 &&, 4 %%, eggs: délevage (MNHN).

Diagnosis: Well characterised by the short median segment of both sexes which is merely half as long as the metanotum, 4–7 prominent and very long spines on the posteroventral carina of the profemora of && and colouration of %%. Related to the Sri Lankan Ph. hypharpax (Westwood, 1859) but in addition to the features mentioned differing by: the broader body of both sexes; smaller lateral lobes of abdominal tergite VII; spinose medioventral carina of the meso- and metafemora and shape of anal segment of &&; the lack of tegmina and alae; broader


HENNEMANN & CONLE

semi-tergites of the anal segment and smooth (at best with 1–3 sub-basal teeth) posteroventral carina of the profemora of %%. The eggs differ from those of Ph. hypharpax by the less distinctly sculptured egg capsule and differently shaped, not conspicuously bi-lobed micropylar plate. Etymology: Dedicated to Father J. Robert de Sinéty a well-known French naturalist in India. Description: && (Fig. 106): Moderately long (body length 175.0–220.0 mm) and robust for the genus (body width ± 10.0 mm) with a short subgenital plate. General colouration of body and legs plain yellowish or greyish mid to dark brown, or pale to dark green. Leg armature reddish brown. Eyes reddish mid to dark brown. Antennae mid to dark brown and becoming blackish towards the apex, the two basal antennomeres of paler colour. Head: Oval, almost 1.5x longer than wide and slightly narrowing from eyes towards posterior; vertex very gently convex. Posterior of vertex with a slightly impressed lateral line. Between the bases of the antennae with two shallow impressions. Eyes distinct, circular and strongly convex; their length contained well 2.5x in that of cheeks. Antennae consisting of about 30 segments and roughly reaching posterior of mesothorax. Scapus dorsoventrally compressed, almost 3x longer than wide and parallel-sided. Pedicellus less than half the length of scapus, round in cross-section. Third antennomere distinctly longer than pedicellus and constricted towards apex. Following antennomeres increasing in length, but the apical four antennomeres distinctly shorter than previous. Thorax: Pronotum slightly shorter and narrower than head, 1.3x longer than wide, anterior half distinctly narrowed. Median transverse depression distinct and almost reaching lateral margins of segment, very gently curved. Mesothorax slightly constricted at anterior and slightly broadened at posterior margin, about 2.2x longer than head and pronotum combined. Mesonotum parallel-sided. Mesosternum with a very faint longitudinal median carina. Metanotum 3x longer than wide, parallel-sided and about half the length of mesonotum. Metasternum simple. Abdomen: Median segment slightly less than half the length of metanotum, about 1.2x longer than wide and widened in posterior half. Segments II–VI increasing in length, II 1.5x, VI 2.5x longer than wide. VII about ¾ the length of VI and as long as III. Tergite VII with lateral margins forming a rather small, rounded lobe posteriorly. Praeopercular organ formed by two flat, acute lobes at posterior margin of sternum VII. Tergite VIII distinctly narrower than previous, about equal in length to VII, almost 3x longer than wide, parallel sided and strongly convex. IX parallel-sided, strongly convex, half the length of VIII and indistinctly longer than wide. Anal segment narrow, slightly longer than IX with a fine median carina and a distinct triangular posteromedian excavation; posterolateral angles roundly triangular (Fig. 250). Supraanal plate very indistinct, rounded and with a median keel. Cerci small, oval in cross- section, bristled and tapered towards a narrow apex. Subgenital plate strongly longitudinally keeled and reaching to or slightly projecting over posterior margin of anal segment (Fig. 249). Legs: All slender and of moderate length, profemora almost as long as head, pro- and mesothorax combined, mesofemora about as long as mesonotum, metafemora projecting over posterior margin of abdominal segment IV and metatarsi slightly exceeding apex of abdomen. Posteroventral carina of profemora armed with 4–7 very prominent and long spines; anterodorsal carina with ± 12 distinct and acute serrations (Fig. 340). Dorsal and posteroventral carinae of protibiae with minute serrations. Meso- and metafemora with all carinae distinctly dentate, although more sparsely and slightly less distinct on dorsal carinae. Medioventral carina well developed and armed with 5–6 distinct spines (Fig. 342). Meso- and metatibiae with all carinae more or less distinctly serrate. Mesotibiae occasionally with a more or less prominently enlarged triangular tooth or lobe one third the way along posterodorsal carina. Probasitarsus as long as remaining segments combined, dorsal carina gently elevated, all carinae unarmed. Meso- and metabasitarsi as long as following three segments combined, dorsal carina gradually elevated towards the apex; all carinae minutely dentate. %% (Fig. 107): Small to medium-sized (body length 112.0–142.5 mm) and rather robust for the genus (maximum body width ± 3.0 mm), apterous. Body surface glabrous. General colouration of body and legs,



189

ferruginous, ochracheous or mid brown. Greater portion of the mesonotum except for the posterior portion and most of metanotum covered by a washed malachite or bluish green marking; may be faint in some specimens. Meso- and metapleurae and sterna pale greyish green. Meso- and metapleurae occasionally with a pale posterior marking. Head with a more or less prominent washed dark brown postocular stripe; the vertex usually of paler colour than rest of head capsule or sometimes pale cream between the eyes. Eyes dark reddish brown with darker brown mottling. Pronotum often with two faint dark longitudinal markings along lateral margins. Antennae dark brown dorsally and black ventrally, two basal antennomeres pale to mid brown. Leg armature distinctly black. Head: Generally as in &&, but more prominently narrowed from eyes towards posterior. Eyes very prominent, circular and projecting hemispherically from head capsule; their length contained about 2x in that of cheeks. Antennae as in && but projecting over posterior margin of metanotum. Thorax: Pronotum generally as in &&. Mesothorax 3.1x longer than combined length of head and pronotum, mesonotum parallel-sided, but slightly widened at posterior margin. Metanotum 5.5x longer than wide, parallel-sided and slightly more than half the length of mesonotum. Meso- and metasternum with very faint longitudinal median carina, although very indistinct on metasternum. Abdomen: Median segment slightly less than half the length of metanotum, 3x longer than wide and slightly widened posteriorly. Segments II to VI slightly increasing in length, II almost 4x and VI 5x longer than wide. VII shorter than previous, about as long as II. Tergite VIII ¾ the length of VII, strongly swollen and widened towards the posterior. IX distinctly narrower and about 2/3 the length of VIII, 1.8x longer than wide, strongly convex, constricted medially and with anterior margin broader than posterior margin. Anal segment strongly keeled, laterally compressed and tectiform. Semi-tergites moderately slender, slightly downcurving and apically rounded (Fig. 251). Interior surface set with several slightly back-curving, dark reddish brown spines. Cerci elongate, round in cross-section, slightly in-curving and tapered towards the apex. Poculum strongly convex, cup-like with the centre conically raised, reaching about 2/3 the way along tergite IX. Legs: All long and slender, profemora longer than head, pro- and mesothorax combined, mesofemora slightly longer than mesonotum, metafemora ± reaching posterior margin of abdominal segment V and metatibiae exceeding distinctly over apex of abdomen. Anterodorsal carina of profemora armed with 9–20 prominent, acutely triangular serrations; posteroventral carina smooth, or at best with 1–3 small teeth (Fig. 341). All carinae of protibiae unarmed. Meso- and metafemora with all carinae distinctly dentate, dorsal carinae with a smaller number of teeth or entirely unarmed. Medioventral carina indistinct and unarmed, at best with 1–3 minute spines. Meso- and metatibiae with the ventral carinae minutely but regularly dentate, and the dorsal carinae sparsely dentate or unarmed. Probasitarsus longer than remaining tarsomeres combined, slender and unarmed. Meso- and metabasitarsi about as long as remaining tarsomeres combined with all carinae minutely dentate. Eggs (Fig. 179–180): Capsule basically lens-shaped, laterally compressed, 1.5x longer than high. General colouration of capsule and operculum plain pale to dark brown. Capsule surface coriaceous and slightly wrinkled. Capsule surrounded by dorsoventral keel, beginning and ending at the operculum and only interrupted at the polar-area and micropylar plate. Seen laterally the posterior section of the capsule is distinctly angular. Dorsal surface with two pairs of distinct, deep impressions on each side of the dorsoventral keel, one anteriorly near the operculum and a second smaller one just below micropylar plate. Micropylar plate shieldshaped, about as long as wide with a prominent posteromedian indention and a gently angulate anterior margin. Median line distinct. Micropylar cup small, at polar end of plate and near centre of dorsal egg surface. Operculum flat, oval and in the centre with a conical, dark brown capitulum on a short stalk. Measurements [mm]: length (including capitulum) 5.0–5.3, length 4.4–4.9, width 2.8–3.0, height 3.4–3.7, length of micropylar plate 1.2–1.4. Comments: Brunner v. Wattenwyl (1907: 184) did not refer to any material in NHMW in the original description of Phobaeticus sinetyi. However, examination of the % and & in NHMW, which bear labels stating


HENNEMANN & CONLE

„Pantel“, and comparison with the % and & syntypes in MNHN (coll. J. Pantel) prove they have been retained in NHMW after publication of the monograph (1906–1908) by Brunner v. Wattenwyl. Furthermore, Brunner v. Wattenwyl (1907: 184) stated the type locality to be “Shembaganor in Pulney Hills, Prov. Madras”, which matches exactly with the data of the % in NHMW. Thus, and as the measurements given by Brunner v. Wattenwyl roughly correspond to the NHMW specimens, they are clearly part of the type series. The % in NHMW with the collection-number 24.762 is here selected as the LT of Ph. sinetyi. The PLT & in MNHN is conspicuously larger than the & in NHMW (→ Table 27) and darker brown, which was however most probably caused by preservation. A further & without data is pinned next to the two PLT’s does not appear to be part of the type series. The collection of J. Pantel and main collection of MNHN both contain several eggs and captive reared specimens from a culture maintained in France between 1911 and 1918. The colouration of these && varies from pale brown to bright green. A % in MNCN has the leg armature very strongly developed. Due to all other known records of Ph. sinetyi are from South India, the locality “Indonesia” recorded for the three %% in RMNH can undoubtedly be regarded as erroneous. Phibalosoma annamallayanum Wood-Mason, 1877a was described based on two && from South India, one from Annamallay Forest and one from the Travancore Hills. This species is here transferred from Pharnacia Stål, 1875 to Phobaeticus Brunner v. Wattenwyl, 1907 (comb. nov.) due to having a distinct praeopercular organ. A % from the Annamallay Hills in MUME is a typical specimen of Ph. sinetyi and along with the very brief diagnosis and measurements of the & provided by Wood-Mason (1877a: 161, 1877b: 346), which widely match with Ph. sinetyi, suggests these two species might be identical. However, any decision on a possible synonymy of Ph. sinetyi with Ph. annamallayanus requires availability of newly collected %% and && from the Annamallay Hills, the type locality of the senior species. If this is confirmed, Ph. sinetyi Brunner v. Wattenwyl will fall as a synonym of Ph. annamallayanus (Wood-Mason). TABLE 28. Measurements [mm] of Phobaeticus sinetyi Brunner v. Wattenwyl, 1907.

%, LT Body

%, PLT (MNHN)

%

&, PLT

&, PLT

(NHMW)

(coll. FH)

(NHMW)

(MNHN)

142.5

140.0

130.8

176.0

220.0

Head

6.9

6.7

5.9

10.1

12.5

Pronotum

5.0

4.3

4.1

7.5

9.7

Mesonotum

30.0

31.8

27.7

36.8

43.5

Metanotum

18.3

17.5

15.2

18.5

24.5

Median segment

8.2

8.0

7.8

9.0

10.6

Profemora

46.0

49.4

39.1

50.0

56.2

Mesofemora

35.2

37.7

33.7

37.0

40.7

Metafemora

42.0

44.9

41.0

42.0

48.5

Protibiae

57.0

-

59.2

57.3

-

Mesotibiae

36.2

-

37.3

38.7

-

Metatibiae

46.0

-

47.8

46.1

-

Antennae

> 49.5

> 42.0

50.0

-

> 42.0

The two %% in ANSP have body lengths of 129.5 mm and 141.5 mm. Captive reared specimens in MNHN measure: %% 112.0–126.0 mm, && 175.0–197.0 mm.

The rather robust body of && resembles the more slender taxa of Pharnacia Stål, 1877 whereas the short median segment and colouration of the %% resemble certain species of Phryganistria Stål, 1875. The typical laterally compressed eggs with their small, shield-shaped micropylar plate, structures of the genitalia and dis-



191

tinct black dorsal serrations of the profemora however clearly confirm the correct placement of Ph. sinetyi in its original genus Phobaeticus Brunner v. Wattenwyl, 1907. Distribution (Fig. 393): Southern India (Annamallay Hills; Madura & Madras Province: Shembagonor & Kodikanel). Number of specimens examined: 36

Phobaeticus sobrinus Brunner v. Wattenwyl , 1907 (Figs. 121, 264, 336, 394) Phobaeticus sobrinus Brunner v. Wattenwyl , 1907: 184, pl. 7: 1a–b (%). LT, %: Sumatra, Si-Rambé, XII. 1890-III.1891, E. Modigliani; Type; Phobaeticus sobrinus Br. (MCSN). [Not: PLT, &: Nias Island, 1897–1898, leg. U. Raap (MCSN). This is clearly a distinct species and not a member of Phobaeticus; see comments below] Brock, 1996: 30. [Designation of LT] Otte & Brock, 2005: 270. Phobaeticus beccarianus Brunner v. Wattenwyl , 1907: 185. HT, & (penultimate instar nymph): Sumatra, Mt. Singalang, Luglio, leg. Beccari VII. 1878; Type, Beccarianus Brunner v. W.; Phobaeticus beccarianus Br. (MCSN) syn. nov. Brock, 1996: 28. Otte & Brock, 2005: 268. Pharnacia beccarianus, Günther, 1935b: 139.

Diagnosis: Related to Ph. serratipes (Gray, 1835) but differing by: the smaller size; shorter median segment and pale orange bases of the profemora of both sexes; less prominent posteromedian excavation of the anal segment and prominent dorsal and ventral lobes of the mesofemora and meso- and metatibiae of &&, as well as the lack of tegmina and alae and dorsally rounded anterodorsal crest of the anal segment of %%. && are only known from a unique penultimate instar nymph but appear to differ from the also related and Sumatran Ph. mucrospinosus spec. nov. by: the smaller size (estimated ± 200.0 mm when adult); slightly more robust body and different armature of the mid and hind legs. Etymology: “Sobrinus” (lat. = inconspicuous). Description: && (penultimate instar nymph): Moderately long (body length 163.0 mm; estimated to be ± 200.0 mm when adult) and slender species (maximum body width ± 4.0 mm) with characteristic leg armature. General colouration ochracheous mid brown, with slightly darker areas on abdomen and very faint darker transverse bands on the mid and hind legs. Head with a slightly paler brown, coronal-line and a faint, broad postocular line. Bases of all femora pale orange, those of the profemora in particular. Eyes dark ochracheous. Antennae brown and becoming paler towards the base. Head: Oval, almost 1.5x longer than wide, oval in cross section and slightly narrowing towards the posterior. Eyes small, circular and slightly projecting from head capsule; length contained less than 3x in that of cheeks. Antennae at least reaching half way along mesonotum (broken in the HT). Scapus dorsoventrally compressed, about 2.5x longer than wide and slightly constricted towards the base. Pedicellus distinctly shorter than scapus, round in cross-section, sub-spherical. Thorax: Pronotum slightly narrower than head but of almost equal length, 1.5x longer than wide and narrowed in anterior portion. Transverse median depression very gently curved and almost reaching lateral margins of segment. Mesothorax almost 4x longer than head and pronotum combined and 2.5x longer than metanotum. Mesonotum parallel-sided and with a very faint longitudinal median line. Metanotum about 4.5x longer than wide, parallel-sided. Pleurae and sterna simple. Abdomen: Median segment slightly more than 2/3 the length of metanotum, gently constricted medially. Segments II–VI cylindrical and increasing in length, II 2x, III and IV 3.5x, V 4x and VI slightly more than 4x longer than wide. II gently narrowing towards the posterior, III–VII parallel-sided. VII slightly narrower than previous and about ¾ the length of VI. Praeopercular organ, formed by two small, laterally compressed teeth


HENNEMANN & CONLE

at posterior margin of sternum VII. Tergite VIII shorter than VII, longer than IX and X combined, strongly convex and parallel-sided. IX less than half the length of VIII, indistinctly longer than wide and trapezoidal, gradually widening towards posterior margin. Anal segment ¾ the length of IX, with a fine median carina and a small triangular notch posteromedially. Supraanal plate very small, rounded and keeled. Cerci very small, cylindrical and tapered towards a pointed tip. Subgenital plate keeled and scoop-shaped (not fully developed in the HT). Legs: All very long and slender, mid and hind legs gently down-curving. Profemora almost as long as head, pro- and mesothorax combined, mesofemora about as long as mesothorax, metafemora slightly projecting over posterior margin of abdominal segment IV and metatarsi exceeding apex of abdomen. Anterodorsal carina of profemora densely and prominently serrate, posteroventral carina densely dentate. Posterodorsal carina very faint and armed with 3–4 minute, triangular teeth.. All carinae of protibiae unarmed except for a few minute teeth on posteroventral carina and a single enlarged, triangular tooth just before middle of tibia on posterodorsal carina. Anterodorsal carina of mesofemora with three prominent, triangular lobes in great but roughly equal distance (Fig. 336). Posteroventral carina slightly dilated sub-basally and in this portion with 2– 3 prominent teeth. All other carinae dentate. Metafemora with all carinae dentate, although more minute and sparsely on dorsal carinae. Medioventral carina of meso- and metafemora indistinct and armed with a few minute spines. Posterodorsal and medioventral carinae of meso- and metatibiae each with a large, roundly triangular lobe about 1/3 off the base; the dorsal lobe considerably larger. Probasitarsus as long as remaining segments combined except claw, unarmed. Meso- and metabasitarsi as long as following three segments combined, dorsal carina slightly rounded and all carinae minutely dentate. %% (Fig. 121): The % LT of Ph. sobrinus is in poor condition with most extremities glued. It lacks the right hind leg, right mesotibia and complete right protarsus as well as the four terminal segments of the left protarsus, complete left metatarsus and great parts of the antennae. Rather small (body length 125.0 mm) and slender for the genus, apterous. General colouration of body and legs yellowish brown with slightly darker areas on abdomen. Head with a pale brown coronal line and a washed, roughly triangular pale marking between the eyes. Meso- and metasternum as well as bases of profemora pale orange-brown. Lateral margins of meso and metathorax with a faint blackish longitudinal line. Armature of legs very dark reddish brown. Eyes reddish brown with darker brown mottling. Antennae dark brown dorsally and black ventrally, two basal antennomeres pale brown. Head: Oval, almost 1.5x longer than wide, broadest at eyes and strongly narrowing towards posterior margin. Between the bases of the antennae with an oval, slightly raised and smooth area. Eyes very prominent and projecting hemispherically from head capsule; their length contained only about 1.7x in that of cheeks. Antennae longer than head and pronotum combined (broken in the LT). Scapus dorsoventrally compressed, constricted towards the base and about 3x longer than wide. Pedicellus distinctly shorter than scapus, subspherical. Thorax: Pronotum generally as in &&, almost equal in length to head but distinctly narrower, the anterior portion narrowed. Median transverse depression distinct, gently curved and reaching to lateral margin of segment. Mesothorax almost 4x longer than head and pronotum combined, mesonotum parallel-sided except for being very slightly broadened at posterior margin. Metanotum less than half the length of mesonotum 6x longer than wide, parallel-sided. Meso- and metasternum with a faint longitudinal median carina. Tegmina represented by a pair of very small, oval lobes (c. 1.0 mm). Abdomen: Median segment 2/3 the length of metanotum, almost 5x longer than wide and slightly broadening towards the posterior. Segment II 4.5x longer than wide slightly shorter than median segment. III–VI in average 5.5x longer than wide and slightly constricted medially. VII shorter than VI and slightly widening towards posterior margin. Tergites V and VI with a small tubercle posteromedially. VIII swollen and gradually broadening towards posterior margin and less than 2.5x longer than wide. IX slightly shorter than VIII, anterior margin broader than posterior margin, strongly constricted medially and convex with a fine median



193

carina. Anal segment as long as VII, strongly keeled, laterally compressed and tectiform. Semi-tergites long, straight and finger-like with a rounded apex (Fig. 264). The interior surface apically set with minute black spines. Cerci moderately long, slender, round in cross-section and incurving in apical portion. Poculum strongly convex, cup-like, not reaching about half way along tergite IX and with a prominent but blunt central spine. Legs: All very long and sender, profemora considerably longer than pro-, meso- and metathorax combined, mesofemora a little longer than pro- and mesothorax combined, metafemora reaching to abdominal segment VI and metatibiae projecting exceeding apex of abdomen. Anterodorsal carina of profemora with 19– 22 prominent, sharply triangular serrations. Posteroventral carina with a similar number of small but pointed teeth. Protibiae unarmed except for a few minute serrations on posteroventral carina. Two outer ventral carinae of meso- and metafemora sparsely armed with small, pointed teeth, dorsal carinae with only very few minute teeth. Medioventral carina of meso and metafemora indistinct and armed with a few very minute spines. Meso- and metatibiae with all carinae minutely dentate, more sparsely on dorsal carinae. Probasitarsus very long and slender, dorsal carina very slightly raised, all carinae unarmed (remaining tarsomeres missing in LT). Mesobasitarsus as long as remaining segments combined and with all carinae minutely dentate. Comments: Brunner v. Wattenwyl (1907: 184) described Phobaeticus sobrinus based on adult % and & both in MCSN. Brock (1996: 28) designated the % from Sumatra (Si-Rambé) as LT and selected Ph. sobrinus Brunner v. Wattenwyl as the type species of Phobaeticus Brunner v. Wattenwyl, 1907. Detailed examination of the & PLT from Nias Island and comparison with the % LT has clearly shown these to represent two distinct specie, which do not even belong in the same genus. The shape of the anal segment and genitalia, lack of an elevated and displaced, ledge-like medioventral carina of the profemora as well as the oval body cross-section place the & PLT in close relation to the genus Nesiophasma Günther, 1934 (tribe Stephanacridini). Clarification if its generic position including a description is in progress by the authors. The HT of Ph. beccarianus Brunner v. Wattenwyl in MCSN is a penultimate instar & nymph, which may have reached an estimated body length of ± 200.0 mm when adult. Examination and careful comparison with the % LT of Ph. sobrinus has shown Ph. beccarianus merely to represent the corresponding & (syn. nov.). Only known from the type specimens. Eggs unknown. Distribution (Fig. 394): W-Sumatra (Si-Rambé & Mount Singalang). Number of specimens examined: 2

TABLE 29. Measurements [mm] of Phobaeticus sobrinus Brunner v. Wattenwyl, 1907.

%, LT (MCSN)

& (penultimate instar nymph), HT of Ph. beccarianus (MCSN)

Body

125.0

163.0

Pronotum

3.3

5.0

Mesonotum

29.0

35.3

Metanotum

12.5

15.5

Median segment

9.8

12.5

Profemora

42.5

44.6

Mesofemora

34.0

37.0

Metafemora

42.0

38.0

Antennae

> 15.0

> 23.0


HENNEMANN & CONLE

The following species, Phobaeticus incertus Brunner v. Wattenwyl, 1907, is provisionally retained in the genus Phobaeticus Brunner v. Wattenwyl but with doubt. It differs considerably from all other members of Phobaeticus Brunner v. Wattenwyl, 1907 by lacking a praeopercular organ in &&. It furthermore differs from entire Pharnaciini by having the medioventral carina of the profemora serrate and about midways on the ventral surface of the femur. The egg as well differs considerably from all other known eggs of Phobaeticus and is rather unlike those of other Pharnaciini, having a very small, rhomboidal micropylar plate and the capsule irregularly impressed and covered with wart-like structures. Unfortunately, the internal micropylar plate is as yet unknown but the rhomboidal shape of the plate is usually associated with closed plates (Figs. 167–168). The supposedly closed internal micropylar plate and structure of the medioventral carina of the profemora would place Ph. incertus in the tribe Stephanacridini and indeed the general appearance and shape of the genitalia, such as the laterally excavated anal segment (Fig. 252), indicate relation to the genera Eucarcharus Brunner v. Wattenwyl, 1907 and Nesiophasma Günther, 1934 (→ 4.7.2). The egg shows similarity to those of e.g. Nesiophasma plateni (Dohrn, 1910) in several aspects (Fig. 60). However, without having at hand %% and && from the same locality and knowledge of the internal micropylar plate of the eggs it is for now difficult to decide about the taxonomic position of Ph. incertus with confirmation. A clarification is in progress by the authors. Despite the unconfirmed taxonomic position a complete treatment of this species is presented below.

Phobaeticus (?) incertus Brunner v. Wattenwyl, 1907 (Figs. 167–168, 252, 395) Phobaeticus incertus Brunner v. Wattenwyl, 1907: 185. HT, & (+ 4 eggs ex abdomen): 3476; Banka v. Mart; Phobaeticus incertus Br., Brunner det.; Holotypus (MNHU). Brock, 1996: 28. Otte & Brock, 2005: 268. Zompro, 2005b: 267. Nearchus grubaueri Redtenbacher, 1908: 448. HT, &: Grubauer 1902, Kwala Kangsar, Perak; Mus. Caes. Vindobon; det. Br. v.W. Pharnacia grubaueri (NHMW, Nr. 857). syn. nov. Bragg, 1995b: 275. Brock, 1995: 95. Brock, 1998a: 32. Seow-Choen, 2000: 35, pl. 92 (&). Bragg, 2001: 393. Brock, 2002: 43. Otte & Brock, 2005: 209. Further material: [2 &&]: PENINSULAR MALAYSIA: 1 &: Malakka, Perak, A. Grubauer leg.; B. Jachau vend. 15.12.1901 (ZMUH); 1 N.C.E. Miller 1928 (BMNH).

&: Malacca, Selangor, Ulu Gombak,

Etymology: “Incertus” (lat. = uncertain, unknown). Description: && (Fig. 105): Long (body length 167.0–209.0 mm; incl. subgenital plate 189.0–227.0 mm) slender species (± 7.5 mm) with a very long, lanceolate subgenital plate (± 40.0 mm). Body surface smooth and slightly glossy. General colouration of body and legs uniformly dark greyish or reddish brown, occasionally with a bold white longitudinal median line, which begins on the head, is very fine on the mesonotum, interrupted on abdominal tergites and terminates on tergite VI. The line may be very broad and form a wedgeshaped marking on the vertex. Bases of profemora pale yellowish or red interiorly. Legs with indistinct whitish annulations, armature dull red. Eyes dark reddish brown. Antennae blackish brown.



195

Head: Head elongate, suboval 1.5x longer than wide, broadest at eyes and slightly narrowing towards the posterior. Vertex flat and smooth. Between the bases of the Antennae with a small but distinct, transverse depression. Eyes large, circular and moderately convex; their length contained a little more than 2x in that of cheeks. Antennae reaching to posterior margin of mesonotum; all antennomeres bristled. Scapus dorsoventrally flattened and about 1.7x longer than wide, apex slightly club-like. Pedicellus distinctly shorter than scapus, cylindrical. Following segments first increasing until mid of antenna, then decreasing in length towards apex. Thorax: Pronotum rectangular, 1.5x longer than wide, shorter and narrower than head. Anterior margin slightly raised, median transverse depression flat and curved, almost reaching lateral margins of segment. Anterior half with an impressed median line. Mesothorax 2.4–2.5x longer than head and pronotum combined, mesonotum with a faint longitudinal median line and very slightly broadened towards posteriorly. Mesosternum with a very shallow longitudinal median keel, which is most decided in anterior portion and gradually terminates towards the posterior margin. Metanotum 3x longer than wide, less than half the length of mesonotum, very slightly constricted medially. Metasternum smooth. Abdomen: Median segment about 2/3 the length of metanotum, rectangular and 1.5x longer than wide. Segments II–VI increasing in length, II 2x, III–IV 2.5x and V–VI 3x longer than wide, lateral margins very gently convex. VII slightly narrower than previous, 3x longer than wide. Tergite VI with a small, transverse posteromedian tubercle; remaining tergites smooth. VIII distinctly narrower and about 3/5 the length of VII, 2.5x longer than wide, strongly convex and constricted medially. IX half the length of VIII, strongly convex, roughly quadrate. Anal segment longer than IX, with a fine median carina and a small, triangular excavation posteromedially; lateral margins widely excavated at bases of cerci. Supraanal plate small, strongly keeled, rounded and not projecting over posterior margin of anal segment. Cerci small, oval in cross-section and slightly tapered towards the apex. Subgenital plate longitudinally keeled, gradually tapered towards an acute apex, very long and lanceolate; projecting over apex of abdomen by more than the combined length of tergites VIII–X (Fig. 252). Legs: All long and slender with all carinae distinctly and more or less uniformly dentate, lacking any considerably enlarged spines or teeth. Profemora about as long as pro- and mesothorax combined, mesofemora a little longer than mesothorax, metafemora projecting over posterior margin of abdominal segment IV and metatarsi reaching to apex of anal segment. Anterodorsal carina of profemora with ± 20 distinct and very sharp, triangular teeth; posteroventral carina with a similar number of slightly smaller teeth. Medioventral carina of profemora distinct and dentate; about midways on ventral surface of femur. Medioventral carina of meso- and metafemora distinct and armed with a longitudinal row of ± 10 pointed spines. Medioventral carina of meso- and metatibiae strongly raised and densely dentate. Probasitarsus slightly shorter than remaining tarsomeres combined, all carinae unarmed, the dorsal carina gradually raised towards the apex. Meso- and metabasitarsi with a slightly raised and rounded dorsal carina, all carinae densely serrate, about as long as following three tarsomeres combined. All basitarsi with a minutely dentate medioventral carina. Eggs (Figs. 167–168): The following description is based on a single fully developed egg which was removed from the ovipositor of the HT of N. grubaueri Redtenbacher in NHMW. Eggs extracted from the abdomen of the HT of Phobaeticus incertus Brunner v. Wattenwyl in MNHU were only used for comparison and confirming the synonymy of these two species. The description below is based on the unique fully developed egg in NHMW. Large, general colouration of capsule dark greyish brown with irregular lighter markings. Capsule with irregularly raised and impressed areas, slightly laterally compressed and longer than high. There is a distinct wide transverse indention across the polar-area and the micropylar plate is conspicuously impressed. Whole capsule surface densely but minutely granulated and covered with numerous small, whitish to pale brown, wart-like structures. Micropylar plate small and roundly rhomboidal, its length contained about 4x in that of capsule. Micropylar cup, distinct, pale brown, cup-like and positioned well away from the posterior margin of


HENNEMANN & CONLE

the plate. Operculum orange-brown slightly oval, strongly concave and with distinct radial lamella-like ridges. Capitulum prominent, conical with deep radial impressions and on very short stalk; brown. Measurements [mm]: length (including capitulum) 4.9, length 4.0, width 3.1, height 3.9, length of micropylar plate 0.9. Comments: Brunner v. Wattenwyl (1907: 185) described Phobaeticus incertus based on a single & from Banka Island, a few kilometres off the northeast coast of Sumatra, in MNHU. Redtenbacher (1908: 448) described Nearchus grubaueri based on a single & from Kuala Kangsar, Perak (Peninsular Malaysia) in NHMW, not being aware that this was identical to Brunner v. Wattenwyl’s species. Detailed examination and comparison of the HT’s of Ph. incertus Brunner v. Wattenwyl and N. grubaueri Redtenbacher has not shown any significant differences, except from the latter being remarkably larger and having distinct white markings on the body and head. Comparison of eggs extracted from the abdomen of the HT of Ph. incertus and a single egg removed from the ovipositor of the HT of N. grubaueri have also not revealed any differences that would allow specific distinction and additionally confirm the synonymy of these two taxa. Consequently, N. grubaueri falls as a junior synonym of Ph. incertus Brunner v. Wattenwyl (syn. nov.). The taxonomic position of Ph. incertus is questionable and certain features indicate relation to Eucarcharus Brunner v. Wattenwyl, 1907 and Nesiophasma Günther, 1934, two genera of the tribe Stephanacridini (see comments above). The specific name “incertus” indicates that already Brunner v. Wattenwyl was uncertain about the generic position of his new species. Two && and eggs from the Andaman Islands were examined from photos kindly sent by P.D. Brock (Slough, England) and represent a new record for Ph. (?) incertus. Both were collected from the canopy of primary rain forest between 20–25 metres above the ground (pers. comm. with P.D. Brock). % unknown. Distribution (Fig. 395): Pulau Banka prope Sumatra (in the Straight of Malacca), Peninsular Malaysia (Malacca; Perak: Kuala Kangsar & Selangor: Ulu Gombak) & Andaman Islands. Number of specimens examined: 4

TABLE 30. Measurements [mm] of Phobaeticus (?) incertus Brunner v. Wattenwyl, 1907.

&, HT (MNHU)

&, HT of N. grubaueri (NHMW)

Body

167.0

209.0


189.0

227.0

Pronotum

6.3

8.0

Mesonotum

33.0

40.5

Metanotum

15.2

18.0

Median segment

9.0

11.0

Profemora

40.5

47.5

Mesofemora

35.0

43.5

Metafemora

43.3

51.0

Protibiae

43.0

53.3

Mesotibiae

31.0

39.0

Metatibiae

42.0

52.3

Antennae

> 36.0

53.0



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6.4 Genus Phryganistria Stål, 1875

FIGURES 126-129: Phryganistria spp. 126. Phryganistria grandis Rehn, 1906, & N-Vietnam, Than-Moi (ZMUH) 127. Phryganistria heusii (Hennemann & Conle, 1997), & N-Vietnam, captive reared (coll. FH, No. 0240-4) 128. Phryganistria heusii (Hennemann & Conle, 1997), % N-Vietnam, captive reared (coll. FH, No. 0240-11) 129. Phryganistria virgea (Westwood, 1848), & N-India, Darjeeling (ZMUH)


HENNEMANN & CONLE

Type species: Bacteria sarmentosa Westwood, 1848: 66, pl. 32: 1–2 (= Bacteria virgea Westwood, 1848: 66), by subsequent designation by Kirby, 1904a: 358. Phryganistria Stål, 1875: 6 & 63 (in part). Kirby, 1904a: 358. Rehn, 1906: 279. Brunner v. Wattenwyl , 1907: 181–183. Otte & Brock, 2005: 272 (in part). Chen & He, 2007: 337, figs. 308-309. Bacteria, Westwood, 1848: 66, pl. 32: 1–2. Lonchodes, Westwood, 1859: 46 (in part). Phobaeticus Brunner v. Wattenwyl , 1907: 183 (in part). Günther, 1934a: 94, fig. 6. Günther, 1935b: 124. Brock, 1996: 28 (in part). Otte & Brock, 2005: 272 (in part).

Description (%, &): Long to very long, mostly very elongate and moderately slender Pharnaciini, both sexes apterous. Sexual dimorphism moderate. %% either much smaller than && or body only indistinctly shorter. Body surface (especially of %%) shiny. Head distinctly longer than wide, vertex flat or very slightly convex and smooth. Antennae of %% indistinctly longer than those of &&, as long as combined length of head, proand mesonotum or slightly longer than complete thorax and head combined. Antennomeres very elongate, usually more than 5x longer than wide. Mesosternum of %% simple, not keeled. Median segment very short, at best 2/5 the length of metanotum. Abdominal tergites occasionally with a posteromedian tubercle and / or lateral margins of II–VII gently rounded; tergite VII at best with small, rounded lobes posterolaterally. Sternum VII of && with a distinct praeopercular organ, formed by two elongate, tooth-like lobes or blunt spines. Semitergites of anal segment of %% raised anterodorsally, of a roughly triangular overall-shape, apex ± elongated. Subgenital plate of %% strongly convex, conical and with a blunt hump at the angle. Subgenital plate of && either slightly projecting over anal segment or conspicuously lanceolate and projecting over anal segment by more than the combined length of tergites VIII–X. Cerci simple, conical and cylindrical or oval in cross-section. All legs very long and ± prominently dentate or serrate; occasionally single slightly enlarged teeth are present on mid-legs. Anterodorsal carina of profemora ± serrate. Meso- and metafemora of %% either slender, or distinctly broadened and with a prominent, dagger-like sub-apical spine on the antero- and posteroventral carinae; medioventral carina indistinct. Mesofemora distinctly longer than metanotum and median segment combined. Probasitarsus longer than remaining tarsomeres combined. Meso- and metabasitarsus at least as long as the following three tarsomeres combined; carinae not conspicuously elevated, except for dorsal carina which may be slightly rounded apically. Eggs (Figs. 186–191): Medium-sized to large. Capsule distinctly longer than high, oval to slightly elliptical in cross-section and with a slight keel on dorsal and ventral surfaces and polar-area. Capsule surface minutely punctate, slightly shiny. External micropylar plate at least 2/3 the length of capsule, elongate, tapering towards anterior end and with a conspicuous notch posteromedially. Operculum flat, oval. Capitulum hatlike and distinctly stalked. Diagnosis (Table 2): Closely related to Phobaeticus Brunner v. Wattenwyl , 1907 and Baculonistria gen. nov.. With the first it shares the prominent praeopercular organ of &&, but it is easily distinguished by: the shorter median segment, which is at best 2/5 the length of the metanotum and and the lack of a median keel on the mesosternum of %%. The eggs differ from those of Phobaeticus by the longer, lanceolate micropylar plate, which covers at least 2/3 of the capsule length, and the less laterally compressed, not lens-shaped capsule. From Baculonistria gen. nov. it differs by: the relatively longer mesonotum, which is distinctly longer than the metanotum; longer median segment (only about 1/8 the length of metanotum in Baculonistria); longer mesofemora, which clearly exceed the metanotum and median segment in length; lack of a sub-apical



199

spine on the medioventral carina of the meso and metafemora and much longer antennae of both sexes. && furthermore differ by: the lack of spines between the eyes; elongate antennomeres and prominent praeopercular organ, which is formed by two spines or pointed lobes at posterior margin of sternum VII of && (a single, transverse lobe in Baculonistria). The eggs differ from those of Baculonistria gen. nov. by the laterally compressed, almost smooth egg-capsule and relatively longer, more slender micropylar plate. Variation: As in the closely related Phobaeticus several features of the insects are subject to strong intrageneric variability. The length of the subgenital plate is considerably variable: it is either short with the tip just reaching the posterior margin of the anal segment or strongly elongated, lanceolate and projects strongly over the apex of the abdomen. The meso- and metafemora of %% are either slender and unarmed subapically, or conspicuously thickened, distinctly broader than the corresponding tibiae and sub-apically armed with a long, dagger-like spine on the two outer ventral carinae. Species, in which %% have strongly thickened meso- and metafemora and a distinct sub-apical spine on the ventral carinae, tend to show less striking sizedimorphism with both sexes being almost equal in body length. In contrast, %% of those species which have the meso- and metafemora not distinctly thickened and lack a prominent sub-apical spine on the ventral carinae, are considerably shorter than the corresponding &&. In all other genera of Pharnaciini the sexual dimorphism is much more striking with %% being conspicuously smaller than &&, reaching at best 65% of their body length. Comments: Stål (1875: 6 & 63) established Phryganistria for Bacteria sarmentosa Westwood, 1848, Bacteria virgea Westwood, 1848 and Phasma (Cladoxerus) acanthopus, de Haan, 1842 (not Burmeister, 1838, → see Tirachoidea biceps Redtenbacher, 1908). Stål placed Phryganistria in close relation to genera currently belonging in the subfamily Lonchodinae, and apart from exhibiting distinctly armed and dentate legs, characterized the genus by the short median segment: “..segmento mediano longiore quam latiore…. (1875: 6)”. Due to the first two species are synonymous and the latter belongs in the genus Tirachoidea Brunner v. Wattenwyl, 1893, Kirby (1904a: 358) designated Bacteria sarmentosa Westwood as the type species of Phryganistria, unaware this had been synonymised with Bacteria virgea Westwood, 1848 by Westwood (1859: 46). Brunner v. Wattenwyl (1907: 181) contained Phryganistria in his tribe Clitumnini and distinguished it from Phobaeticus Brunner v. Wattenwyl, 1907 by the smooth medioventral carina of the meso- and metafemora, which is however a variable feature. It is true for Ph. virgea, the only species that Brunner v. Wattenwyl (1907: 181) contained in Phryganistria, but other members have the medioventral carina set with minute spines. Although Phobaeticus Brunner v. Wattenwyl has subsequently been transferred from Baculini Bradley & Galil, 1977 (= Clitumnini) to Pharnaciini by Brock (1996: 26) no author has so far identified the close relation between Phryganistria and Phobaeticus, why Phryganistria has as yet remained in Clitumnini. It’s affiliation to Pharnaciini is however obvious, why Phryganistria is here transferred to Pharnaciini. All former authors have erroneously interpreted the short median segment as a link to Clitumnini, as did Stål (1875: 6) himself. Distribution (Fig. 85): So far recorded from NE-India, the southern flanks of the Himalaya (Bhutan, Sikkim, Darjeeling), Bangladesh, Myanmar, N-Thailand, Laos, Vietnam and southern China (Guangxi). One species has most certainly been erroneously recorded from Peninsular Malaysia and Java (these two records are excluded in Fig. 85). Species included: 1. Phobaeticus fruhstorferi Brunner v. Wattenwyl , 1907: 184. 2. Phryganistria grandis Rehn, 1906: 279. 3. Phryganistria guangxiensis Chen & He, 2007: 337, figs. 308 a-b. 4. Phobaeticus heusii Hennemann & Conle, 1997b: 505 figs. 1–5. 5. Phryganistria longzhouensis Chen & He, 2007: 338, figs. 309 a-d.


[Myanmar & S-China (Guangxi)] [Vietnam, Laos & Myanmar] [S-China (Guangxi)]* [Northern Vietnam] [S-China (Guangxi)]*

HENNEMANN & CONLE

6. Bacteria virgea Westwood, 1848: 66, pl. 32: 2. = Bacteria sarmentosa Westwood, 1848: 65, pl. 32: 1.

[NE-India, Bhutan, Sikkim, Darjeeling, Bangladesh & Pulau Penang]

Keys to the species of Phryganistria Stål, 1875

&& 1. Tergites III–VI parallel-sided; subgenital plate short, indistinctly projecting over apex of anal segment, not lanceolate ................................................................................................................................................2 - Tergites III–VI laterally expanded and gently rounded; subgenital plate long and lanceolate, distinctly projecting over apex of anal segment (Fig. 286) ..................................................................................virgea 2. Median segment at best 1/3 the length of metanotum; dorsal carina of meso- and metabasitarsus smooth; very large (body length > 207.0 mm) ...........................................................................................................3 - Median segment > 1/3 the length of metanotum; dorsal carina of meso- and metabasitarsus serrate; smaller species (body length 197.0 mm) ..................................................................................... fruhstorferi 3. Tergites II–VII smooth; lobes of praeopercular organ very long and back-curving (Fig. 280); posterolateral angles of anal segment tapered (Fig. 289) .................................................................................. grandis - Tergites II–VII with a posteromedian granule; lobes of praeopercular organ shorter, triangular, (Fig. 282); posterolateral angles of anal segment expanded and rounded (Fig. 288)............................................. heusii

%% 1. Meso- and metafemora robust and considerably broader than corresponding tibiae; outer ventral carinae with a prominent sub-apical spine ................................................................................................................3 - Meso- and metafemora slender, not considerably broader than corresponding tibiae; no sub-apical spine on outer ventral carinae ................................................................................................................................2 2. Median segment about 1/3 the length of metanotum; body plain brown; posteroventral carina of profemora dentate (Fig. 130)............................................................................................................................virgea - Median segment 2/5 the length of metanotum; thorax black laterally; posteroventral carina of profemora unarmed (at best with two single teeth); N-Myanmar (Fig. 132) ................................................ fruhstorferi 3. Profemora black with base pale straw; posteroventral carina of profemora sparsely dentate; cerci short and conical, not reaching apex of anal segment (Fig. 283) ......................................................................... heusii - Profemora plain brown; posteroventral carina of profemora unarmed; cerci elongate and slender, ± reaching apex of anal segment (Fig. 281) .................................................................................................. grandis

Phryganistria fruhstorferi (Brunner v. Wattenwyl , 1907) comb. nov. (Figs. 131–132, 284–285, 396) Phobaeticus fruhstorferi Brunner v. Wattenwyl , 1907: 184. LT [by present designation], &: Mt. Cariani, D. Tornatore 1898; Phobaeticus fruhstorferi Br., Type, Birma (MCSN). [Not: PLT’s in NHMW; all are Phryganistria grandis Rehn, 1906, see comments below] Brock, 1996: 28. Brock, 1998a: 30. Otte & Brock, 2005: 268. Phryganistria grandis, Günther, 1935b: 124. [Erroneous synonym] [Not: Phobaeticus frühstorferi, Vanschuytbroeck & Cools, 1981: 16. Misidentification = Phryganistria grandis Rehn, 1906] [Not: Phobaeticus fruhstorferi, Chen, 1993: 87. Misidentifiaction. Specimens from China (Guangxi) are most likely



201

Phryganistria grandis Rehn, 1906] Further material: [2 %%]: N-MYANMAR: 1 %: Myanmar, Tenasserim, leg. Lehmann 20.III.1994 (coll. FH, No. 0210-1); 1 %: N-Myanmar, Tenasserim Highland, Dawna, leg. Lehmann 27.–28.V.1996 (coll. FH, No. 0210-2).

Diagnosis: Distinguished from all other species in the genus by the relatively longer median segment, which is about a little more than 1/3 the length of the metanotum. && are similar to Ph. grandis Rehn, 1906 and Ph. heusii (Hennemann & Conle, 1997) but easily distinguished from both species by: the smaller size (body length < 200 mm); smaller lobes of the praeopercular organ (Fig. 284); relatively shorter and broader legs and tarsi and having only 4–5 prominent spines on the posteroventral carina of profemora. %% are similar to the type species Ph. virgea (Westwood, 1848) with which they have the slender meso- and metafemora and lack of a prominent sub-apical spine on the two outer ventral carinae of these femora in common. They however differ by: the slightly longer median segment; black lateral surfaces of the thorax; unarmed posteroventral carina of the profemora (at best with two single teeth); less numerous and less distinct serrations of the anterodorsal carina of the profemora; slightly more tapered semi-tergites of the anal segment and longer cerci (Fig. 285). Etymology: Dedicated to H. Fruhstorfer who collected vast amounts of insects in the Oriental Region during the early 19th century, which he sold to most major museums in Europe. Description: && (Fig. 131): Rather small (body length 197.0 mm) and moderately slender for the genus (maximum body width 6.5 mm), surface smooth and slightly glabrous. General colouration of body and legs almost plain ochraceous mid brown, abdomen slightly darker (slightly discoloured in the LT caused by provisional conservation in alcohol). Eyes dark reddish brown. Head: Oval, 1.6x longer than wide and slightly narrowed posteriorly with the vertex very gently rounded. Posterior margin with a pair of impressions which continue towards the eyes as an impressed, longitudinal line and a slightly impressed coronal line. Between the bases of the antennae with an oval, transverse impression and two very faint impressions between the eyes. Eyes of average size, circular and moderately convex; their length contained almost 3x in that of cheek. Scapus dorsoventrally flattened, 2x longer than wide and rectangular (remaining antennomeres lacking in the LT). Thorax: Pronotum narrower and shorter than head, anterior margin slightly narrower than posterior margin, 1.4x longer than wide. Anterior margin raised and followd by a deep transverse furrow. Median transverse depression indistinct and slightly curved, not reaching lateral margins of segment. Mesothorax 2.2x longer than head and pronotum combined, mesonotum parallel-sided except for being very indistinctly broadened posteriorly. Metanotum less than half the length of mesonotum, parallel-sided and almost 3x longer than wide. Pleurae and sterna simple. Abdomen: Median segment slightly more than 1/3 the length of metanotum, rectangular, 1.3x longer than wide. Segments II–VI slightly increasing in length and very slightly, gradually narrowing (II–IV broadest), II 1.5x, VI almost 3x longer than wide. Tergite VII about ¾ the length of VI and dilated into a narrow, rounded lobe posterolaterally. Tergite VI with a minute posteromedial tubercle. Praeopercular organ formed by a pair of moderately sized, carinated spines at posterior margin of sternum VII. Tergite VIII distinctly narrower than previous, slightly shorter than VII, strongly convex and constricted medially, 3.5x longer than wide. IX half the length of VIII, almost rectangular. Anal segment longer than IX, parallel-sided, with a faint median carina and a bold triangular median incision at posterior margin. Posterolateral angles narrowed and rounded apically. Supraanal plate small, triangular and with a fine median carina, not projecting over apex of anal segment. Cerci small, oval in cross-section, tapered towards the apex. Gonapophyses elongate, up-curving and slightly projecting over apex of anal segment. Subgenital plate keeled, boat shaped and reaching to apex of anal segment (Fig. 284).


HENNEMANN & CONLE

Legs: All moderately long and rather robust. Profemora longer, mesofemora as long as mesonotum, metafemora reaching about half way along abdominal segment IV and hind legs projecting over apex of abdomen. Anterodorsal carina of profemora armed with 10–11 prominent, triangular serrations. Posteroventral carina smooth except for 4–5 distinct, pointed teeth. Posteroventral carina of protibiae minutely serrate, remaining carinae unarmed. Two outer ventral carinae of meso- and metafemora each set with 10–14 triangular, black tipped teeth in almost regular distance to each other. Dorsal carinae smooth except for 2–4 small and paired teeth. Medioventral carina of femora faint and set with 5–6 minute but pointed spines. Ventral carinae of meso- and metatibiae serrate, dorsal carinae very sparingly and minutely dentate. Probasitarsus as long as remaining tarsomeres combined, dorsal carina strongly raised but of uniform height. Meso- and metabasitarsi as long as following three segments combined, with the dorsal carina gently raised and rounded, all carinae serrate. %% (Fig. 132): Rather small (body length 143.5–144.0 mm) and very slender for the genus (maximum body width 2.3 mm), body surface smooth and glabrous, legs all very long and slender. General colouration of body and legs dark yellowish brown, legs with a slight greenish was. Cheeks occasionally with a black postocular line (coll. FH, No. 0210-2). Pronotum black with a bold, longitudinal greyish stripe medially. Mesoand metathorax with a bold black longitudinal lateral stripe, which covers the complete pleurae, sterna as well as the extreme lateral margins of the meso- and metanotum. Abdominal tergites dark brown at their hinder margin. Tergites IX and X each with a longitudinal, pale marking laterally. Tibiae each with three faint greyish transverse bands, the basal one being the most prominent. Basal half of all basitarsi and bases of profemora pale greyish to straw. Antennae dark brown, the ventral surfaces off all segments, except the scapus and pedicellus, black. Head: Generally as in && but slightly less than 1.5x longer than wide. Eyes large, very prominent, circular and projecting hemispherically; their length contained about 2x in that of cheek. Antennae reaching to posterior margin of metanotum (broken in both examined specimens). Scapus dorsoventrally flattened, 2x longer than wide, rectangular. Pedicellus cylindrical and less than half the length of scapus. Following antennomeres increasing in length towards apex of antennae. Thorax: Pronotum slightly shorter and narrower than head, 1.4x longer than wide, posterior margin broader than anterior margin; the latter raised. Median transverse depression distinct, gently curved but not reaching lateral margins of segment. Mesothorax more than 3x longer than head and pronotum combined; mesonotum parallel-sided except for being very slightly broadened posteriorly. Mesosternum in posterior portion with a hint of a median carina, otherwise smooth. Metanotum about half the length of mesonotum, about 8.5x longer than wide and very gently constricted medially. Posterior margin at each outer angle with a small, rounded hump (residues of the alae). Metasternum simple. Abdomen: Median segment about 2/5 the length of metanotum, narrowed medially and almost 4x longer than wide. Segment II–VI roughly of equal length, about 4.5x longer than wide and gently constricted medially. VII ¾ the length of previous, 3.5x longer than wide and slightly widened towards posterior margin. Tergite VIII ¾ the length of VII, 2.5x longer than wide, strongly convex and broadened towards posterior margin. IX slightly shorter and distinctly narrower than VIII, 3x longer than wide and strongly constricted medially. Anal segment conspicuously longer than IX, laterally compressed, strongly keeled and split. Semi-tergites elongate and distinctly tapered towards a narrow, rounded apex (Fig. 285). Interior surfaces swollen apically and densely covered with minute black teeth. Cerci very elongate, slightly lanceolate, slightly in-curving at the apex and almost reaching apex of anal segment. Poculum moderately convex, cup-like and with a blunt, rounded central hump; reaching about half way along tergite IX. Legs: All very long and slender. Profemora slightly longer than head, pro- and mesonotum combined, mesofemora as long as combined length of pro- and mesonotum, metafemora reaching about half way along abdominal segment VI and metatibiae projecting considerably over apex of abdomen. Armature generally as in && but less prominent. The spination of the meso- and metatibiae in particular considerably less distinct.



203

Anterodorsal carina of profemora with 6–11 black, moderately sized triangular teeth. Posteroventral carina smooth (coll. FH, No. 0210-1) or with two single black teeth (left profemur of coll. FH, No. 0210-2). Probasitarsus slightly longer than remaining tarsomeres combined, unarmed. Meso- and metabasitarsi as long as remaining tarsomeres combined, unarmed. Comments: When Brunner v. Wattenwyl (1907: 184) described Phobaeticus fruhstorferi he did not recognize that his type series consisted of two distinct species. Nor did Günther (1935b: 124), who synonymised Ph. fruhstorferi Brunner v. Wattenwyl with Phryganistria grandis Rehn, 1906 but without designating a lectotype. Critical and detailed examination of the complete type-series, a & from Myanmar in MCSN as well as 3 %% and a penultimate instar & nymph from Tonkin (= Northern Vietnam) in NHMW have confirmed Günther’s synonymy only in part. Only Brock (1996: 28, 1998b: 30) stated that the & from Myanmar in MCSN was very unlikely to represent the same species as the remaining type series. In fact, the MCSN & is obviously a distinct species while the remaining syntypes from Vietnam in NHMW all represent Ph. grandis Rehn. Thus, to fix Brunner v. Wattenwyl’s species and in order to prevent the need of a new specific name, the & in MCSN specimen is here selected as the lectotype of Ph. fruhstorferi Brunner v. Wattenwyl. Brock (1998b: 30) stated the specimens from Tonkin, Chiem Hoa in ISNB to be likely syntypes, but as these were not listed in the original description by Brunner v. Wattenwyl (1907: 184) nor bear any hand-written labels of that author, their type-status is rather doubtful. Examination has shown all specimens traced in ISNB (2 %%, 1 &) to represent Ph. grandis Rehn. Eggs unknown. Distribution (Fig. 396): N-Myanmar (Highland of Tenasserim: Mount Cariani & Dawna). Number of specimens examined: 3 TABLE 31. Measurements [mm] of Phryganistria fruhstorferi (Brunner v. Wattenwyl, 1907) comb. nov.

&, LT

%%

(MCSN)

(coll. FH)

Body

197.0

143.5–144.0

Head

7.0

5.0–5.1

Pronotum

6.2

4.1–4.2

Mesonotum

41.0

32.2–32.8

Metanotum

20.9

16.8–17.3

Median segment

7.0

7.8–8.0

Profemora

55.0

46.5–48.0

Mesofemora

41.0

37.8–40.5

Metafemora

49.5

44.5–46.5

Protibiae

61.0

56.0–64.0

Mesotibiae

40.5

38.0–45.0

Metatibiae

50.0

47.6–55.0

Antennae

-

> 51.0

Phryganistria grandis Rehn, 1906 (Figs. 126, 186–187, 280–281, 289, 397) Phryganistria grandis Rehn, 1906: 279, figs. 1 & 2 (%). HT, %: Tonkin, Indo-China; Phryganistria grandis Rehn, Type (ANSP). Otte, 1978: 78.


HENNEMANN & CONLE

Hennemann et al., 2008: 24. Phobaeticus grandis, Günther, 1935b: 124. Brock, 1996: 28. Otte & Brock, 2005: 268. Phibalosoma acanthopus, Wood-Mason, 1877a: 161. Wood-Mason, 1877b: 346. Phobaeticus fruhstorferi Brunner v. Wattenwyl , 1907: 184 (in part – only PLT’s in NHMW). PLT, %: Tonkin, Montes Mahokon, April. Mai, 2–3000m, H. Fruhstorfer; Collectio Br. v. W.; det. Br. v. W. Phobaeticus fruhstorferi; 24.418 (NHMW, No. 307); PLT, 2 %%: Central-Tonkin, Chiem-Hoa, Aug. Sept., H. Fruhstorfer; Collectio Br. v. W.; det. Br. v. W. Phobaeticus fruhstorferi (NHMW, No. 307); PLT, & (penultimate instar nymph): Tonkin, Than-Moi, H. Fruhstorfer ; Collectio Br. v. W. ; 24.419, det. Br. v. W. Phobaeticus fruhstorferi (NHMW, No. 307). [Not: LT, &: Mt. Cariani, D. Tornatore 1898, Phobaeticus fruhstorferi Br., Type, Birma (MCSN)—erroneous synonym of Günther, 1935b: 124, see comments below] Brock, 1996: 28. Phobaeticus frühstorferi, Vanschuytbroeck & Cools, 1981: 16. ? Phobaeticus fruhstorferi, Chen, 1993: 87. [Record from South China (Guangxi), see comments below] Further material: [22 &&, 21 %%, 1 egg]: VIETNAM: 1 %, 1 &: Tonkin, Chiem-Hoa, leg. Fruhstorfer VIII–IX. (SMNS); 1 %: Tonkin, Than Moi, leg. Fruhstorfer (ZMUH); 1 &: Tonkin, Chiem Hoa, leg. Fruhstorfer 15.IV. 1902 (ZMUH); 2 %%, 2 &&: Tonkin, Chiem-Hoa, Aug. Sept., leg. H. Fruhstorfer (MHNG); 1 %: Tonkin, Than-Moi, Juni-Juli, H. Fruhstorfer; 1 %: Tonkin, coll. Le Moult; 1 &: Tonkin, Chiem Hoa, Aug.–Sept., H. Fruhstorfer (ISNB); 3 %%, 2 &&: Tonkin, Juni–Juli, H. Fruhstorfer; 1 &: Tonkin, Chiem Hoa, Aug.– Sept., H. Fruhstorfer (RMNH); 1 %: Central-Tonkin, Chiem-Hoa, Aug. Spet., leg. H. Fruhstorfer, L. 845 (SMTD); 1 &: Tonkin, Fruhstorfer (SMTD); 1 %: Tonkin, Than Moi, Fruhstorfer (MNHU); 1 %: Tonkin, det. Willemse (MNHU); 1 %, 1 &: Central-Tonkin, Chiem-Hoa, Aug./ Sept., H. Fruhstorfer (ETHZ); 1 %: 27, Central-Tonkin, Chiem-Hoa, Aug. Sept., H. Fruhstorfer (SMFM); 1 & + 1 egg (ex ovipositor), 1 %: Cochinchina, Mt. Bavi, 1935, S. Massey eff., BM 1937-658 (BMNH); 1 &: Cochinchina, Tuyen Quang, 1936, S. Massey eff., BM 1937-658 (BMNH); 1 %: Tonkin, 1935-448, Phryganistria sarmentosa BMNH (BMNH); 1 &: Tonkin, 1935-448, Phryganistria fruhstorferi (BMNH); 1 &: Vietnam: 55 km NNW Tam Dao, 26.VII.–10.VIII.1998, A. Napolov, BMNH(E) 2005-98, Phobaeticus grandis det. P. Brock 2005 (BMNH); 1 %: Museum Paris, Haut-Tonkin et Bas-Yunnan, Entre Mau-Hoa, Huong-Num (Prés Lao-Kay) et Bau-HauKhun Lieut, Lesourt 1905 (MNHN); 1 %: Museum Paris, Tonkin Hajiank, H. Donchier 1924 (MNHN); 1 &: Museum Paris, Tonkin N., Reg. Ma-Giang, Siebens Olivier 1916 (MNHN); 1 &: Museum Paris, Haut-Tonkin, Reg. De Bac Ken, Ma-Giang Guam-Ba et Yen Minh, F. de Broissia 1918 (MNHN); 1 %, 1 &: Tonkin (ZSMC); 1 % : Tonkin, Than Moi, Juni–Juli, H. Fruhstorfer (ZSMC); 1 &: Tonkin, Chiem Hoa, Aug.–Sept., H. Fruhstorfer (ZSMC); 1 %, 1 &: Central Tonkin, Chiem Hoa, Aug. Sept. H. Fruhstorfer (MNCN). MYANMAR: 1 &: Upper Burma, Nam Tamai Valley, 22.9.1938, R. Kaulbach leg., BM 1938-741, att. 3000ft., Lat. N 27°42`, Long 97°54` (BMNH). NO DATA: 1 %: no data (MNHU); 1 %, 1 &: no data (ZSMC); 1 &: no data (SMFM); 1 &: no data (BMNH).

Diagnosis: Very closely related and certainly the adelphotaxon of Ph. heusii (Hennemann & Conle, 1997), with which it is sympatric in Northern Vietnam. It however frequently differs by: the more slender body; relatively longer and more slender legs and tarsi as well as the slightly more globose head of both sexes. %% furthermore differ by: the plain brown profemora which have the posteroventral carina unarmed; relatively smaller eyes; more narrow apices of the semi-tergites of the anal segment and longer, more slender cerci which almost reach to the apex of the anal segment (Fig. 281). && may be distinguished by: the lack of a posteromedian tubercle on abdominal tergites II–VI; longer and more slender lobe-like spines of the praeopercular organ (Fig. 280) and much deeper posteromedian excavation of the anal segment (Fig. 289). Eggs differ from those of Ph. heusii by the smaller dimensions, slightly more ovoid capsule and relatively smaller micropylar plate (Figs. 186–187). Etymology: “Grandis” (lat. = large) refers to the large size of %%.



205

Description: &&: Long to very long (body length 234.5–253.0 mm) and slender species (maximum body width 8.0–9.0 mm) with the body surface slightly glossy. General colouration of body and legs plain mid to dark brown, ochracheous, brownish green or dull green, the abdomen in particular furnished with several small blackish speckles. Armature of legs dark greyish green with black points. Antennae dark brown. Head: Globose, oval in dorsal aspect, roughly 1.4x longer than wide and vertex gently convex. Coronal line distinct, impressed and on each side accompanied by a pair of faint dark brown, longitudinal lateral lines. Posterior margin of head capsule with three short, longitudinal impressions. Between the bases of antennae with an oval, slightly transverse depression followd by two slightly raised, mid brown areas and a pair of very faint impressions. Eyes rather small, circular and moderately convex; their length contained about 3.3x in that of cheek. Antennae reaching to posterior margin of mesonotum, with the antennomeres increasing in length towards apex of antennae. Scapus dorsoventrally compressed, rectangular, almost 3x longer than wide. Pedicellus slightly spherical and about 1/3 the length of scapus. Thorax: Pronotum shorter and narrower than head, 1.6x longer than wide, posterior margin slightly broader than anterior margin. Anterior margin arcuate and followd by a transverse furrow. Anterior half with a faint trace of an impressed longitudinal median line. Median transverse depression distinct, slightly curved but not reaching lateral margins of segment. Mesothorax almost 3x longer than head and pronotum combined. Mesonotum parallel-sided except for being slightly widened posteriorly. Metanotum parallel-sided, about 4x longer than wide and 3/5 the length of mesonotum. Pleurae and sterna unarmed. Abdomen: Median segment indistinctly longer than wide, roughly ¼ the length of metanotum and slightly trapezoidal, with posterior margin broader than anterior margin. Segments II more than 2x longer than median segment, II–VI increasing in length, II 2x, VI more than 3x longer than wide, all parallel-sided and of uniform width. Tergite VII about ¾ the length of VI with lateral margins elevated to form a slight, rounded lobe posteriorly. Praeopercular organ formed by a pair of very prominent, long, carinate, back-curving, lobe-like spines at posterior margin of sternum VII. Tergite VIII half the length of VII, constricted anteriorly and 2x longer than wide. IX indistinctly longer than wide, slightly more than half the length of VIII, parallel-sided. Anal segment slightly longer than IX, with a very faint, longitudinal median carina and a deep, narrow triangular excavation posteromedially; posterolateral angles triangular, slightly in-curving and swollen (Fig. 289). Supraanal plate very small, rounded and strongly keeled dorsally. Cerci small, sub-oval in cross-section and tapered towards a narrow apex; finely bristled. Subgenital plate moderately convex, strongly keeled, boatshaped and projecting over apex of abdomen by at least 1/3 the length of anal segment (Fig. 280). Legs: All long and moderately slender. Profemora almost as long as head, pro- and mesonotum combined, mesofemora slightly shorter than mesothorax, metafemora reaching about half way along abdominal segment V and metatibiae slightly projecting over apex of abdomen. Anterodorsal carina of profemora armed with 11– 17 prominent triangular serrations which decrease in size towards apex of femur, posterodorsal carina set with 5–9 minute teeth. Posteroventral carina with 13–20 small to medium-sized but acute, triangular teeth. Medioventral carina of protibiae sparingly and rather minutely dentate, remaining carinae unarmed. Two outer ventral carinae meso- and metafemora armed with 12–20 triangular teeth of average size. Dorsal carinae each with only 2–6 teeth, the basal one of the posterior carina sometimes conspicuously enlarged and triangular. Medioventral carina armed with a longitudinal row of 6–12 minute spines. All carinae of meso- and metatibiae serrate, although considerably more sparsely on dorsal carinae. Posterodorsal carina often with a ± prominently enlarged, triangular tooth pre-medially and gently rounded apically. Probasitarsus a little longer than remaining tarsomeres combined, dorsal carina very Anacardiaceae raised, all carinae unarmed. Meso- and metabasitarsi as long as following three tarsomeres combined, dorsal carina gradually raised towards the apex, ventral carinae minutely dentate. %% (Fig. 126): Long to very long (body length 171.5–232.0 mm) and moderately slender species (maximum body width 7.0–8.5 mm), with body surface smooth and glabrous. General colouration of body and legs, ferruginous, reddish or yellowish pale to mid brown. Greater portion of the mesonotum except anterior and


HENNEMANN & CONLE

posterior margin as well as the anterior ¾ of the metanotum covered by a washed malachite green, bluish green or greyish marking; may be very faint in some specimens. Two outer ventral carinae of meso- and metafemora dull green. Complete armature of legs tipped with black. Eyes reddish brown. Antennae dark reddish brown to dark brown. Head: Generally as in && but slightly more decidedly narrowed posteriorly. Posterior margin with four faint impressions, which anteriorly continue as four sub-parallel, slightly impressed longitudinal lines. Eyes rather small, circular and projecting hemispherically; their length contained about 3x in that of cheek. Antennae projecting over posterior margin of median segment, otherwise as in &&. Thorax: Pronotum slightly shorter but distinctly narrower than head, 1.5x longer than wide, anterior portion narrowed, otherwise as in &&. Mesothorax 3.6x longer than head and pronotum combined and slightly swollen posteriorly, mesonotum parallel-sided except for being slightly broadened at posterior margin. Metanotum less than 2/3 the length of mesonotum and slightly constricted medially, more than 7x longer than wide. Pleurae and sterna smooth. Abdomen: Median segment about 1/4 the length of metanotum, 1.8x longer than wide and slightly trapezoidal being gently widened towards the posterior. Segment II about 1.7x longer than median segment, II–VII slightly constricted medially. III–IV subequal in length and slightly longer than II, almost 6x longer than wide. V–VI subequal and slightly longer than III–IV, almost 6.5x longer than wide. VII about 2/3 the length of VI and broadened posteriorly. Tergite VIII half the length of VII, strongly convex and gradually widening towards posterior margin. IX strongly convex and slightly shorter than VIII, anterior margin distinctly broader than posterior margin. Anal segment laterally compressed, strongly tectiform and carinate dorsally. Semi-tergites roughly triangular, very broad anteriorly and strongly tapered in apical portion, the apices narrow and rounded (Fig. 281). Interior surfaces apically swollen and covered with several minute, black and back-curving teeth. Cerci elongate, sub-oval in cross-section, gradually tapered towards apex and roughly reaching apex of anal segment. Poculum strongly convex with a distinct transverse carina and a blunt rectangulate extension anteriorly; reaching about 2/3 the way along tergite IX. Legs: All very long and slender, meso- and metafemora robust, conspicuously swollen and considerably broader than corresponding tibiae. Profemora longer than combined length of head, pro- and mesonotum, mesofemora a little longer than mesothorax, metafemora almost reaching posterior margin of abdominal segment V and metatibiae projecting considerably over apex of abdomen. Anterodorsal carina of profemora with 14–20 small triangular serrations, posterodorsal and posteroventral carinae unarmed. All carinae of protibiae unarmed. Two outer ventral carinae of meso- and metafemora armed with 12–20 moderately sized, triangular teeth, each sub-apically with a prominent, acutely dagger-like spine. Medioventral carina with a longitudinal row of 5–10 very minute spines. Dorsal carinae smooth except for 1–3 minute teeth. Ventral carinae of mesoand metatibiae densely, dorsal carinae rather more sparsely dentate. Probasitarsus very elongate and slender, about 1.5x longer than remaining tarsomeres combined, all carinae smooth. Mesobasitarsi about as long as, metabasitarsi a little longer than remaining tarsomeres combined; ventral carinae with a few minute teeth in basal half. Eggs (Fig. 188–189): One fully developed egg was removed from the ovipositor of the & from Cochinchina (Mount Bavi) in BMNH (accession No. BM 1937-658) and served for the following description. Medium-sized, capsule oval in lateral aspect and slightly laterally compressed, oval in cross-section. Capsule with a very faint keel on dorsal and ventral surfaces and polar-area, beginning and ending at operculum. Capsule surface slightly glossy and minutely granulated (70x). General colouration of capsule and micropylar plate creamish dark brown, operculum black. Micropylar plate elongate and covering about 2/3 the length of capsule, tapered towards anterior end and with a wide gap posteromedially. Outer margin of plate black. Micropylar cup placed in the posterior gap of plate, small and dark reddish brown. Median line on a distinctly raised hump, almost reaching polar-area. Operculum oval and slightly convex. Capitulum large, dark brown, irregularly conical and with several large lateral impressions. Stalk obvious, whitish.



207

Measurements [mm]: length (incl. capitulum) 5.4, length 4.4, width 2.9, height 3.6, length of micropylar plate 2.7 . Comments: Brunner v. Wattenwyl (1907: 184) appears to have not been aware of Rehn’s 1906 publication, described Phobaeticus fruhstorferi from a series of specimens from N-Vietnam and Myanmar in NHMW and MCSN and did furthermore not recognize that the type-series consisted of two distinct species (see comments on Phryganistria fruhstorferi). The & from Birma (= Myanmar) in MCSN is clearly distinct from the remaining types from N-Vietnam in NHMW and here selected as the lectotype of Ph. fruhstorferi Brunner v. Wattenwyl, 1907. All paralectotypes (3 %%, 1 & penultimate instar nymph) in NHMW clearly represent Phryganistria grandis Rehn, 1906. Günther (1935b: 124) had obviously not seen the specimen in MCSN and synonymised both species. However, as Günther (1935b) did not designate a lectotype his synonymization is invalid, and Ph. fruhstorferi Brunner v. Wattenwyl here treated as a valid species with the MCSN specimen selected as the LT. Wood-Mason (1877a: 161) recorded a & of what he believed to be Phibalosoma acanthopus Burmeister, 1838 (= Phobaeticus serratipes (Gray, 1835)) from Moolai to Moolat in Upper Tenasserim, Northern Myanmar. Already Redtenbacher (1908: 453) doubted the identity of Wood-Mason’s specimen and designated it as a syntype of his newly described Pharnacia ingens. The brief diagnosis provided by Wood-Mason however clearly shows this & to represent a distinct species and to be even generically different from the LT of Ph. ingens from the Malabar Coast of SW-India. The prominent, bifid praeopercular organ, short median segment, locality and body length which Wood-Mason stated to be 260.0 mm obviously place it in Phryganistria Stål, 1875. TABLE 32. Measurements [mm] of Phryganistria grandis Rehn, 1906.

%, HT (ANSP)

%%

%

Body

198.0

171.5–232.0

207.5

Head

7.5

-

Pronotum

7.0

Mesonotum

&&

&

&&**

(SMFM)

(China, Guangxi)

234.5–253.0

236.0

207.0–214.0

8.2

-

10.2

-

5.1–7.8

6.8

7.7–9.5

8.0

8.5–9.0

46.0

38.0–47.5

49.0

51.5–55.0

51.0

41.0–41.5

Metanotum

26.0

20.0–31.3

28.3

28.0–30.5

29.0

16.0–18.0

Median segment

8.5

6.6–8.0

6.8

7.0–8.5

7.3

12.5–13.0

Profemora

63.8

54.2–69.0

69.0

56.5–72.5

63.1

50.0–51.0

Mesofemora

48.5

41.5–51.0

49.4

48.0–53.5

49.8

38.0–41.0

Metafemora

58.0

48.0–60.0

59.0

55.5–63.0

53.7

49.0

Protibiae

87.0

76.0–94.0

88.8

-

80.4

-

Mesotibiae

58.0

48.5–60.0

59.5

-

54.3

-

Metatibiae

70.5

62.0–73.5

78.9

-

68.0

-

Antennae

> 70.0

100.0–115.0

> 91.0

66.5–67.0

-

-

(SMFM)

* including median segment ** according to Chen (1993: 87)

Chen (1993: 87) recorded specimens, which he believed to be Ph. fruhstorferi (Brunner v. Wattenwyl, 1907), from the Guangxi autonomous region in South China close to the border to Northern Vietnam and provided a brief description of the && along with a set of measurements. Due to having a relatively longer median segment compared to both, Ph. grandis Rehn and Ph. fruhstorferi Brunner v. Wattenwyl (Tables 31– 32) the exact identity of these Chinese specimens deserves confirmation. Although they are shorter than specimens of Ph. grandis from N-Vietnam (body length 207.0–214.0 mm) the distribution rather associates them


HENNEMANN & CONLE

with Ph. grandis or perhaps Ph. guangxiensis Chen & He, 2007 rather than Ph. fruhstorferi. Unfortunately, the specimens could not be examined for this study. Almost every European museum contains specimens of Ph. grandis Rehn, most of which were collected by H. Fruhstorfer in Vietnam during the late 19th century. According to the large number of specimens collected in only a short period of time this species appears to be quite abundant in certain areas of Northern and Central Vietnam. Distribution (Fig. 397): Northern Vietnam (Mount Mahokon 2000–3000 m; Chiem-Hoa; Than Moi; MaGiang Region & Mau-Hoa: Huong-Num), Southern Vietnam (Cochinchina: Mount Bavi & Tuyen Quang) and Northern Myanmar (Upper Tenasserim: Moolai to Moolat 4000–6000 ft. [Wood-Mason, 1877a: 161] & Nam Tamai Valley 3000 ft.). South China (Guangxi)—with doubt [Chen, 1993: 87]. Number of specimens examined: 48

Phryganistria heusii (Hennemann & Conle, 1997) comb. nov. (Figs. 127–128, 190–191, 277–279, 282–283, 288, 355, 359, 398) Phobaeticus heusii Hennemann & Conle, 1997b: 505, figs. 1–5. HT, %: N-Vietnam, Cuc Phuong, ex. Zucht P. Heusi 1996 (ZSMC, ex coll. FH, No. 0240-1); AT, & (+ 4 eggs): N-Vietnam, Cuc Phuong, ex. Zucht P. Heusi 1996 (ZSMC, ex coll. FH, No. 0240-2). Otte & Brock, 2005: 268. Bresseel, 2007: 8. Further material [7 &&, 10 %%, eggs]: N-VIETNAM: 6 %%, 6 &&, eggs: ex Zucht: F. Hennemann 2002, Herkunft: N-Vietnam, Ninh Binh Prov., Nho Quan District, Cuc Phuong (coll. FH, No’s 0240-3 to 14, ED); 1 &, 4 %%: ex Zucht: F. Hennemann 2002, N-Vietnam, Ninh Binh Prov., Nho Quan District, Cuc Phuong (coll. OC).

Diagnosis: Very closely related and certainly the sister-taxon of Ph. grandis Rehn, 1906, with which it is sympatric in Northern Vietnam, but differing by: the more robust body; relatively shorter and more robust legs and tarsi as well as the slightly less globose head of both sexes. %% clearly differ by: the black profemora which have the basal portion pale straw and the posteroventral carina sparsely dentate; relatively larger eyes; broader apical portion of the semi-tergites of the anal segment and distinctly shorter, more obtuse and conical cerci which clearly stay beyond the apex of the anal segment (Fig. 283). && may be distinguished by: the posteromedian tubercle on abdominal tergites II–VI; shorter and more broader lobe-like spines of the praeopercular organ (Fig. 282) and much more shallow posteromedian excavation of the anal segment (Fig. 288). Eggs differ from those of Ph. grandis by the larger dimensions, less ovoid capsule and relatively larger, more elongate micropylar plate (Figs. 188–189). Etymology: Dedicated to Peter Heusi (Uster, Switzerland), who first imported culture-stock of this magnificent species to Europe and provided the type specimens. Description: && (Fig. 127): Very long (body length 239.0–293.0 mm) and slender species (maximum body width 7.0–9.5 mm), complete surface smooth and slightly glabrous. General colouration of body and legs variable, ranging from more or less plain pale to dark brown or green. Abdomen in particular ± decidedly furnished with small black or brown speckles. Lateral surfaces of head usually with an indistinct longitudinal dark brown stripe above the eyes, which consists of numerous small dark brown speckles. Eyes dark reddish brown. Antennae brown with ventral surfaces except scapus and pedicellus black. Head (Fig. 278): Oval, 1.3x longer than wide, broadest behind the eyes and slightly narrowed posteriorly, vertex almost flat and with a well decided impressed coronal line. Posterior margin usually with a further longitudinal, sub-parallel impression. A deep, transverse impression between the bases of antennae. Eyes of averREVISION OF PHARNACIINI


209

age size, circular, and projecting hemispherically; their length contained almost 3x in that of cheek. Antennae almost reaching posterior margin of mesonotum, consisting of about 38 antennomeres. Length of antennomeres first increasing in length, then decreasing owards apices of antennae. Scapus roundly rectangular, dorsoventrally compressed and about 2.5x longer than wide. Pedicellus slightly spherical, 1/3 the length of scapus. Third antennomere cylindrical, narrower but slightly longer than pedicellus. Thorax: Pronotum almost as long but conspicuously narrower than head, about 1.7x longer than wide, slightly gradually broadening towards the posterior. Anterior margin gently arcuate and followd by a transverse furrow; dorsal surface otherwise with an impressed longitudinal median line. Median transverse depression distinct, slightly curved and almost reaching lateral margins of segment; median line clearly visible. Mesothorax about 2.7x longer than head and pronotum combined, mesonotum parallel-sided. Metathorax slightly broadened posteriorly, metanotum 4x longer than wide with posterior margin slightly broader than anterior margin. Sterna and pleurae smooth. Abdomen: Median segment slightly longer than wide, rectangular, slightly more than ¼ the length of metanotum. Segments II–VI increasing in length, II 2x, VI 3.5x longer than wide. Tergite VII ¾ the length of IV with lateral margins gently rounded posteriorly. Tergites II–VIII with a transverse posteromedial tubercle, which is most prominent on IV–VI. Praeopercular organ formed by a pair of prominent, triangular lobe-like spines at posterior margin of sternum VII (Fig. 359). Remaining sternites smooth. Tergite VIII 2/3 the length, 2.5x longer than wide, strongly convex, parallel-sided and slightly narrower than VII. IX a little more than half the length of VIII, 1.3x longer than wide and strongly convex. Anal segment slightly longer than IX flattened and slightly broadening posteriorly, with a fine longitudinal median carina and a wide and shallow excavation poseromedially (Fig. 288). Supraanal plate small, transverse and slightly projecting over apex of anal segment. Cerci small and conical, finely bristled. Gonapophyses up-curving and almost reaching tip of subgenital plate. Subgenital plate keeled, boat-shaped and at best very slightly projecting over posterior margin of anal segment (Fig. 282). Legs: All very long and moderately slender. Profemora about as long as head, pro- and mesonotum combined, mesofemora Anacardiaceae shorter than mesonotum, metafemora reaching about 1/3 the way along abdominal segment V and metatarsi slightly projecting over apex of abdomen. Anterodorsal carina of profemora with 11–18 prominent, acutely triangular serrations which decrease in size towards apex of femur, posterodorsal carina set with 3–5 very small teeth. Posteroventral carina with 17–20 medium-sized but pointed teeth. Posteroventral carina of protibia sparsely and minutely serrate, remaining carinae unarmed. Two outer ventral carinae of meso- and metafemora armed with 13–19 pointed triangular teeth of average size, dorsal carinae each with only 2–5 minute but acutely pointed teeth. Medioventral carina with a longitudinal row of 10–18 minute spines. All ventral carinae of meso- and metatibiae densely serrate, dorsal carinae with only a few rather small teeth. Posterodorsal carina rounded and bi-dentate apically. One specimen (coll. FH, No. 0240-5) has the posterodorsal carina very gently but widely raised and terminating in a small tooth pre-medially. Probasitarsus elongate and with the dorsal carina gently raised and rounded apically, slightly longer than remaining tarsomeres combined; all carinae smooth. Meso- and metabasitarsi slightly longer than following three tarsomeres combined, dorsal carina gradually raised and conspicuously rounded at the apex; ventral carinae set with a few minute teeth. %% (Figs. 128): Very long (body length 206.0–250.5 mm) and rather robust for the genus (maximum body width 8.0–9.0 mm), the meso- and metafemora conspicuously swollen and broader than corresponding tibiae. Body surface entirely smooth and glabrous. General colouration of body, mid and hind legs pale to mid ferruginous or yellowish brown. Greater portion of the mesonotum except anterior and posterior margin as well as the anterior ¾ of the metanotum covered by a washed malachite green or creamish grey marking. Profemora black except for the compressed basal portion and extreme apex both of which are pale straw; the latter occasionally with a slight pinkish wash. Protibiae dark brown and marbled with black ventrally. Eyes greyish or reddish brown. Scapus and pedicellus of same colour as body. Remaining tarsomeres very dark brown dor-


HENNEMANN & CONLE

sally, black ventrally and becoming black towards apex of antennae. Spination of legs dull red with black tips. Head (Fig. 279): Generally as in &&, but more decidedly narrowed towards the posterior; posterior margin with three small, longitudinal impressions. Transverse impression between bases of antennae less distinct. Eyes very prominent, circular and projecting hemispherically; their length contained only about 2.3x in that of cheek. Antennae generally as in && but relatively longer and slightly projecting over posterior margin of median segment. Thorax: Pronotum generally as in &&, but anterior margin more distinctly narrowed. Mesothorax very elongate, cylindrical and more than 3x longer than head and pronotum combined. Mesothorax slightly broadened posteriorly, the mesonotum gently widened in that section; a little more than 3x the length of head and pronotum combined. Metathorax constricted medially, metanotum 6x longer than wide and widened in posterior portion. Meso- and metasternum smooth, the first strongly convex. Abdomen: Median segment, a little more than ¼ the length of metanotum, 1.5x longer than wide and trapezoidal being gently broadened towards the posterior. Segment II about 1.8x longer than median segment but shorter than following. II–VI distinctly constricted medially and increasing in length, II 3.3x, VI 5x longer than wide. Tergite VII 3/5 the length of VI, slightly broader than previous and widened posteriorly. VIII strongly convex, slightly longer than wide and conspicuously broadening towards posterior margin; broadest segment. IX slightly shorter than VIII and tapered towards the posterior and with a longitudinal impression laterally. Anal segment laterally compressed, strongly tectiform and carinate dorsally. Semi-tergites roughly triangular, very broad basally and gradually tapered in apical half, the apices rounded (Fig. 283). Interior surfaces swollen apically and densely armed with minute black teeth. Cerci stout, conically tapered towards an acute tip and staying clearly beyond apex of anal segment. Poculum strongly convex, cup-like with a blunt rectangulate hump basally and reaching to posterior margin of tergite IX. Legs: All very long and rather robust, armature generally as in &&. Profemora almost as long as head, proand mesonotum combined, mesofemora about as long as mesonotum, metafemora reaching almost posterior margin of abdominal segment V and metatibiae projecting considerably over apex of abdomen. Profemora with serrations of anterodorsal carina smaller than in && and the posteroventral carina just sparsely and rather minutely dentate. Protibiae at best with a few minute teeth on posteroventral carina, otherwise unarmed. Meso- and metafemora prominently swollen and considerably broader than corresponding tibiae. Two outer ventral carinae each with a very prominent and acute, black dagger-like spine sub-apically (Fig. 277); this is sometimes accompanied by one or two conspicuously enlarged spines on metafemora. Dorsal carinae of meso- and metafemora at best with 1–3 minute teeth and those of corresponding tibiae very sparsely and minutely dentate. Probasitarsus very elongate and slender, almost 1.5x longer than remaining tarsomeres combined and with the dorsal carina very gently raised; all carinae smooth. Meso- and metabasitarsus slightly longer than remaining tarsomeres combined, otherwise as in && but dorsal carina less decidedly raised. Eggs (Fig. 190–191, 355): Large, capsule oval in lateral aspect and slightly laterally compressed, oval in cross-section. Capsule with a very shallow keel on dorsal and ventral surfaces and polar-area, beginning and ending at the operculum. Capsule surface slightly shiny and very minutely rugulose and granulated (70x). General colouration of capsule and micropylar plate creamish grey-brown, operculum blackish brown. Micropylar plate large, elongate and covering about ¾ the length of capsule, tapered towards anterior end and with a wide gap posteromedially. Outer margin of plate dark brown. Micropylar cup placed in the gap of plate, small, dark reddish brown. Median line slightly raised and short, not considerably projecting over posterior margin of plate. Operculum slightly cordiform and convex. Capitulum prominent, slightly conical and with several deep lateral impressions; greyish or reddish mid brown. Stalk obvious, whitish. Measurements [mm]: Length including capitulum 5.5–6.2, length 4.9–5.8, width 3.0–3.2, height 3.9–4.0, length of micropylar plate 4.1. Comments: Hennemann & Conle (1997b: 505) originally described Phobaeticus heusii from an F1-generation captive reared % HT, & AT and four eggs (PT’s). The original eggs were obtained from a Nature



211

Reserve Project in Cuc Phuong, Nho Quan District, Ninh Binh Province (about 100 km south of Hanoi) by Peter Heusi (Uster, Switzerland) in 1996. The type specimens have subsequently been deposited in ZSMC. With a maximum recorded body length of 250.5 mm the %% of Ph. heusii represent the longest of all known extant male Phasmatodea (see Hennemann & Conle, 1997b: 508) even being the world’s longest known male insect. && reach overall lengths of almost 500.0 mm if legs are included, a & in coll. FH (No. 0240-3) measuring 479.0 mm from the tip of the protarsi to the tip of the hind tarsi, thus being among the world’s longest extant insects. %% occur in two distinct colour morphs, either having the dorsal surface of the mesonotum and anterior ¾ of the metanotum washed malachite green or creamish grey. Eggs were imported along with stock of Tirachoidea jianfenglingensis (Bi, 1994) by P. Heusi (Switzerland) on at least two occasions. In captivity in Europe Ph. heusii has proven rather difficult to maintain in a stable culture for more than a few generations, but it still is successfully reared by several enthusiasts why it was recently included on the Phasmid Study Group culture-list as PSG 277. Oak (Quercus spp., Fagaceae) and bramble (Rubus spp., Rosacea) are accepted as alternative foodplants and very large cages are necessary to ensure successful moulting of large nymphs. Notes on breeding were provided by Bresseel (2007). Distribution (Fig. 398): Northern Vietnam (Ninh Binh Province: Nho Quan District, Cuc Phuong). Number of specimens examined: 19 TABLE 33. Measurements [mm] of Phryganistria heusii (Hennemann & Conle, 1997) comb. nov.

%, HT

&, AT

%%

&&

(ZSMC)

(ZSMC)

(coll. FH)

(coll. FH)

Body

250.5

255.0 (293.0*)

206.0–230.0

239.0–281.5

Head

10.0

13.4

8.1–9.2

11.1–14.8

Pronotum

9.5

11.3

6.6–7.2

8.6–9.5

Mesonotum

57.8

53.0

43.7–48.0

49.5–60.1

Metanotum

34.0

29.8

28.3–33.2

29.0–34.8

Median segment

9.0

8.0

7.9–8.5

7.2–8.9

Profemora

72.0

74.5

66.0–73.0

62.3–76.0

Mesofemora

61.0

55.0

48.5–54.8

48.0–57.6

Metafemora

71.5

61.0

60.1–63.7

54.5–68.0

Protibiae

101.0

83.3

82.0–87.3

75.1–94.0

Mesotibiae

63.6

56.0

52.1–54.9

47.5–58.2

Metatibiae

78.2

66.0

68.0–70.7

62.0–77.6

Antennae

> 105.0

> 68.0

94.0–110.0

70.0–73.0

Phryganistria virgea (Westwood, 1848) (Figs. 129–130, 190–191, 286–287, 399) Bacteria virgea Westwood, 1848: 66, pl. 32: 2. HT, %: Silhet; Type, Westwood, Bacteria virgea 1848, pl. 32, fig. 2, Type: Orth 535, Bacteria virgea Westwood (OXUM, No. 535). Lonchodes virgeus, Westwood, 1859: 46. Phryganistria virgea, Brunner v. Wattenwyl , 1907: 183. Günther, 1934b: 94, fig. 6 (&, abdomen) Otte & Brock, 2005: 272. Bacteria sarmentosa Westwood, 1848: 65, pl. 32: 1. HT, &: Silhet; Type, Westwood, Bacteria sarmentosa 1848, pl. 32, fig. 1, Type: Orth. 534, Bacteria sarmentosa Westwood (OXUM, No. 534). [Synonymised by Westwood, 1859: 46,


HENNEMANN & CONLE

see comments below]. Phryganistria sarmentosa, Stål, 1875: 63. Kirby, 1904a: 358. Giglio-Tos, 1910: 16. Otte & Brock, 2005: 272. [As a synonym of Ph. virgea (Westwood, 1848)] Further material: [43 &&, 20 %%, 2 nymphs, eggs]: NE-INDIA: 1 &: Coll. Br. v. W. Assam, Thorey, det. Br. v.W. Phryganistria virgea Westw., 2.701 (NHMW, No. 306); 1 &: Coll. Br. v. W. Assam, Thorey, det. Br. v.W. Phryganistria virgea Westw., 3.513 (NHMW, No. 306); 1 %, 1 &: Coll. Br. v. W. Assam, Thorey, det. Br. v.W. Phryganistria virgea Westw. (NHMW, No. 306); 1 &: Coll. Br. v. W. Assam, leg. Brunner 1866 (NHMW, No. 306); 1 %, 1 &: without data, &, det. Br. v.W. Phryganistria virgea Westw., 25. 043; 2 &&: Samarang, leg. Mason (NHMW, No. 306); 2 %%, 2 && Coll. Br. v. W. Silhet, Deyrolle, det. Br. v.W. Phryganistria virgea Westw. (NHMW, No. 306); 1 &: Silhet (NHMW, No. 306); 1 &: Indien, Assam, Liman, H. Rolle, Berlin S.W.M., ex coll. Werner, ex coll. Ebner (NHMW, No. 306); 2 && + glas-tube with eggs: Darjeeling, leg. Gutmann 28.I.1905, vend. 28.7.1905, det. F. Hennemann 1.1998 (ZMUH); 2 %%: Assam, leg. Fink 1878 (SMNS); 3 &&: Assam, leg. Dingenbach 1874 (SMNS); 1 %, 1 &: Assam, Nr. 21 & 34 (SMNS); 4 &&: Assam, leg. Diefenbach 1834 (SMNS); 1 &: Assam, 34; 1 %: Assam, 21; 1 %: Assam, leg. Finkh 1878 (SMNS); 2 %%, 3 &&: Silhet (BMNH); 1 &: N-India (BMNH); 1 &: Rothschild, Bequest B.M. 1939.-I. (BMNH); 1 &: Bacteria sarmentosa Westw. Sylhet, 48.55 (BMNH); 1 &: Silhet (OXUM); 1 &, 1 %: Assam, leg. Jerkins (OXUM); 1 &: Assam, Khasi Geb., Pres. 1907 by Dr. Brancsick (OXUM); 3 &&: Inde, Matloffinum, Musea de Stuttgart (MHNG); 1 &: Assam, Khasi Hills, Lait Pyng Kot, 16.III.1966, F. Schmid, Don F. Baud 1978 (MHNG); 1 &: Indie orient., D. Henfrey, coll. Giglio-Tos (MRSN); 1 %: Silhet (MNHN); 1 &: Lamin, Assam, H. Rolle leg., Berlin (MNHU); 1 %: Sylhet, Nr. 3200, „virgea“ det. (MNHU); 1 egg: Darjeeling, leg. Gutmann 28.I.1905, vend. 28.7.1905, ex coll. ZMUH (coll. FH, No. 0347-E); 1 &: Assam (ZSMC); 1 %: Khasi Hills, Assam, Phryganistria sarmentosa (Westw.), Det. Rehn 1934, Hebard Collection (ANSP). THAILAND [with doubt]: 1 &: Siam (BMNH). PENINSULAR MALAYSIA [with doubt]: 1 &: W-Malaysia, Pulau Penang ?, leg. Rolle (NHMW, Nr. 306). JAVA [in error ?]: 2 &&: Java, Samarang, leg. Mason 1874, coll et det. Br. v.W. Phryganistria virgea Westw. (NHMW, No. 306); 1 %: Java (MHNG). NO DATA: 1 &: 45,33 (BMNH); 1 %: 5b,8 (BMNH); 2 %%: (BMNH); 1 %: data unreadable, [5] (ZMUH); 1 &, 1 & (nymph): "Ashmol. Mus." (OXUM); 1 &, 1 %, 1 & (nymph): no data (OXUM).

Diagnosis: && are easily distinguished from all other members of the genus by the laterally broadened and rounded abdominal tergites III–VI and long, lanceolate subgenital plate, which considerably projects over the apex of the abdomen (Fig. 286). %% are similar to Ph. fruhstorferi (Brunner v. Wattenwyl , 1907) but differ by: the plain brown body colouration; slightly shorter median segment; sparsely dentate posteroventral carina of the profemora; more numerous and distinct serrations of the anterodorsal carina of the profemora; less decidedly tapered semi-tergites of the anal segment; shorter cerci (Fig. 287) and shorter antennae which project no further than to the posterior margin of the mesonotum. Etymology: The specific name virgea is derived from the Latin “virga” (= twig) and refers to the slender body of %%. Description: && (Fig. 129): Very long (body length 190.0–234.0 mm; including subgenital plate 220.0– 248.0 mm) and moderately slender species (maximum width of thorax 8.0–9.5 mm), with lateral margins of abdominal tergites II–VI gently rounded and the subgenital plate very long and lanceolate. Colour variable; general colouration of body and legs usually yellowish, ochracheous or greyish mid to dark brown, but occasionally dull green specimens occur. Body surface smooth and slightly glabrous. Spination of legs dark red with black points. Bases of tarsi pale brown or grey. Eyes dark brown. Antennae greyish mid brown with ventral surface darker.



213

FIGURES 130-132: Phryganistria spp. 130. Phryganistria virgea (Westwood, 1848), % HT, N-India, Silhet (OXUM) 131. Phryganistria fruhstorferi (Brunner v. Wattenwyl , 1907), & LT, N-Myanmar, Mount Cariani (MCSN) 132. Phryganistria fruhstorferi (Brunner v. Wattenwyl , 1907), %, N-Myanmar, Tenasserim (coll. FH, No. 0210-1)

Head: Oval, broadest just behind the eyes, 1.5x longer than wide, vertex very slightly rounded and posterior margin gently elevated. A pair of oval impression between the eyes and a deep, transverse depression between the bases of antennae. Posterior margin of head capsule with a longitudinal impression on both sides of the coronal line, the latter one very faint. Eyes small, circular and moderately convex; their length contained about 3x in that of cheek. Antennae almost reaching posterior margin of mesonotum. Scapus dorsoventrally compressed, 1.5x longer than wide and slightly narrowed basally. Pedicellus cylindrical, slightly longer than wide and about 2/3 the length of scapus. Thorax: Pronotum shorter and narrower than the head, 1.3x longer than wide, posterior margin slightly broader than anterior margin. Anterior margin raised and followd by transverse furrow, posterior margin rounded. Median transverse depression distinct, slightly rounded but not reaching lateral margins of segment. Mesothorax slightly dorsoventrally compressed and oval in cross-section, about 2.8x longer than head and pronotum combined. Mesonotum very indistinctly, gradually widened towards the posterior. Metanotum parallel-sided and about 3.2x longer than wide. Pleurae and sterna smooth. Abdomen: Median segment about 1/3 the length of metanotum, slightly longer than wide and rectangular. Segments II–V increasing in length, II 2x, V about 2.5x longer than wide, VI as long as V. Tergite II rectangu-


HENNEMANN & CONLE

lar, III–VI with lateral margins ± distinctly deflexed and gently rounded. VII a little longer than II and parallel-sided except for a slight lobe posterolaterally, roughly 3x longer than wide and distinctly narrower than previous. Praeopercular organ formed by a pair of distinct, slender, back-curving spines at posterior margin of sternum VII. Remaining sternites smooth. Tergite VIII slightly narrower than VII, about 2.3x longer than wide, strongly convex and very gently constricted medially. IX less than half the length of VIII, quadrate. Anal segment longer than IX, with a very fine median carina and a wide triangular excavation posteriomedially; posterolateral angles tapered and roundly triangular. Supraanal plate very small, triangular and sharply keeled. Cerci small, cylindrical and tapered towards a pointed tip; finely bristled. Subgenital plate very long and lanceolate, gradually tapered towards a pointed apex and projecting over anal segment by more than the combined length of tergites VIII–X (Fig. 286). Legs: All long and moderately slender. profemora slightly longer than head, pro- and mesonotum combined, mesofemora longer than mesonotum, metafemora reaching about half way along abdominal segment V and metatibiae slightly projecting over apex of anal segment. Anterodorsal carina of profemora with 14–26 prominent and acute black serrations. Posteroventral carina armed with 8–18 rather more slender and distinctly pointed teeth. Posteroventral carina of protibiae sparingly and minutely serrate, other carinae unarmed. Two outer ventral carinae of meso- and metafemora set with 8–14 distinct but rather slender and pointed teeth, which decrease in size towards the apex of femur. Medioventral carina indistinct, unarmed. Dorsal carinae with a similar number of conspicuously smaller teeth; posterodorsal carina of mesofemora occasionally with an enlarged triangular, lobe-like tooth one quarter off the base. All carinae of meso- and metatibiae minutely dentate, although more sparsely on dorsal carinae. Posterodorsal carina of mesotibiae often with a slightly enlarged, triangular tooth about 1/3 off the base of tibia. Probasitarsus slightly longer than remaining tarsomeres combined, very slender and with all carinae unarmed. Meso- and metabasitarsi about as long as following three tarsomeres combined, dorsal carina slightly rounded apically and ventral carinae very minutely serrate. %% (Fig. 130): Medium-sized (body length 141.0–154.0 mm) and moderately slender for the genus (maximum body width 3.0 mm), body surface smooth and glabrous. General colouration of body and legs plain greyish, yellowish or reddish mid to dark brown and but occasionally dark green specimens may occur; mesoand metathorax slightly darker than rest of body. Cheeks sometimes with a faint dark postocular stripe. Tibiae often with a few faint, pale transverse bands; the basal one most distinct. Antennae dark brown. Head: Generally as in &&. Eyes very prominent, convex and projecting hemispherically; their length contained a little more than 2x in that of cheek. Antennae as in && but relatively longer, projecting over posterior margin of metanotum. Thorax: Pronotum slightly shorter and distinctly narrower than head, 1.5x longer than wide, otherwise as in &&. Mesothorax 3.7x longer than combined length of head and pronotum; mesonotum parallel-sided and very gently broadened at posterior margin. Metanotum about 3/5 the length of mesonotum, parallel-sided and almost 9x longer than wide. Posterior margin at each outer angle with a small, rounded hump (residues of the alae). Pleurae and sterna smooth. Abdomen: Median segment slightly more than 1/3 the length of metanotum, rectangular and about 1.6x longer than wide. Segments II–VII gently constricted medially, II–V slightly increasing in length, II 2.5x, V 4.5x longer than wide, VI as long as V. Tergite VII about ¾ the length of VI and roughly 2.5x longer than wide. VIII 2/3 the length of VII, strongly broadened towards posterior margin and 1.5x longer than wide. IX equal in length but considerably narrower than VIII, 2x longer than wide and slightly constricted medially. Anal segment a little longer than IX, laterally compressed and strongly tectiform. Semi-tergites elongate, slightly gradually tapered towards a rounded apex (Fig. 287). Interior surfaces apically swollen and densely covered with minute black teeth. Cerci small, cylindrical, tapered towards a pointed apex and finely bristled; staying clearly beyond apex of anal segment. Poculum moderately convex, cup-like, with a blunt hump at the angle and reaching about half way along tergite IX.



215

Legs: All very long and slender, armature generally as in && but less prominent. Profemora considerably longer than head, pro- and mesonotum combined, mesofemora about as long as combined length of pro- and mesonotum, metafemora reaching about half way along abdominal segment VI and metatibiae projecting considerably over apex of abdomen. Posteroventral carina of profemora just sparsely dentate. Probasitarsus very elongate and slender, longer than remaining tarsomeres combined. Meso- and metabasitarsi as long as or a little longer than remaining tarsomeres combined, dorsal carina not raised apically. Eggs (Fig. 190–191): Medium-sized, capsule oval in lateral aspect, with a flat impression near polar-area, slightly laterally compressed and oval in cross-section. Capsule with a slight keel on dorsal and ventral surfaces and polar-area, beginning and ending at operculum. Capsule surface minutely granulated (70x), not glossy. General colouration of capsule and micropylar plate dark brown, operculum black. Micropylar plate, elongate, covering slightly less than 2/3 the length of capsule, tapered towards anterior end and posteromedially with a distinct gap. Outer margin black. Micropylar cup placed in the posteromedial gap of plate, small and dark brown. Median line distinct and projecting over posterior margin of plate. Operculum oval and slightly concave, outer margin raised. Capitulum large, dark reddish brown, funnel-shaped and with several large lateroventral impressions. Stalk obvious, whitish. Measurements [mm]: Length including capitulum 5.2, length 4.5, width 3.1, height 4.0, length of micropylar plate 2.6. Comments: When Westwood (1848) described Bacteria sarmentosa from the & (Westwood, 1848: 65, pl. 32: 1) and Bacteria virgea from the % (Westwood, 1848: 66, pl. 32: 2) he already noticed they might just represent the opposite sexes of one and the same species and commented on Bacteria virgea: “The proportions and general appearance of this insect indicate that it is most probably the male of the proceeding. It is on this account that I have represented them both on one plate.”. Subsequently, Westwood (1859: 46) synonymised both species and listed Lonchodes virgeus (Westwood) as the valid species. Although sarmentosa has page priority Westwood’s synonymy as the first reviser is valid and in accordance to Article 24.2.2 of the Code (ICZN, 1999) on the precedence between simultaneously published names spelling or acts. Stål (1875: 63) Kirby (1904a: 358) and Giglio-Tos (1910: 16) have erroneously listed Phryganistria sarmentosa as the valid name. Distribution (Fig. 399): Northeastern India (Assam: Khasi Hills, Lait Pyng Kot; Liman; Samarang and Silhet). The records from “Siam” (= Thailand), Peninsular Malaysia (Pulau Penang) are doubtful and excluded from Fig. 399. Java as a locality for a & in NHMW and a % in MHNG is most certainly wrong, since all other confirmed records are from Northeastern India, thus from the asian continent. Number of specimens examined: 65 TABLE 34. Measurements [mm] of Phryganistria virgea (Westwood, 1848).

&&

%%

Body

190.0–234.0

141.0–154.0


220.0–248.0

-

Pronotum

7.0–8.5

4.0–5.0

Mesonotum

40.0–49.5

32.5–35.0

Metanotum

25.5–27.5

18.5–19.2

Median segment

6.5–8.2

6.4–6.9

Profemora

58.5–70.0

52.0–54.0

Mesofemora

47.0–51.5

39.0–41.0

Metafemora

53.0–59.5

45.5–48.0

Antennae

64.0–68.0

> 65.0


HENNEMANN & CONLE

6.5 Genus Tirachoidea Brunner v. Wattenwyl , 1893 stat. rev. Type species: Phibalosoma cantori Westwood, 1859: 74, pl. 37: 1, 1a–b & pl. 38: 1, 1a–b by subsequent designation of Kirby, 1904a: 359. Tirachoidea Brunner v. Wattenwyl, 1893: 83 (in part). Kirby, 1904a: 359 (in part—only species No’s 1–2). Karny, 1923: 240 (in part). Günther, 1953: 555. Qi & Liu, 1992: 250. Eucarcharus Brunner v. Wattenwyl, 1907: 186 (in part). (Type species: Lonchodes feruloides Westwood, 1859: 45, pl. 6: 5, 5a–b, by subsequent designation of Günther, 1935b: 138; see comments below) Otte & Brock, 2005: 136 (in part). Lobophasma Günther, 1935b: 138 (in part). Pharnacia, Redtenbacher, 1908: 449 (in part). Giglio-Tos, 1910: 43. (in part). Brock, 1995: 95 (in part). Brock, 1996: 27 (in part). Seow-Choen, 1998c: 184. Brock, 1999: 132 & 181 (in part). Seow-Choen, 2000: 35 (in part). Bragg, 2001: 398. Otte & Brock, 2005: 264 (in part). Phibalosoma, Westwood, 1859: 71 (in part). Wood-Mason, 1875: 216. Wood-Mason, 1877a: 161 (in part). Wood-Mason, 1877b: 347, pl. 3: 1.

Description (%, &): Moderately slender to very robust Pharnaciini (&& in particular) with strong sexual dimorphism. Head indistinctly longer than wide, vertex strongly convex, either rounded and almost spherical or conical. Usually armed with a pair of (sometimes several) ± distinct tubercles, blunt spines or humps; the right one being larger than the left. Antennae shorter than mesonotum (&&) or about as long as head and complete thorax combined (%%). Mesothorax of moderate length, 2.0–2.4x longer than head and pronotum combined, in && constricted anteriorly. Mesosternum of %% with a more or less distinct median carina. Alae of %% fully developed and at least reaching tergite IV. Tegmina ovate or scale-like, slightly conical and narrowed basally, at best 2x longer than wide. Abdomen including median segment considerably longer than combined length of head and complete thorax. Median segment equal in length or slightly longer than metanotum in &&, distinctly longer than metanotum in %%. Abdominal segments II and III of && often ± swollen. Tergite VII dilated into a rounded or truncate lobe posterolaterally; more distinct in &&. Praeopercular organ of && very indistinct and formed by two short carinae or ridges close to posterior margin of abdominal sternum VII. Tergite VIII of && at least as long as IX and X combined. Semi-tergites of anal segment of %% with apex ± conspicuously elongated and finger-like. Cerci obtuse, oval or cylindrical in cross-section; ± distinctly in-curving in %%. Subgenital plate of && strongly keeled, boat-shaped and at best slightly projecting over apex of anal segment; never lanceolate. Poculum of %% convex and with a ± acute conical hump or spine at the angle. Legs of moderate length, femora ± broadened. All carinae ± prominently spinose or serrate; occasionally with single enlarged teeth and lobes. Anterodorsal carina of profemora ± serrate. Mesofemora of && ± strongly broadened, especially towards the base. Medioventral carina of meso- and metafemora distinct and armed with a several prominent spines (&& in particular). Mesofemora distinctly longer than metanotum and median segment combined. Probasitarsus of && at best as long as remaining tarsomeres. Meso- and metabasitarsus about as long as following three tarsomeres combined. All carinae elevated, either smooth or dentate; the dorsal carina of the meso- and metabasitarsus ± strongly raised and rounded. Eggs (Figs. 157–166): Medium-sized to large. Capsule ± spherical or slightly longer than wide, circular



217

in cross-section and lacking any depressions or keels. Capsule surface almost smooth, strongly shiny and with a conspicuous and large dark lateroventral marking. Polar-area with a minute blunt spine or tubercle and a bold blackish spot. External micropylar plate covering more than 2/3 of capsule length and shaped like an inverted “Y”. Apices of posterior arms of plate parallel-sided and rounded. Operculum slightly convex, circular and with an irregularly shaped hat- or knob-like capitulum on a distinct stalk. Diagnosis (Table 2): Well characterized amongst Pharnaciini by the strongly convex and armed vertex and ± spherical, smooth and strongly shiny egg capsule (Fig. 415). Very closely related to Pharnacia Stål, 1877 but distinguished by: the strongly convex, either conically raised or almost spherical vertex and rounded dorsal carina of the basitarsi of both sexes which bears a pair of (or more) tubercles, blunt spines or rounded humps, curved cerci of %% and tergite VIII of && being at least as long as but usually longer than IX and X combined. Furthermore, the eggs differ by: the capsule being almost spherical or slightly elongate and circular in cross-section, lacking depressions or a dorsoventral keel; smooth and strongly shiny capsule surface; parallel-sided and rounded apices of posterior arms of the micropylar plate (broadened and truncate in Pharnacia); minute blunt spine and bold blackish spot of the polar-area and large lateroventral marking of the capsule. Comments: Brunner v. Wattenwyl (1893: 83) originally established Tirachoidea for Phibalosoma cantori Westwood, 1859, Phibalosoma hypharpax Westwood 1859, Phibalosoma tirachus Westwood, 1859 and Cyphocrana tamyris Westwood, 1859. The latter species is not closely related and has subsequently been selected as the type species of Diesbachia Redtenbacher, 1908, a genus of the subfamily Necrosciinae (Brock, 1995). Tirachoidea was first misinterpreted by Kirby (1904a: 359) who designated Phibalosoma cantori Westwood as the type species. Although Brunner v. Wattenwyl did not make any ranking among the species that he listed as belonging to Tirachoidea, the generic name obviously indicates that Brunner v. Wattenwyl originally based it on Phibalosoma tirachus Westwood. There are several other cases in which Brunner v. Wattenwyl based new generic names on already described specific names and the concerned species were then always included in the genus. Examples include Marmessoidea Brunner v. Wattenwyl, 1893 and Necroscia marmessus Westwood, 1859 or Sipyloidea Brunner v. Wattenwyl , 1893 and Necroscia sipylus Westwood, 1859. Apart from selecting a type species, Kirby (1904a: 359) included one further recently described species in Tirachoidea, Phibalosoma westwoodii Wood-Mason, 1875. Redtenbacher (1908: 449) synonymised Tirachoidea with Pharnacia Stål, 1877 but omitted Kirby’s catalogue (1904a), hence not being aware of the typedesignations established in that publication. Karny (1923: 240) re-established Tirachoidea and transferred the first four more robust species of Redtenbacher’s key to Pharnacia, which have a swollen, convex and bituberculate head, to Tirachoidea. Brock (1996: 25) confirmed Redtenbacher’s synonymy of Tirachoidea and Pharnacia and synonymised Tirachoidea yet again, mainly based on %% of both type species having fully developed alae. Brock (1996: 25) stated “Tirachoidea is a synonym of Pharnacia on the basis that T. cantori belongs to the same genus as Pharnacia ponderosa Stål, the type species of Pharnacia” and commented Karny (1923) to have not identified “the link between P. cantori Westwood and P. ponderosa Stål”. His key states Pharnacia ponderosa Stål to show the same strongly rounded or elevated head seen in Ph. cantori (Westwood) “Back of head rounded or elevated; often with two swollen tubercles. (Brock, 1996: 26)”, which is not the case; it is actually just gently rounded and entirely smooth in Ph. ponderosa Stål. Thus, the strongly convex or conical and bi-tuberculate head is not a typical character of Pharnacia Stål, but one of the key features that distinguish Tirachoidea from Pharnacia. Consequently, all those taxa placed in Pharnacia by Brock (1996) and which have a distinctly convex and bi-tuberculate head like the type species T. cantori (Westwood), actually belong in Tirachoidea Brunner v. Wattenwyl (→ 6.2). Careful examination of all concerned taxa and comparison with Pharnacia have, in addition to the shape of the head, revealed several further features of the insects and eggs that clearly distinguish Tirachoidea from Pharnacia (see Table 2 and differentiation above). Consequently, Karny’s treatment of the two genera is confirmed and Tirachoidea here re-established as a valid genus (stat. rev.).


HENNEMANN & CONLE

Otte & Brock (2005: 136) erroneously stated that Günther (1935b: 138) had subsequently designated Eucarcharus inversus Brunner v. Wattenwyl, 1907 as the type species of Eucarcharus Brunner v. Wattenwyl, 1907, which would place Eucarcharus as a junior synonym of Tirachoidea. In fact however, Günther (1935b: 138) designated Lonchodes feruloides Westwood, 1859 as the type species and moreover stated the two Bornean Pharnacia annulata Redtenbacher, 1908 and Pharnacia nigricornis Redtenbacher, 1908 to be congeneric. Due to E. feruloides (Westwood, 1859) has a rather indistinct medioventral carina midways on the ventral surface of the profemora, not split anal segment and produced vomer in %%, as well as a closed internal micropylar plate in the eggs, Eucarcharus Brunner v. Wattenwyl is here removed from the tribe Pharnaciini and transferred to Stephanacridini (→ 4.7.2). Distribution (Fig. 86): So far recorded from NE-India, Bangladesh, Myanmar, Thailand, Vietnam, South China, Peninsular Malaysia, and N-Borneo (Sarawak & Brunei) as well as the Greater and Lesser Sunda Islands (Sumatra, Java & Lombok). Species included: 1. Pharnacia biceps Redtenbacher, 1908: 451.

[Java, Sumatra, Lombok, Peninsular Malaysia]

= Pharnacia chiniensis Seow-Choen, 1998c: 184 syn. nov. 2. Phibalosoma cantori Westwood, 1859: 74, pl. 37: 1, 38: 1. 3. Tirachoidea herberti spec. nov. 4. Eucarcharus inversus Brunner v. Wattenwyl , 1907: 186, pl. 8: 1. = Pharnacia semilunaris Redtenbacher, 1908: 452. syn. nov. 5. Pharnacia jianfenglingensis Bi, 1994: 10. 6. Tirachoidea siamensis spec. nov. 7. Phibalosoma westwoodii Wood-Mason, 1875: 216.

[Peninsular Malaysia, Thailand & SMyanmar] [Northern Borneo: Sarawak & Brunei] [Java] [S-China (Hainan & Yunnan) & NVietnam] [Thailand & S-Vietnam] [NE-India & N-Myanmar]

Keys to the species of Tirachoidea Brunner v. Wattenwyl, 1893

&& 1. Vertex with a pair of tubercles, blunt spines or distinct rounded to conical humps; meso-femora without a sub-basal lobe on posteroventral carina........................................................................................................2 - Vertex strongly globose and almost spherical, only with a pair of very shallow swellings, (Fig. 302); mesofemora with a prominent, dentate sub-basal lobe on posteroventral carina (Fig. 325) Java ......inversa 2. Head longer than high, with a pair of tubercles, spines or humps................................................................3 - Head almost as high as long; vertex prominently conically raised and with two or more prominent conical humps or spines (Figs. 137, 299 & 300).............................................................................................. biceps 3. Mesothorax < 2.5x longer than head and pronotum combined; subgenital plate reaching or projecting over posterior margin of anal segment; not Borneo .............................................................................................4 - Mesothorax 2.6x longer than head and pronotum combined; subgenital plate short, not reaching posterior margin of anal segment; Borneo (Figs. 135 & 215) ........................................................ herberti spec. nov. 4. Robust insects; vertex rounded and with a pair of granules or spiniform tubercles; meso- and metasternum conspicuously darker than rest of body ........................................................................................................6 - Slender insects; vertex conically raised and with two low humps; meso- and metasternum coloured like rest of body ...................................................................................................................................................5



219

5. Colouration of head plain; abdominal tergite with a posteromedian tubercle; subgenital plate projecting over apex of anal segment; NE-India & N-Myanmar ..................................................................westwoodii - Posterior of head with two conspicuous dark transverse stripes (Fig. 296); abdominal tergite VII smooth; subgenital plate not considerably projecting over anal segment; N-Vietnam & S-China (Fig. 143) ............. ............................................................................................................................................ jianfenglingensis 6. Very broad species (body width 15.0–18.0 mm); mesothorax < 2x longer than head and pronotum combined; ventral surface of antennae black; armature of mid and hind legs green; abdominal tergites V–VI smooth (Fig. 133)................................................................................................................................ cantori - More slender species (body width 12.0–14.0 mm); mesothorax > 2x longer than head and pronotum combined; antennae entirely brown; armature of mid and hind legs orange or red; tergites V–VI with a posteromedian tubercle (Fig. 142)...........................................................................................siamensis spec. nov.

%%* 1. Robust insects; mesonotum at best 2.5x longer than head and pronotum combined ...................................3 - Slender insects, mesonotum 3x longer than head and pronotum combined.................................................2 2. Vertex gently convex (Fig. 303); semi-tergites of anal segment very short, broad and truncate apically (Fig. 211); NE-India & N-Myanmar.............................................................................................westwoodii - Vertex strongly conically raised and armed with a pair of minute spines; semi-tergites of anal segment elongate and straight (Fig. 217); Borneo (Fig. 136) ........................................................ herberti spec. nov. 3. Head distinctly longer than high; vertex convex and with a pair of small granules or spines; anterior margin of tegmina and alae white or yellowish ..................................................................................................4 - Head almost as high as long; vertex strongly conically raised and with two or more prominent conical humps or spines (Fig. 301); anterior margin of tegmina and alae green (Fig. 138) ............................ biceps 4. Robust insects; abdominal tergites III–V < 5x longer than wide; tegmina plain .........................................5 - Slender species; abdominal tergites III–V almost 6x longer than wide; tegmina with a conspicuous white diagonal stripe (Fig. 144).................................................................................................... jianfenglingensis 5. Mesonotum 1.7x longer than head and pronotum combined; apex of semi-tergites elevated or hook-like ventrally (Fig. 214); ventral surfaces of antennae black (Fig. 134).................................................... cantori - Mesonotum almost 2x longer than head and pronotum combined; apex of semi-tergites rounded (Fig. 220); antennae plain brown (Fig. 139) ..........................................................................siamensis spec. nov. * %% of T. inversus (Brunner v. Wattenwyl , 1907) are not known.

Tirachoidea biceps (Redtenbacher, 1908) stat. rev. (Figs. 137–138, 165–166, 221–223, 299–301, 323, 363, 400, 428) Pharnacia biceps Redtenbacher, 1908: 451. LT [by present designation], &: Coll. Br. v. W., Java, Dr. Dohrn; det. Br.v. W. Pharnacia biceps (NHMW, No. 861); PLT, 2 &&: Coll. Br. v. W., Java, Dr. Dohrn; det. Br. v. W. Pharnacia biceps (NHMW, No. 861); PLT, &: 99; 18.445; Coll. Br. v. W., Java, Dr. Dohrn; det. Br. v. W. Pharnacia biceps (NHMW, No. 861); PLT, %: Coll. B. v. W. Java, Malang, Staud., det. Br. v. W. Pharnacia biceps; 20.534 (NHMW, No. 861); PLT, %: Coll. Br. v. W., Java, Malang, Staud.; det. Br. v. W. Pharnacia biceps, Malang (NHMW, No. 861); PLT, %: Coll. Br. v. W., Soekabaia, Fruhstorfer; det. Br. v. W. Pharnacia biceps, 18.477 (NHMW, No. 861); PLT, 2 && (subadult): Coll. Br. v. W., Java, Dr. Dohrn; det. Redtenb. Pharnacia biceps, Pharnacia biceps Redt. (NHMW, No. 861); PLT, 2 nymphs (3rd instar): Coll. Br. v. W., Tengger-Geb. Java, Fruhstorfer; det. Br. v. W. Pharnacia biceps; 18.158 & 18.490.b (NHMW, No. 861); PLT, &: Java orient. Montes Tengger, 4000´ IX.–X.1890 H. Fruhstorfer 10. II. 1894 (ZMUH); PLT, &: Java orient. Montes Tengger 4000´, 1890 H. Fruhstorfer; 2, Java H. Fruhstorfer vend. 10. II. 1894; Tirachoidea biceps Redt. &, Jos. Redtenbacher determ. 1899; public. 1906–08, Bestimm.-Verz. Nr. 2 „T.westwoodi W.M.“ (ZMUH); PLT, 2 &&:Java, Montagnes le 27., M. Fruhstorfer (MHNG); PLT, %: Java, Soekaboemi, M. E. Walsh (MHNG); PLT, 1 %: Java, Or., Passeroean, 622/20 (MHNG, No. 69); PLT, 1 %, 1 &: Museum Paris,


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Java, Plaboan Ratoé, R. Oberthur 1898; alcohol (MNHN); PLT, &: Museum Paris, Java, Toussaint 21-62. (MNHN); PLT, &: 3013, Java, Nuhn; Tirachoidea biceps Redt.; Brunner det. (MNHU); PLT, &: Tengger-Geb., Ostjava, Fruhstorfer S.; Java, Phryganistria acanthopus de Haan, Pharnacia biceps Redt. (MNHU); PLT, & (penultimate instar nymph): Ostjava, Tenggergeb., 4000’, Fruhstorfer S.; 50, Tirachoidea biceps Redt. (MNHU); PLT, 1 &: Assam; 88. Tirachoidea biceps Redt., Type (SMNS); PLT, 2 %%, 1 &: Java, Fruhstorfer, Tirachoidea biceps Brunner det. (ISNB); PLT, %: Jawa ‘Java occident.’, Mons Gede, 4000 ft., 1898; ex. H. Fruhstorfer coll. (ZMAS); PLT, number of specimens unknown: Java (HNHM – destroyed by fire). Weidner, 1966: 231. Brock, 1996: 27. Brock, 1998a: 17. Zompro, 2002a: 182. Zompro & Brock, 2003: 7. Hennemann & Conle, 2003a: 5. Otte & Brock, 2005: 264. Zompro, 2005b: 255. Brock, 2006: 49. Tirachoidea biceps, Karny, 1923: 240. Pharnacia cantori, Redtenbacher, 1908: 480 (in part—only the % from Java in SMNS). Pharnacia chiniensis Seow-Choen, 1998c: 184. HT, &: Malaysia, Pahang, Tasek Chini, 1.I.97, F. Seow-Choen & I. Seow-En (ZRCS); PT, 1 %: Malaysia, Pahang, Tasek Chini, 20.V.97, F. Seow-Choen (FSC); PT, 1 %: Malaysia, 19.II.1995, F. Seow-Choen (ZRCS); PT, 1 %: Malaysia, Pahang, Tasek Chini, II.94, F. Seow-Choen & Tay, E.P. (FRIM); PT, 1 %: Malaysia, Pahang, Tasek Chini, 3.VI.97, F. Seow-Choen (UKM); PT, 1 %: Malaysia, Pahang, Tasek Chini, 9.VIII.97, F. Seow-Choen (UKM); PT, 1 %: Malaysia, Pahang, Tasek Chini, 20.V.97, F. Seow-Choen (UKM); PT, 1 % & 1 &: Malaysia, Pahang, Tasek Chini, 20.XI.97, F. Seow-Choen (FSC); PT, 1 &: Malaysia, Pahang, Tasek Chini, 20.XI.97, F. Seow-Choen (UKM); PT, 1 &: Malaysia, Pahang, Tasek Chini, 21.XI.97, F. SeowChoen (UKM); PT, 1 &: Malaysia, Pahang, Tasek Chini, 20.XI.97, F. Seow-Choen (FSC); PT, 1 &: Malaysia, Pahang, Tasek Chini, 20.XI.97, F. Seow-Choen. syn. nov. Seow-Choen, 2000: 38, pl. 95 (%, &, egg). [Erroneously named Pharnacia cantori (Westwood) on pl. 95] Otte & Brock, 2005: 265. Phasma (Cladoxerus) acanthopus, de Haan, 1842: 131. [Misidentification. Subsequently described as Pharnacia biceps by Redtenbacher, 1908: 451] Tirachoidea hypharpax, Vanschuytbroeck & Cools, 1981: 17. [Misidentification] Further material: [32 &&, 12 %%, 1 nymph]: JAVA: 1 &: Java (ZSMC); 1 %: Kenden-Geb., O-Java, A. Heyne Berlin Wilm., Tirachoidea biceps Redt. K. Günther det. (MNHU); 1 &: Phryganistria Stål, Pharnacia westwoodi W.-Mas. K. Günther det., Pharnacia biceps Redt. det. F. Hennemann & O. Conle, IV.1998 (MNHU); 1 %: Radjamandala, Mt. Ofanpanja 1200 m, 8. 1936 via. Reinbek, Eingangs-Nr. 1/1957, det. F. Hennemann 1.1998 (ZMUH); 2 %%, 6 &&, eggs: ex Zucht: F. Hennemann, 1999, urspr.: Java (coll. FH, No’s 0193-2 to 9 & E1); 3 %%, 6 &&, eggs: ex Zucht: F. Hennemann, Herkunft: E-Java, F1-Generation, 2008 (coll. FH, No’s 0193-10 to 18 & E2) 1 &: Java orient., Montes Tengger 4000 m, 1890, H. Fruhstorfer; ex. verz. Z.L.Groningen; det. P.E. Bragg (RMNH); 1 %: Java (RMNH); 1 &: Banjoewangi, Java, Kahic, Barve, Aug. 1902 (RMNH); 2 &&, 1 nymphs: Buitenzorg, "specimens of acanthopus de Haan" det P.E. Bragg (RMNH); 1 &: Java (ISNB); 1 & Java, Vulkan Gede; coll. A. Finot, Phryganistria hypharpax, Westw.; Ex coll. Finot (ISNB); 1 &: Java, Mt. C. Picket; det. Brunner v. W. (MHNG); 1 &: Museum Paris, Java, Plaboan Ratoé, R. Oberthur 1898, alcool (MNHN); 1 &: Monts-Tengger Java, 1890, envoi Fruhstorfer; Collection A. Finot, Phryganistria hypharpax, Westw. (MNHN, coll. Finot, Box. No. 296); 1 &: Montagnes de Java, 12.1890, envoi Fruhstorfer; Collection A. Finot, Phryganistria hypharpax, Westw. (MNHN, coll. Finot, Box. No. 296); 1 &: Pengalengan, Java, 5. 1893; coll. A. Finot, Phryganistria hypharpax, Westw. (MNHN, coll. Finot, Box. No. 296); 1 %: Soekaboemi (Java) E. Walsh (MNCN). SUMATRA: 1 &: P.O. Stolz, Solok (Sum), Juni 1914; Tirachoidea cantori K. Günther det. (RMNH); 1 &: Ponteboek, Bekui-Deli; Tirachoidea westwoodi K. Günther det. (RMNH); 1 &: G.J. Weymann, Telokbetong, Sumatra, 23.5.1931 (RMNH); 1 &: „Sapama“, Br. / Redtb. Det. 1903 (ETHZ). PENINSULAR MALAYSIA: 1 %: W-Malaysia, Pulau Penang, leg. M.K.P. Yeh X. 1993 (coll. FH, 0193-1); 1 %: Fed. Malay State, Kedah, Kuala Ketil, J. Cadman (BMNH); 1 %: Malay State, Bukit Kutu, 3300ft., A.R. Sanderson (BMNH). NO / ERRONEOUS DATA:



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2 &&: no data (ETHZ); 1 &: no data (BMNH); 1 &: Philippines (?), W.L. Distant, 1903-365 (BMNH).

Diagnosis: Closely related to the second Javanese species, T. inversa (Brunner v. Wattenwyl, 1907), and T. cantori (Westwood, 1859) from Peninsular Malaysia and South Myanmar. It is however easily distinguished from these two species and all other members of the genus by: the namely strongly conically raised, prominently bi-tuberculate or bi-spinose vertex (Figs. 299–301) and strongly raised, distinctly rounded dorsal carina of the basitarsi of both sexes; large lobes of the mid and hind-legs of &&; very prominent and acute spine of the poculum (Fig. 223) and bright apple green anterior margin of the tegmina and alae of %% as well as the comparatively small eggs (capsule length 4.5–4.9 mm). Etymology: Neuter. The specific name biceps refers to the two characteristic conical elevations of the vertex of this species. Description: The description of the colouration is combined from photos of live captive reared insects and preserved specimens. && (Fig. 137): Medium-sized to large (body length 152.0–218.0 mm), moderately slender species (maximum width 9.0–13.0 mm). Colouration of body and legs variable, ranging from almost uniformly pale yellowish or greyish brown to dark brown often with dark greenish mottling and speckles, more rarely dark green specimens are encountered. Occasionally specimens may have conspicuous bold white markings or diagonal white lines on the dorsal surface of the thorax and abdomen. Mesopleurae with a fine orange-brown longitudinal line. Armature of the legs reddish brown. Antennae brown with ventral surfaces of all segments except scapus and pedicellus glossy black. Eyes dark reddish brown. All gonapophyses bright red. Head (Figs. 299–300): About 1.7x longer than wide, oval, vertex strongly conically raised and armed with a pair or blunt tubercles or spines; the sinistral one considerably larger than the dextral one. Occasionally the two central tubercles or humps may be accompanied with a variable number of smaller, irregularly set tubercles. Just behind the bases of the antennae with a broad, transverse depression. Eyes rather small and circular, their length contained about 3x in that of cheek. Antennae consisting of about 30 segments and slightly projecting over posterior margin of mesonotum. Scapus dorsoventrally flattened, almost 3x longer than wide and very gently narrowing towards the base. Pedicellus cylindrical, ¼ the length of scapus. Third antennomere longer than pedicellus, IV as long as pedicellus, V–VIII increasing in length, following roughly of equal length. Thorax: Pronotum distinctly narrower and a little shorter than head, almost 2x longer than wide and gently widening towards the posterior. Median transverse depression distinct and curved, but short and not reaching lateral margins of segment. Mesothorax slightly constricted at anterior margin and very gently widened posteriorly, 2.3x longer than head and pronotum combined. Mesonotum occasionally with a pale longitudinal median line which is most decided in posterior portion of segment. Metanotum about 2.3x longer than wide, rectangular and with a very minute posteromedian tubercle. Abdomen: Median segment slightly shorter than metanotum, almost 2x longer than wide, rectangular. Tergites II–VI increasing in length, II 1.5x, V almost 2.3x longer than wide and VI al little shorter than V; all with a small posteromedian tubercle. Tergite VII ¾ the length of VI, lateral margins in posterior half elevated to form a rounded lobe, which bears a minute spine at the angle and laterally extends by as much as 1/3 the width of the segment. Praeopercular organ formed by two short, curved carinae at posterior margin of sternum VII (Fig. 363). Tergite VIII longer than VII and about as long as IX and X combined, distinctly narrower than previous, constricted medially and almost 3x longer than wide. IX half the length of VIII, longer than wide, parallel-sided. Anal segment a little longer than IX and with a fine median carina; posterior margin with a deep, triangular excavation and the outer angles rounded (Fig. 222). Supraanal plate very small, rounded and with a fine median carina. Gonapophyses reddish brown, elongate, up-curving and reaching about to apex of anal segment. Cerci oval in cross-section, slightly tapered towards apex and almost reaching posterior margin of anal segment. Subgenital plate strongly keeled and boat-shaped, apex rounded and slightly projecting over


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posterior margin of anal segment (Fig. 221). Legs: All moderately long and slender, strongly spinose and armed with several prominent lobes. Profemora slightly longer than pro- and mesonotum combined, mesofemora a little shorter than mesothorax, metafemora reaching about half way along abdominal tergite V and metatibiae reaching apex of abdomen. Anterodorsal carina of profemora raised and armed with 16–22 distinct, triangular serrations. Posteroventral carina with 10–14 pointed triangular teeth. Posterodorsal carina with 3–8 prominent triangular teeth; one of these occasionally strongly enlarged and lobe-like. Posterodorsal carina of protibiae unarmed and often with a large, foliaceous, ± triangular tooth just before middle. Anterodorsal carina sparingly dentate. Posteroventral carina densely but minutely serrate, anteroventral carina smooth. Meso- and metafemora very gently downcurving with the two outer ventral carinae densely and acutely dentate. Anterodorsal carina only with a very few triangular teeth of variable size. Posterodorsal carina sparingly dentate; in mesofemora usually with one distinct triangular tooth sub-basally and two further, ± enlarged, triangular lobes in apical half (Fig. 323). Medioventral carina with 5–7 pointed spines of variable size and increasing in length towards base of femur. Ventral carinae of meso- and metatibiae densely but sharply dentate, anterodorsal carina almost smooth. Posterodorsal carina sparsely dentate and with a ± prominent, broad triangular lobe just before the middle and a rounded, dentate lobe apically. All carinae of probasitarsus moderately elevated and smooth, about as long as remaining tarsomeres combined. Meso- and metabasitarsi with dorsal carina distinctly raised and rounded and all carinae minutely dentate; slightly longer than following three tarsomeres combined. %% (Fig. 138): Small to medium-sized (body length 116.0–145.0 mm) and slender for the genus with long alae (53.0–71.0 mm). General colouration of body, legs, tegmina and costal region of alae ranging from almost uniformly pale to dark greenish or greyish brown, often with numerous yellowish speckles in darker specimens. Anterior margin of tegmina and alae bright apple green. Anal region of alae transparent greyish brown with brown veins. Antennae brown and, except scapus and pedicellus, blackish brown ventrally. Eyes reddish brown. Head (Fig. 301): Generally as in &&, but eyes much more prominent, strongly convex and projecting hemispherically; their length only a little more than 2x in that of cheek. Sometimes the vertex may bear five minute blunt spine-like tubercles of variable sizes. Antennae projecting over posterior margin of tergite II, otherwise as in &; densely sethose. Median antennomeres strongly elongated. Thorax: Pronotum generally as in &&, 2x longer than wide, narrower but about as long as head. Mesothorax gently broadened at posterior margin and about 2.5x longer than head and pronotum combined. Metanotum slightly longer than wide, broader than mesonotum. Meso- and metasternum with a fine longitudinal median keel; less decided on metasternum. Tegmina oval, strongly narrowed towards base and with a prominent conical central hump. Alae reaching at least half way along abdominal tergite V. Abdomen: Median segment considerably longer than metanotum, narrowing towards posterior margin. Segments II–V slightly increasing in length, II 3.5x, V almost 5x longer than wide. VI slightly shorter than V. Tergites V–VII often with a minute posteromedian granule. VII shorter than VI, 3x longer than wide and with posterior angles gently elevated and rounded. VIII about as long as VII, strongly swollen and gradually widening towards the posterior. IX strongly convex, constricted in posterior half and slightly shorter than previous. Anal segment a little longer than IX, laterally compressed and strongly tectiform. Semi-tergites moderately elongate, very gently down-curving and gradually tapered with the apex rounded dorsally and acute ventrally (Fig. 223). Interior surfaces densely covered with minute dark reddish brown teeth. Cerci elongate, round in cross-section, tapered and slightly in-curving apically; almost reaching apex of anal segment. Poculum strongly convex, cup-like and with a very prominent, elongate and acute, backward pointing central spine (Fig. 223) Legs: All long and slender, with all carinae, except anteroventral carinae of profemora- and tibiae, minutely but densely dentate and finely bristled. Profemora longer than head, pro- and mesonotum combined, mesofemora about as long as pro- and mesonotum combined, metafemora reaching about half way along



223

abdominal tergite V and metatibiae reaching to apex of anal segment. Anterodorsal carina of profemora with about 20 distinct and acute, triangular serrations. Posteroventral carinae with a slightly smaller number of pointed teeth. Posterodorsal carina with 4–8 minute teeth. Medioventral carina of meso- and metafemora with 10–13 minute spines, increasing in size towards the base. Posterodorsal carina occasionally with a single conspicuously enlarged tooth some 2/3 off the base. Mesotibiae with a more or less prominent triangular tooth or lobe about half way along posterodorsal carina, and a rounded, dentate lobe at the apex. Probasitarsus with all carinae ± prominently elevated and smooth, longer than remaining tarsomeres combined. Meso- and metabasitarsi as long as remaining tarsomeres combined except claw, dorsal carina strongly rounded and all carinae minutely serrate. Variation: T. biceps shows considerable variation concerning to the size, shape and number of the tubercles or spines of the vertex, armature of the legs and tarsi and colouration. A % from Pulau Penang in the first author’s collection (coll. FH, No. 0193-1) is remarkable for its dark greyish brown colour, strongly developed leg armature, strongly raised dorsal carina of the basitarsi and very prominently raised vertex, bearing five distinct but irregularly set spiniform tubercles. Furthermore, the left mesofemur of this specimen has a prominent triangular lobe about 2/3 the way along the posterodorsal carina. Captive reared specimens from a culture stock imported to Europe in the 1990’s from Java are remarkable for their comparatively small size, which may be due to the rearing conditions and alternative foodplants offered. The six reared && from this stock in the first author’s collection (coll. FH, No’s 0193-2 to 7) measure body lengths of only 152.0–170.0 mm, while most other && examined exceed 180.0 mm. In contrast to &&, the two captive reared %% of the concerned stock in the same collection (coll. FH, No’s 0193-8 & 9) are not considerably shorter than other specimens, measuring 120.5 mm and 125.0 mm. A & from Pengalengan (Java) in the collection of A. Finot (MNHN) exhibits remarkable bold white diagonal lines on the dorsal surface of the body. Eggs (Figs. 165–166): Medium-sized, almost spherical, general colouration of capsule and micropylar plate pale creamish grey, operculum blackish brown. Lateroventrally with a conspicuous, bold olive brown marking and slightly darker grey areas around micropylar plate. Whole capsule surface shiny and very unevenly granulose (70x). Micropylar plate slightly more than 2/3 the length of capsule, shaped like a bold inverted “Y” and with a bold reddish brown band along outer margin. Apices of all micropylar extension rounded. Micropylar cup placed in posteromedial gap of plate, cup-like. Median line slightly raised, not reaching apices of posterior extensions of plate. Polar-area with an oval, dark brown spot and a minute, blunt granule in the centre. Operculum almost circular, slightly convex and with a slight central impression. Capitulum yellowish brown, very prominent and irregularly funnel-shaped. Stalk indistinct, blackish. Measurements [mm]: Length including capitulum 4.5–4.9, length 3.7–4.0, width 3.0–3.2, height 3.2–3.8, length of micropylar plate 2.4–3.0. Comments: Redtenbacher (1908: 451) originally described this characteristic species from a syntype series consisting of at least 27 specimens deposited in NHMW, MHNG, MNHN, ISNB, MNHU, ZMUH, ZMAS and HNHM all from Java, with the exception of a & from Assam (= Northeast India) in SMNS and a specimen stated to be from Tonkin (= Vietnam) in MNHN. The latter specimen in MNHN was not traced, or is represented by one of the three && from Java contained in that collection, with the locality mistaken by Redtenbacher. Although the locality (Bangladesh) of the SMNS specimen is questionable it is clearly conspecific with the remaining type-series. Both localities are here regarded as erroneous, due to there are no recent records from Vietnam or Bangladesh The specimen(s) that Redtenbacher listed from HNHM appear to have been destroyed by fire (Brock, 1988b: 17) and so the exact number of specimens remains unknown. A & in NHMW is here selected as the lectotype, as the NHMW specimens were the only ones that Redtenbacher had at hand himself with confirmation. Examination of the % from Java in SMNS, which Redtenbacher (1908: 480) recorded as Pharnacia cantori (Westwood, 1859), has shown this to be a typical % T. biceps. The % from Java, two && and & nymph from Buitenzorg (= Bogor, Java) in RMNH, which de Haan (1842: 131) erroneously identified as Phasma


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(Cladoxerus) acanthopus Burmeister, are also typical representatives of this characteristic species. The &, which de Haan (1842: 131) termed “Varietas feminea” from Tapos (Java) is not traced in RMNH. Pharnacia chiniensis Seow-Choen 1998 is undoubtedly a junior synonym of this characteristic but strongly variable species and the first published record from Peninsular Malaysia (syn. nov.). The synonymy is confirmed by a % from Pulau Penang (Peninsular Malaysia) in the first author’s collection (coll. FH, No. 0193-1). Although the type specimens of Ph. chiniensis were not examined, the rather detailed descriptions and nice illustrations provided by Seow-Choen show all the characteristic features of T. biceps, e.g. the conically elevated and distinctly bi-tuberculate head of both sexes, characteristic legs armature; strongly dorsally rounded basitarsi and green anterior margin of tegmina and alae of %%. T. biceps is widespread throughout great parts of Java and Sumatra and appears to be quite common in the Tasek Chini area in central Peninsular Malaysia. The record “Lombok” listed below concerns to a culturestock imported to Europe and reared in captivity for a few generations in the late 1990’s (see below). According to its wide range, this species shows considerable variation concerning to the size, shape of the tubercles or horns of the vertex, armature of the legs and colouration. TABLE 35. Measurements [mm] of Tirachoidea biceps (Redtenbacher, 1908) stat. rev.

&&

%%

(Java)* 152.0-218.0 9.2-11.5 7.0-9.2 28.5-45.5 12.5-19.0 9.3-14.0 40.0-60.5 27.5-42.5 38.5-53.0 46.8-72.5 29.2-42.0 42.5-63.0 38.5-68.5

(Java)* 116.0-145.0 5.1-6.3 4.0-4.8 19.8-29.5 7.6-9.1 11.5-13.5 9.1-11.0 53.0-71.5 36.5-49.5 27.5-31.5 35.0-42.8 42.0-53.0 27.3-36.5 38.7-54.0 58.0-86.0

Body Head Pronotum Mesonotum Metanotum Median segment Tegmina Alae Profemora Mesofemora Metafemora Protibiae Mesotibiae Metatibiae Antennae * including the type specimens ** according to Seow-Choen, 1998c: 185

&&, PT of. chiniensis** 170.0-205.0 13.0-16.0 8.0-9.0 36.0-44.0 15.0-18.0 42.0-46.0 29.0-38.0 41.0-48.0 47.0-55.0 31.0-35.0 50.0-54.0 40.0-60.0

%%, HT & PT of. chiniensis** 138.0 7.5-9.0 4.0 23.0 17.0 9.0-11.0 70.0-71.0 34.0-39.0 31.0-34.0 39.0-44.0 43.0-50.0 26.0-34.0 46.0-52.0 60.0-84.0

T. biceps is being successfully cultured in Europe since the late 1990’s from stock originating from an unknown locality in Java and was included on the Phasmid Study Group culture-list as culture No. 203. A second stock from E-Java is currently being imported and successfully reared in Europe. In captivity it readily accepts oak (Quercus spp., Fagaceae), bramble (Rubus spp., Rosaceae), rose (Rosa spp., Rosaceae) and Salal (Gaultheria shallon, Ericaceae) as alternative foodplants. Seow-Choen (2000: 38) stated Hopea nutans (Dipterocarpaceae) to be the natural food-plant at Tasek Chini (Peninsular Malaysia) and Mangifera indica (Anacardiaceae) and Psidium guajava (Myrtaceae) to be accepted as alternative foodplants in captivity in Singapore. In the early 1990’s stock from Lombok was imported to Europe, but the culture was not maintained for any longer than a few generations and died out soon. This was included on the Phasmid Study Group culture-list as culture No. 137 “Pharnacia sp.”. && of the first Javanese stock are remarkable for their very small size (see comments on variation above). This species is apparently easy to rear in very large and well ventilated cages with nymphs taking some 5 months to reach maturity. && produce an average of eight eggs per day. REVISION OF PHARNACIINI


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Distribution (Fig. 400): Java (Malang; Soekabaia; Tengger Mountains 4000 ft.; Kenden Mountains; Radjamandala, Mt. Ofanpanja 1200 m; Passeroean; Kahic, Barve; Bogor [= Buitenzorg]; Pengalengan; Nuhn & Mount Picket), Sumatra (Solok; Ponteboek, Bekui-Deli; Telokbetong & Sapama), Lombok and Peninsular Malaysia (Pulau Penang; Kedah: Kuala Ketil; Pahang: Tasek Chini & Bukit Kutu 3300 ft.). Northeast India [= Assam] (with doubt). Number of specimens examined: 73

Tirachoidea cantori (Westwood, 1859) stat. rev. (Figs. 133–134, 157–158, 212–214, 272, 401) Phibalosoma cantori Westwood, 1859: 74, pl. 37, fig. 1, 1a–b (%), pl. 38, fig. 1, 1a–b (&). LT, %: Cantor, Malacca; Type, Westwood. Phibalosoma cantori, Type Orth. 621, Phibalosoma cantori Westw. Hope Dept. Entom.; Tirachoidea cantori (Westwood) Lectotype % det. P.D. Brock, 1995 (OXUM, Nr. 621); PLT, &, Cantor, Malacca; Type, Westwood. Phibalosoma cantori, Type Orth. 621, Phibalosoma cantori Westw. Hope Dept. Entom.; Tirachoidea cantori (Westwood) & Paralectotype P. D. Brock, 1995 (OXUM, Nr. 621). Wood-Mason, 1877a: 161 Wood-Mason, 1877b: 347. Tirachoidea cantori, Brunner v. Wattenwyl , 1893: 83. Kirby, 1904a: 359. Karny, 1923: 240. Rehn, 1904: 70. Brock, 1995: 95. [Designation of LT] Tay & Seow-Choen, 1996: 186. Pharnacia cantori, Redtenbacher, 1908: 480 (in part—only &, → % from Java = Tirachoidea biceps Redtenbacher). Brock, 1996: 27. Brock, 1999: 132 (in part—only description of %, Figs. 88a–d (% and &)). Seow-Choen, 2000: 35, pl. 94 (%, &). Otte & Brock, 2005: 264. [Not: Pharnacia cantori ?, Bragg, 2001: 398. Bornean specimens are Tirachoidea herberti spec. nov., see below ] Further material: [12 &&, 26 %%, eggs]: PENINSULAR MALAYSIA: 3 &&: W-Malaysia, Perak, Tapah Hills, ca. 600 m, via M.K.P. Yeh VIII. 1994 (coll. FH 0038-1 bis 3); 22 eggs: ex Zucht F. Hennemann 1993, urspr.: W-Malaysia (coll. FH 0038-ED); 1 %: Cameron Highlands, 2000ft., 19.6.1972, C.C.Chua; BM 1975-604 (BMNH); 2 &&: Perak, Rothschild, Bequest, BM 1939-1 (BMNH); 2 &&: Malaisie, Perak, Cameron Highl., 1200–1600 m, leg. Pfanner (MHNG); 1 &: Zoo Frankfurt, Zucht 18.12.84 (SMFM); 1 %: Malaysia, Cameron Highlands, leg. A. E. Selner, Augsburg 41990 (ZSMC); 1 %, 2 &&, eggs: Captive reared by M. Herbert, culture stock from West Malaysia (coll. MH). S-THAILAND: 1 %: Peninsular Siam, Nakon Sri Tamorat, Khao Ram 1200–3000ft., 3. 1922 H. M. Pendlebury; Nr. 717 (RMNH). S-MYANMAR: 2 %%: S-Myanmar, Mergui District, Tenasserim, 20.III.1994, leg. Lehmann (coll. FH, 0038-4 & 5); 6 %%: S-Myanmar, Mergui District, Tenasserim, VI.1996, leg. Lehmann (coll. FH, 0038-6 to 11); 14 %%: S-Myanmar, Tenasserim, leg. Lehmann VI.1996 (coll. OC); 1 &: Tenasserim, Mekane, 90 km, E. of Móulmein, 200m, 2.-8.XI.1934 (RMNH). NO DATA: 1 &: without data (BMNH).

Diagnosis: Closely related to T. siamensis spec. nov. but distinguished by: the relatively shorter mesothorax; less globose vertex and black ventral surface of the antennae of both sexes; broader body (maximum width 15.0–18.0 mm); deeper and more triangular posteromedian excavation of the anal segment (Fig. 213); smaller


HENNEMANN & CONLE

FIGURES 133-136: Tirachoidea spp 133. Tirachoidea cantori (Westwood, 1859), & West Malaysia, Perak, Tapah Hills (coll. FH, No. 0038-2) 134. Tirachoidea cantori (Westwood, 1859), % S-Myanmar, Mergui District, Tenasserim (coll. FH, No. 0038-6) 135. Tirachoidea herberti spec. nov., & HT, Brunei, Badas (BMNH) 136. Tirachoidea herberti spec. nov., % PT, Brunei, Badas (coll. MH)



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FIGURES 137-141: Tirachoidea spp. 137. Tirachoidea biceps (Redtenbacher, 1908), & Java, captive reared (coll. FH, No. 0193-6) 138. Tirachoidea biceps (Redtenbacher, 1908), % Java, captive reared (coll. FH, No. 0193-8) 139. Tirachoidea siamensis spec. nov., % PT, N-Thailand, Dansai Loei (coll. FH, No. 0142-2) 140. Tirachoidea westwoodii (Wood-Mason, 1875), % N-Myanmar, Highland of Tenasserim (coll. FH, No. 0211-1) 141. Tirachoidea inversa (Brunner v. Wattenwyl , 1907), & Java (HT of Ph. semilunaris Redtenbacher, NHMW)

and roundly triangular posterolateral lobes of abdominal tergite VII (Fig. 272); less distinct praeopercular organ; smooth abdominal tergites V–VII and dull green armature of the legs of &&; as well as the ventro-apically elevated or slightly hook-like semi-tergites of the anal segment (Fig. 214); shorter meso- and metabasitarsi and more regular and dense serrations of the legs of %%. The eggs are easily distinguished from those of


HENNEMANN & CONLE

T. siamensis spec. nov. by the creamish brown, almost spherical egg capsule (Figs. 157–158). In addition to these morphological features, T. cantori is distributed more southward than siamensis spec. nov. which is restricted to the northern and central portions of Thailand. Also related to T. biceps (Redtenbacher, 1908) but at once distinguished by lacking the prominent conical elevation of the vertex, relatively shorter mesothorax and more robust body and legs both sexes. && furthermore differ by the relatively shorter mesothorax, broader and rounded posterolateral angles of the anal segment and more prominent ventral armature of the meso- and metafemora, %% by the broader semi-tergites of the anal segment, less acute central spine of the poculum and white or pale yellow instead of apple green anterior margin of the alae. The eggs are considerably larger than those of T. biceps and have the micropylar plate relatively longer and more slender. Etymology: Dedicated to Dr. Theodore Cantor, member of the East India Company, who first collected this species in 1840. Description: && (Fig. 133): Large to very large (body length 190.0–246.5 mm), very robust species (maximum body width 15.0–18.0 mm). All legs of moderate length, robust and heavily spinose. General colouration of body either dull green with numerous darker speckles or greyish, greenish or reddish brown with irregular darker and paler speckles, often with washed whitish areas on abdomen. Back of head with four ± decided dark brown longitudinal lines and a shallow dark postocular line. Abdominal tergites with 1–3 irregular pairs of slightly darker patches in posterior portion. Ventral surface of abdomen with ± distinct white mottling. Mesonotum with a longitudinal row or small dark brown speckles on a whitish background along lateral margins. Meso- and metasternum dark green or brown and set with a few small, oval yellow spots. Legs dull green, brown or slightly aubergine. All carinae of mid and hind legs and entire spination green with black points (may be yellowish green on tibiae). Antennae brown with ventral surfaces of all antennomeres except scapus and pedicellus black. Eyes reddish brown. Head: About 1.8x longer than wide, oval, vertex roundedly convex and with a pair of ± distinct tubercles or small, blunt spines (the sinistral one smaller). Between the bases of antennae with a deep, transverse impression and between the eyes a larger, slightly raised and smooth region. Eyes rather large, higher than long with the anterior margin triangularly elevated; their length contained length contained about 3.5x in that of cheek. Antennae reaching to posterior margin of mesonotum, segments first distinctly increasing then decreasing in length towards apices of antennae; consisting of about 37 antennomeres. Scapus dorsoventrally flattened, 2.7x longer than wide and parallel-sided, ventrally with a longitudinal median furrow. Pedicellus about 1/3 the length of scapus, cylindrical and indistinctly longer than wide. III longer than pedicellus, IV strongly shortened. Thorax: Pronotum distinctly narrower and shorter than head, trapezoidal, and widening towards posterior margin. Anterior margin slightly raised and with a distinct pit at each outer lateral angle. Median transverse depression gently curved and just not reaching lateral margins of segment. Mesothorax about 1.8x longer than head and pronotum combined, slightly constricted anteriorly and conspicuously broadened at posterior margin. Mesosternum smooth except for a very indistinct longitudinal carina along lateral margins and two parallel longitudinal median ridges which terminate shortly before the apex. Metanotum distinctly shorter than mesonotum, about 1.7x longer than wide and slightly constricted medially. Metasternum smooth. Tegmina and alae present as very small (ca. 1.0 mm), sclerotised scale-like structures. Abdomen: Median segment a little shorter than metanotum, slightly trapezoidal with anterior margin rounded and about 1.4x longer than wide. Tergite II about as long as median segment, as long as wide and slightly broadening towards the posterior. III–VI slightly increasing in length, III indistinctly longer than wide, VI about 2.2x longer than wide. VII slightly shorter than VI, lateral margins gradually dilated and forming a roundly triangular lobe posteriorly, which projects by about 1/3 the width of the segment (Fig. 272). Sternites II–VI smooth. Praeopercular organ formed by a pair of very shallow and low, converging ridges close to posterior margin of sternum VII. Tergite VIII about as long as VI and longer than XI and X combined,



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distinctly narrower than previous, gently constricted medially, strongly convex and about 3x longer than wide. IX strongly convex, indistinctly longer than wide and about 1/3 the length of VIII. Anal segment longer than IX, with a distinct median carina and a broad and deep, more or less triangular median excavation, outer angles forming a narrow, rounded lobe (Fig. 213). Supraanal plate short, rounded and with a distinct longitudinal keel. Cerci short, oval in cross-section, gently up-curving and tapered at the apex. Gonapophyses elongate, up-curving and slightly projecting over apex of anal segment. Subgenital plate elongate, strongly keeled, boat-shaped and slightly projecting over anal segment (Fig. 212). Legs: All of moderate length, very robust and heavily spinose, basal half of meso- and metafemora distinctly swollen with the two outer ventral carinae gently elevated. Profemora about as long as pro- and mesonotum combined, mesofemora about as long as mesothorax, metafemora reaching about one third the way along abdominal segment V and hind tarsi just not reaching apex of abdomen. Anterodorsal carina of profemora armed with 15–18 prominent and pointed serrations. Posterodorsal carina with 6–7 acute triangular teeth. Posteroventral carina armed with 9–12 pointed spines. Anterodorsal carinae of protibiae with low but regular, wide-spaced serrations, posteroventral carina with ± 20 rather small and low serrations. The posterodorsal carina occasionally with an triangular, tooth-like lobe just before the middle. Dorsal carinae of mesoand metafemora sparingly dentate, each usually with a ± enlarged triangular tooth or lobe 1/3 off the base. Two outer ventral carinae with 10–15 very prominent, long and pointed spines which decrease in size towards the apex. Medioventral carina becoming increasingly acute but rounded towards the base and armed with 6–8 very prominent, slightly hooked spines; the three basal ones largest. Dorsal carinae of meso- and metatibiae sparingly but acutely dentate. Posterodorsal carina occasionally with a ± prominent triangular lobe just before the middle and forming a rounded, acutely dentate lobe at the apex. Ventral carinae densely armed with prominent, slender and pointed spines, those of the median carina largest. Probasitarsus with all carinae slightly elevated and about as long as remaining segments combined. Meso- and metabasitarsi with all carinae bearing 2–4 small teeth and the dorsal carina gently rounded; about as long as following three tarsomeres combined. %% (Fig. 134): Of moderate size (body length 119.0–152.5 mm) and rather robust for the genus with very long alae (77.5–83.0 mm). General colouration of body and legs yellowish, greenish or greyish dark brown, often with a few small white spots on meso- and metanotum. Mesonotum usually with a slight greyish wash. Mesopleurae with a fine longitudinal orange line along upper margin. Meso- and metasternum dark brown, abdominal sternites I–VIII with ± distinct white mottling and a sepia brown transverse streak posteriorly. Abdominal tergite usually with an irregular white lateral marking. Tegmina and costal region of alae either plain yellowish, greenish or greyish brown, anterior margin of tegmina and basal quarter of anterior margin of alae with a bold, longitudinal whitish line. Anal region of alae transparent greyish brown with sepia brown veins. Outer ventral carinae of meso- and metafemora dull green, spination of legs dark orange with black points. Cheeks with a shallow dark brown postocular streak. Eyes reddish brown. Antennae mid to dark brown dorsally and except for scapus and pedicellus black ventrally. Head: Generally as in &&, but eyes much more prominent and projecting hemispherically; their length contained no more than 2.5x in that of cheek. Antennae projecting over posterior margin of tergite II, otherwise as in &&. Thorax: Pronotum distinctly narrower and shorter than head, gently widened towards the posterior, otherwise as in && but relatively longer; 1.6x longer than wide. Mesothorax about 2x longer than head and pronotum combined and slightly broadened at posterior margin. Mesonotum parallel-sided and with a very faint longitudinal median line. Mesosternum with a distinct longitudinal median carina which terminates slightly before posterior margin. Metasternum just with a very faint longitudinal median line. Metanotum slightly longer than wide, considerably broader than mesonotum. Tegmina oval, narrowed towards the base and with a prominent and rather acute, conical central hump; reaching to posterior margin of metanotum. Alae varying in length, reaching at least half way along abdominal tergite VI and at best half way along tergite VII. Abdomen: Median segment almost 3x longer than wide and 2x longer than metanotum, slightly gradually


HENNEMANN & CONLE

narrowing towards the posterior. Segments II–IV increasing in length,V–VII decreasing, IV longest and VII shortest segment. II 3.5x, IV more than 5x and VII only about 2.5x longer than wide, parallel-sided. Tergite VII slightly constricted medially and with a ± prominent, rounded posterolateral lobe. VIII slightly longer than VII, strongly swollen and broadening towards the posterior; almost 2x the width of II–VII. IX a little longer than VIII, almost 3x longer than wide and decidedly constricted medially. Anal segment shorter than IX, laterally compressed and strongly tectiform. Semi-tergites rather broad, slightly tapered with the apex rounded and ± decidedly truncate or hook-like ventrally (Fig. 214). Interior surfaces densely covered with minute black teeth in the apical portion. Cerci small but obtuse, sub-cylindrical and slightly in-curving. Poculum strongly convex, cup-like, posteriorly marginated and with a small, ± acute backward pointing central spine (Fig. 214). Legs: All of moderate length and rather robust. Profemora slightly longer than head, pro- and mesonotum combined, mesofemora about as long as pro- and mesonotum combined, metafemora slightly projecting over posterior margin of abdominal tergite IV and hind legs slightly projecting over tip of abdomen. Armature similar as in &&, but considerably less prominent but with the number of spines larger. Spines rather uniform in size on each carina. The meso- and metafemora in particular are much less heavily spinose and not considerably broadened basally. Posterodorsal carina of profemora only with 4–6 very minute teeth. The posterodorsal and posteroventral carina of the mesofemora may bear a few slightly enlarged teeth in the basal portion. Occasionally there may be abroad triangular pre-median lobe on the posterodorsal carina of the mesotibiae. Probasitarsus as long as remaining tarsomeres combined except claw, all carinae slightly elevated. Meso- and metabasitarsi with all carinae minutely dentate and the dorsal carina decidedly rounded; slightly longer than following three tarsomeres combined. Eggs (Figs. 157–158): Large, capsule almost spherical. General colouration of capsule creamish pale brown, the lateral and ventral surfaces slightly darker. Micropylar plate pale greyish brown, operculum black, capitulum dull orange. Lateroventrally with a faint, slightly darker marking and slightly darker areas around micropylar plate. Capsule surface strongly shiny and very unevenly granulose (70x). Micropylar plate shaped like an inverted “Y” with outer margin dark brown and covering more than ¾ of capsule length. Apices of all extensions rounded. Micropylar cup placed in posteromedial gap of plate; small, cup-like and with a median carina. Median line slightly raised and reaching to apices of posterior extensions of plate. Polar-area with a circular dark brown spot and a very minute, blunt granule in the centre. Operculum slightly oval, convex and impressed in centre. Capitulum distinct, conical and on an obvious black stalk. Measurements [mm]: Length including capitulum 5.0–5.2, length 4.6–4.8, width 4.1–4.2, height 4.0–4.1, length of micropylar plate 3.8–3.9. Comments: Westwood (1859: 74) originally described Phibalosoma cantori from a % and & in OXUM collected in Peninsular Malaysia and provided nice figures of both sexes (Westwood, 1859: plate 37 (%), 38 (&)). Brock (1995: 95) designated the % in OXUM as the lectotype. Brunner v. Wattenwyl (1893: 83) included Westwood´s taxon and three further species in his new genus Tirachoidea for which subsequently Kirby (1904: 359) designated Phibalosoma cantori Westwood as the type species. Redtenbacher (1908) re-transferred these three species to Pharnacia Stål, 1877 and erroneously synonymised the two genera (→ see: comments on Pharnacia Stål and Tirachoidea Brunner v. Wattenwyl). Examination of Westwood’s type specimens in OXUM clearly showed the & PLT to exhibit a pair of granules on the vertex, which are not indicated in the illustration neither mentioned in the original description by Westwood, a fact which has caused confusion about the status of T. cantori (Westwood) and the genus Tirachoidea Brunner v. Wattenwyl . This error of Westwood has however already been pointed out by WoodMason (1877a: 347) who stated “I have examined with Professor Westwood the typical specimens of P. Cantori preserved in the Hopeian collection at Oxford and find that the insects have been legitimately united by their describer; the female having small equal blunt representatives of the conspicous cephalic tubercles seen in the male; these tubercles not having been represented by Professor Westwood in his figure of the former.”.



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In his book on Malaysian Phasmatodea Brock (1999: 132, fig. 88) reproduced the illustrations of both sexes of Tirachoidea cantori (Westwood) of Westwood (1859, pl. 37: 1, 38: 1) . The photographs of the & (Brock, 1999, pl. 26) and egg (pl. 24) stated to be this species however show Pharnacia sumatranus (Brunner v. Wattenwyl ). Brock (1999: 133) stated there to be a “second type of egg” which he briefly characterized as “[…] glossy whitish brown with a black operculum,…”. These are in fact the egg of Tirachoidea cantori (Westwood). Seow-Choen (2000: 35) provided illustrations and characterizations for both species as well as a list of features which distinguish the insects and eggs. Seow-Choen (2000: 36) cited the following bodylengths for T. cantori: %% 130.0–145.0 mm, && 190.0–230.0 mm. Examination of the % from Java which Redtenbacher (1908: 480) recorded from SMNS has shown this to be a typical specimen of Tirachoidea biceps (Redtenbacher, 1908). Therefore, “Java” can be regarded an erroneous record of T. cantori. The specimens which Bragg (2001: 398) listed from Borneo are not conspecific with T. cantori (Westwood) but all represent Tirachoidea herberti spec. nov., which is the only known representative of Tirachoidea Brunner v. Wattenwyl in Borneo (see comments on this species). TABLE 36. Measurements [mm] of Tirachoidea cantori (Westwood, 1859) stat. rev.

&, PLT

%, LT

&&

%%

(OXUM)

(OXUM)

Body

200.0

137.0

204.0–246.5 (190.0)*

119.0–152.5

Head

13.0

6.8

14.3–16.0

5.7–7.1

Pronotum

8.5

4.7

9.5–11.0

3.7–5.3

Mesonotum

34.5

22.5

39.5–44.0

20.0–24.3

Metanotum

15.0

9.5

15.5–18.0

7.0–10.5

Median segment

10.5

13.5

11.0–12.0

10.5–14.5

Tegmina

-

-

11.0–12.0

10.9–14.0

Alae

-

79.0

-

61.0–83.0

Profemora

43.0

36.0

51.5–57.5

34.0–39.0

Mesofemora

32.8

31.5

41.0–44.5

29.5–31.5

Metafemora

39.5

41.0

51.5–54.3

37.5–41.6

Protibiae

47.5

43.0

54.0–65.0

37.5–48.5

Mesotibiae

35.5

29.0

38.0–43.8

27.3–34.0

Metatibiae

44.0

43.5

51.5–62.0

35.0–43.6

Antennae

> 33.0

> 6.0

50.5–64.0

61.0–77.0

* according to Seow-Choen (2000: 36)

T. cantori used to be successfully cultured in Europe during the 1980’s from stock originating in the Tapah Hills (Peninsular Malaysia). It was included on the Phasmid Study Group species list as culture No. 58 “Pharnacia sp.”. In captivity in Europe the insects readily accepted bramble (Rubus spp., Rosaceae) and oak (Quercus spp., Fagaceae) as alternative foodplants. Seow-Choen (2000: 36) listed Mangifera indica (Anacardiaceaee), Psidium guajava (Myrtaceae), Rubus fruticosus and Rubus moluccanus (Rosaceae) to be natural foodplants of T. cantori in Peninsular Malaysia. Distribution (Fig. 401): Peninsular Malaysia (Perak: Tapah Hills & Cameron Highlands), South Thailand Peninsular (Nakon Sri Tamorat, Khao Ram 1200–3000ft.) and South Myanmar (Mergui District: Tenasserim & Mekane, 90 km east of Móulmein, 200 m). Number of specimens examined: 40


HENNEMANN & CONLE

Tirachoidea herberti spec. nov. (Figs. 135–136, 163–164, 215–217, 304, 366, 402) Pharnacia cantori ?, Bragg 2001: 398, distribution map 23C. HT, &: Brunei, Badas, swamp forest, leg. M. Herbert I.1993 (BMNH, ex coll. MH, No. 11-2). PT, %: Brunei, Badas, swamp forest, leg. M. Herbert I.1993 (coll. MH, No. 11-1). PT, %: Brunei, Badas swamp forest, M. Herbert, III.1994 (coll. PEB, No. 2407). PT, 2 eggs: Brunei, Badas, swamp forest, leg. M. Herbert I. 1993 (coll. FH, No. 0358-E). PT, %: Sabah, Telupid S’Kam, Tawai F. Res, Ento Staff, 1-13 Sept, 94 (FRCS). Further material [not examined]: SARAWAK: 1 %: Sarawak, Sadong, Cranston (SMSM-272); 1 %: Sarawak, Melinau, near Mt. Mulu, XI.1910 (SMSM-275); 1 Sarawak, Mount Meranja Kecil, 23.V.1914 (SMSM-279); 1 &: Sarawak, Kuching, 08.V.1900 (SMSM-280). NO DATA: 1 %: no data (SMSM-276).

&:

Diagnosis: This is the only representative of the genus known to occur in Borneo. It is closely related to T. cantori (Westwood, 1859) and T. biceps (Redtenbacher, 1908) but readily distinguished from both by the raised and distinctly serrate anterodorsal carina of the profemora of both sexes. && furthermore differ from those of both species by: the relatively smaller and more elongate head (Fig. 304); considerably more elongate and slender body (maximum width 10.5 mm); much broader and rounded lateral lobes of tergite VII which extend by more than half of the body width; short subgenital plate which stays beyond the apex of anal segment (Fig. 215) and much less prominent leg armature. %% are readily distinguished from those of the two mentioned species by the relatively longer mesothorax and more elongate and slender, straight semi-tergites of the anal segment (Fig. 217). Eggs differ by the considerably more slender and relatively longer micropylar plate and smaller, funnel-like capitulum (Fig. 163). Concerning to the shape of the vertex it represents an intermediate between T. cantori and. T. biceps. Etymology: This impressive new species is dedicated to Mel Herbert (Hütschenhausen, Germany) for providing his two specimens and eggs for examination and donating two eggs to the first author. Description: && (Fig. 135): Very long (body length 229.0 mm) and slender for the genus (maximum body width 9.5 mm). General colouration of body irregularly dark greyish brown with numerous lighter grey or whitish shadings and white speckles; last five abdominal tergites blackish dorsally. Head with a very faint greenish postocular line. Mesonotum furnished with a longitudinal band of irregular black and white speckles along lateral margins. Abdominal tergite III with a bold white lateral marking. Meso- and metasternum plain dark greyish brown and set with several small white spots. Great parts of abdominal sternites II-III whitish with brown speckles and markings. Lateral surfaces of subgenital plate white in anterior half, posterior portion dark brown with white speckles. Legs pale greyish with all carinae green, femora with a very slight pink wash. All femora and tibiae with a bold white transverse sub-apical band. Entire spination of legs dark red with black points. Basitarsi blackish brown with white bases. Antennae greyish mid brown with bases of all antennomeres darker. Eyes creamish brown. Head (Fig. 304): Almost parallel-sided, 2.3x longer than wide, vertex strongly conically raised and armed with a pair of small but acute spines, the dextral more decided than the sinistral. Between the bases of the antennae with a small, oval impression. Eyes prominent, circular and strongly convex; their length contained about 2.5x in that of cheek. Antennae reaching about ¾ the way along mesonotum, consisting of about 40 antennomeres. Scapus dorsoventrally flattened, 3x longer than wide and roughly parallel-sided. Pedicellus less than half the length of scapus, oval in cross-section and slightly longer than wide. Following antennomeres cylindrical, III slightly longer than pedicellus, IV distinctly shortened, V–X increasing in length. Fol-



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lowing antennomeres very elongate but distinctly decreasing in length towards apex of antennae. All finely bristled. Thorax: Pronotum slightly shorter and narrower than head, about 1.6x longer than wide and gently widening towards the posterior. Median transverse depression rather distinct, straight and covering about half the width of segment. Anterior margin raised and followd by a deep transverse furrow. Mesothorax 2.6x longer than head and pronotum combined, slightly constricted anteriorly and very slightly broadened posteriorly. Metanotum 2.2x longer than wide, gently constricted medially and with a very minute scale-like tubercle posteromedially. Meso- and metasternum smooth. Abdomen: Median segment slightly more than ¾ the length of metanotum, conspicuously tapering towards a rounded anterior margin. Segment II shorter than median segment. II–VI increasing in length, II 1.3x, VI almost 2.3x longer than wide. Tergites V and VI with a distinct, rounded and densely bristled posteromedian tubercle; a minute posteromedian tubercle present on VII. Tergite VII ¾ the length of VI with lateral margins dilated into a broad, rounded lobe which extends by more than half the width of segment. Sternum VII with praeopercular organ formed by a pair of ridge-like, converging median humps on a common base near to the posterior margin (Fig. 366). Tergite VIII distinctly longer than VII or XI and X combined, considerably narrower than previous, constricted medially and about 4x longer than wide. IX less than half the length of VIII, 1.5x longer than wide, constricted medially and broadened posteriorly. Anal segment a little longer than IX, with a fine longitudinal median carina and a broad, roundly truncate posteromedian excavation, outer angles conspicuously elevated and forming a rounded, slightly scoop-shaped lobe (Fig. 216). Supraanal plate small, rounded and with a median keel. Cerci short, almost cylindrical in cross-section and tapered towards a narrow apex; finely bristled. Gonapophyses yellowish brown and slightly projecting over posterior margin of subgenital plate. Subgenital plate boat-shaped, narrow in the anterior half, strongly convex and keeled in the posterior portion, apex staying posterior margin of anal segment (Fig. 215). Legs: All long and slender, meso- and metafemora very slightly down-curving. Profemora about as long as pro- and mesonotum combined, mesofemora slightly shorter than mesothorax, metafemora reaching about half way along abdominal segment V and metatibiae almost reaching tip of abdomen. Anterodorsal carina of profemora with 19–20 prominent and pointed triangular serrations. Posterodorsal carina set with 8–9 minute teeth. Posteroventral carina with 13–16 acutely pointed teeth of moderate size. Anterodorsal carina of protibiae gently raised and roughly serrate. Posterodorsal carina with 8–11 minute teeth. Posteroventral carina minutely but densely dentate. All carinae of meso- and metafemora sparingly armed with small, pointed teeth; medioventral carina with 6–7 moderately prominent spines which increase in size towards the base of femur. Ventral carinae of meso- and metatibiae densely dentate, dorsal carinae with a smaller number of minute teeth. Posterodorsal carina with a flat and broad, roughly triangular lobe about 1/3 off the base and forming a rounded, dentate lobe at the apex. Probasitarsus with all carinae strongly dilated and gently rounded, posteroventral carina a with a single minute tooth, other carinae smooth; about as long as following tarsomeres combined except claw. Dorsal carina of meso- and metabasitarsus rounded, all carinae minutely dentate; as long as following three tarsomeres combined. %% (Fig. 136): Moderately sized to large (body length 139.0–165.0 mm) and slender for the genus with long alae (70.0–77.0 mm). Body, legs tegmina and costal region of alae creamish to reddish mid brown, mesonotum with a slight greenish wash and a few irregularly set small, white spots. Anterior margin of tegmina broadly white and costal region of alae with a white longitudinal stripe along anterior margin, which terminates about half way. Anal region of alae transparent brown and becoming slightly darker towards the apex, veins brown. Meso- and metasternum dark brown and furnished with several white spots, the latter with a conspicuous white marking close to posterior margin. Abdominal sternum III with distinct white markings, the remaining sternites with whitish speckles and mottling. Meso- and metafemora with an indistinct pale, transverse sub-apical band. Leg armature dark red with black points. Eyes dark reddish brown. Antennae plain yellowish brown to greyish mid brown.


HENNEMANN & CONLE

Head: Generally as in && but relatively larger, more globose and distinctly broader than pronotum. Eyes prominent, circular and projecting hemispherically; their length contained more than 2.5x in that of cheek. Antennae strongly sethose and projecting over posterior margin of tergite III, otherwise as in &&. Thorax: Pronotum distinctly narrower and shorter than head, generally as in &&. Mesothorax 3x longer than head and pronotum combined, parallel-sided and very slightly widened at posterior margin. Mesosternum with a fine longitudinal median carina, metasternum smooth. Tegmina strongly constricted basally and broadly rounded apically, central hump ± prominent and roundly conical. Alae reaching half way along abdominal tergite VI. Abdomen: Median segment almost 2x longer than metanotum, tapering towards posterior margin. Segments II–V slightly increasing in length, II 4x and V about 5.5x longer than wide, VI slightly shorter than V. VII 2/3 the length of VI and gently widening towards the posterior. VIII strongly swollen and widened towards the posterior, trapezoidal in dorsal aspect and about 2x longer than wide. IX conspicuously constricted medially, 2x longer than wide and slightly longer than VIII. Anal segment strongly laterally compressed and tectiform. Semi-tergites very elongate, straight and finger-like with the apex narrow and rounded (Fig. 217). Interior surfaces of semi-tergites covered with numerous small hooked spines apically. Cerci obtuse, oval in cross-section, tapered towards a narrow apex and very slightly in-curving. Poculum strongly convex and cup-like with a blunt central projection, reaching about 2/3 the way along tergite XI (Fig. 217). Legs: All very long and slender. Profemora about as long as head, pro- and mesonotum combined, mesofemora as long as pro- and mesonotum combined, metafemora reaching about half way along abdominal tergite V and metatibiae reaching to apex of abdomen. Anterodorsal carina of profemora with about 20 sharply triangular serrations. Posterodorsal carina set with 5–6 minute teeth. Posteroventral carina with about 20 small but pointed spines. Anterodorsal carina of protibiae slightly raised and with 30–40 distinct serrations; posteroventral carina with a similar number of very minute teeth. Posterodorsal carina set with about 10 small teeth. All carinae of meso- and metafemora minutely dentate; teeth of the two outer ventral carinae becoming increasingly prominent towards base of femur. Two outer ventral carinae with about 20–25, dorsal carinae with 10–15 considerably smaller teeth. Medioventral carina armed with 6–8 minute spines which increasing in size towards base of femur. All carinae of meso- and metatibiae minutely dentate. Posterodorsal carina of meso- and metatibiae forming a rounded, dentate lobe apically. Probasitarsus with all carinae smooth and slightly elevated, as long as remaining tarsomeres combined. Dorsal carina of meso- and metabasitarsi slightly rounded, all carinae minutely dentate; as long as remaining segment combined except claw. Variation: The % PT from Tawai Forest Reserve (Sabah) in FRCS is remarkable for the large size (body length 165.0 mm), comparatively flat central hump of the tegmina and distinctly enlarged sub-basal teeth of the two outer ventral carinae of the mesofemora. Eggs (Figs. 163–164): Several eggs were laid by the & HT. The following description is based on two eggs (PT) in the first author’s collection (coll. FH, No. 0358-E). Large, almost spherical, general colouration of capsule and micropylar plate creamish pale brown, operculum dark brown. Lateroventrally with a distinct, large reddish brown marking and areas of similar colour around posterior extensions of micropylar plate. Lateral surfaces with several small dark brown spots. Whole capsule surface glossy and very unevenly granulose (70x). Micropylar plate almost as long as capsule, shaped like an elongate inverted Y. Apices of all extensions rounded, outer margin reddish brown. Micropylar cup placed in a posteromedial gap of plate, indistinct and posteriorly bordered by a pale W-shaped line. Median line slightly raised, almost reaching to polar-area. Polar-area with a circular blackish spot and a minute, blunt spine in the centre. Operculum almost circular, slightly convex and impressed centrally. Capitulum brown, very small, funnel-shaped and with numerous small radial furrows dorsally. Stalk very short, dark brown Measurements of PT [mm]: Length including capitulum 5.1, length 4.8, width 4.7, height 4.7, length of micropylar plate 4.2. Comments: This is the only species of the genus to be recorded from Borneo. Bragg (2001: 398) provided



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a description of the % but was in doubt about the identity of the Bornean specimens in SMSM and coll. PEB, provisionally allocating them to T. cantori (Westwood). The five specimens in SMSM listed above as “Further material” have not been examined. They do however most certainly all represent T. herberti spec. nov.. Distribution (Fig. 402): Northern Borneo, endemic. Sarawak (Sadong [Bragg, 2001: 398]; Kuching [Bragg, 2001: 398]; Mount Meranja Kecil [Bragg, 2001: 398] & Melinau near Mount Mulu [Bragg, 2001: 398]), Brunei (Badas, swamp forest) and Sabah (Tawai Forest Reserve, Telupid S’Kam). Number of specimens examined: 4 TABLE 37. Measurements [mm] of Tirachoidea herberti spec. nov.

&, HT (BMNH)

%, PT (coll. MH)

%, PT

Body

229.0

139.0

145.0

Head

12.6

5.6

6.0

Pronotum

9.0

4.3

5.0

Mesonotum

48.3

25.8

27.0

Metanotum

18.8

7.5

8.0

Median segment

14.5

13.5

14.0

Tegmina

14.5

11.3

10.5

Alae

-

72.0

70.0

Profemora

57.0

36.0

38.0

Mesofemora

42.0

30.0

31.0

Metafemora

50.5

38.0

39.0

Protibiae

66.8

46.0

48.0

Mesotibiae

46.0

32.0

33.0

Metatibiae

63.3

45.0

47.0

Antennae

58.0

80.0

> 27.0

(coll. PEB)

Tirachoidea inversa (Brunner v. Wattenwyl , 1907) comb. nov. (Figs. 141, 210, 302, 325, 404) Eucarcharus inversus Brunner v. Wattenwyl , 1907: 186, pl. 8: 1. HT, &: Museum Paris, Java, Ouest (Tougou), J. D. Pasteur 3-96; 6. Carcharus inversus Br. Type ! (MNHN). Otte & Brock, 2005: 136. Lobophasma inversus, Günther, 1935b: 138. Pharnacia semilunaris Redtenbacher, 1908: 452. HT, &: Baron Warsberg, Java, 1868; Mus. Caes. Vind., Java, Baron Warmsberg; det. Redtenb. Pharnacia semilunaris; Pharnacia semilunaris Redt., Phasm. Inv. Nr. 822 (NHMW, No. 859). syn. nov. Phobaeticus semilunaris, Brock, 1996: 29. Brock, 1998a: 56. Otte & Brock, 2005: 269.

Further material [1 &]: JAVA: 1 &: Java, Sukabami 2000 m, leg. Fruhstorfer 1893; [4] (ZMUH).


HENNEMANN & CONLE

Diagnosis: Closely related to the second Javanese species T. biceps (Redtenbacher, 1908) but at once distinguished from this and all other members of the genus by the strikingly rounded, almost spherical vertex which only has two very shallow swellings towards the frons (Fig. 302) and prominent sub-basal lobe of the posteroventral carina of the mesofemora (Fig. 325). It furthermore differs from T. biceps by: the more slender and elongate body and legs; larger and slightly truncate lateral lobes of abdominal tergite VII; distinct rounded posteromedian tubercle of tergites I–VI and lack of prominent dorsal lobes of the mesofemora and protibiae. Etymology: “Inversus” (lat. = inverse, contrary). Description: && (Fig. 141): Of moderate size (body length 180.6–200.0 mm) and rather slender for the genus (maximum body width 7.0–8.0 mm). General colouration of body and legs yellowish mid to greyish dark brown with the entire body to a various degree furnished with pale greyish and yellowish speckles. Legs with broad but irregular whitish to pale grey transverse bands. Median segment with several distinct white spots in posterior half. Eyes dark reddish brown. Antennae brown and becoming darker towards the apex. Head (Fig. 302): Strongly globose, 1.5x longer than wide, vertex very prominently convex and almost spherical, towards the frons with a pair of very shallow, rounded swellings. Between the bases of the antennae with a distinct oval impression. Eyes of moderate size, convex and with the anterior margin slightly truncate; length contained about 2.5x in that of cheek. Antennae at least reaching half way along mesonotum (broken in all examined specimens). Scapus dorsoventrally flattened, 2.5x longer than wide and gently narrowing towards the base. Pedicellus cylindrical, about 1/3 the length of length of scapus. III 2.5x longer than scapus, IV slightly shorter than scapus, V–XIII increasing in length. Following antennomeres strongly elongated. All finely bristled. Thorax: Pronotum about ¾ the length and distinctly narrower than head, about 1.3x longer than wide and gently widened towards the posterior. Median transverse depression moderately distinct, slightly curved and almost reaching lateral margins of segment. Mesothorax about 2.3x longer than head and pronotum combined, mesonotum almost parallel-sided. Metanotum about 2/5 the length of mesonotum, 2.3x longer than wide and parallel-sided, posteromedially with a small granule. Meso- and metasternum smooth. Abdomen: Median segment about ¾ the length of metanotum, parallel sided and with a transverse posteromedian tubercle. Segment II distinctly shorter than median segment and following segments, slightly longer than wide. Segments III–V increasing, VI–VII decreasing in length, IV longest and about 3x longer than wide. Tergites II–VI each with a distinct, roundly transverse tubercle posteromedially. Tergite VII shorter than previous, posterior half laterally dilated into a very prominent but slender apically gently truncate lobe, which projects by 2/3 of the body width. Praeopercular organ indistinct and formed by two very minute ridges at posterior margin of sternum VII. Tergite VIII slightly shorter and distinctly narrower than previous, a little longer than IX and X combined, strongly convex, 3x longer than wide and gently constricted medially. IX strongly convex, about half the length of VIII and 1.5x longer than wide. Anal segment as long as IX with a fine median carina and a broad, concave excavation posteromedially; posterolateral angles slightly elevated and broadly rounded, slightly scoop-shaped lobe (Fig. 210). Supraanal plate very small, tapered towards the apex and keeled. Cerci sub-cylindrical in cross-section, tapered towards a pointed tip and finely bristled. Gonapophyses elongate and up-curving, reaching to posterior margin of anal segment. Subgenital plate rather elongate, boat-shaped and strongly keeled in posterior portion; apex slightly projecting over apex of abdomen (Fig. 210).



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Figures 142-144: Tirachoidea spp. 142. Tirachoidea siamensis spec. nov, & PT, N-Thailand, Lemgao Petchabun (coll. FH, No. 0142-7) 143. Tirachoidea jianfenglingensis (Bi, 1994), & N-Vietnam, Cuc Phuong, captive reared (coll. FH, No. 0405-2) 144. Tirachoidea jianfenglingensis (Bi, 1994), % N-Vietnam, Cuc Phuong, captive reared (coll. FH, No. 0405-28)


HENNEMANN & CONLE

Legs: All long and very slender for the genus, meso- and metafemora slightly thickened near base. Profemora about as long has pro- and mesonotum combined, mesofemora slightly shorter than mesothorax, metafemora reaching about half way along abdominal segment V and hind legs reaching tip of abdomen. Anterodorsal carina of profemora armed with 16–19 prominent, triangular and slightly foliaceous serrations. Posteroventral carina with a similar number of small teeth. Posterodorsal carina set with 12–14 minute triangular teeth. Anterodorsal carina of protibiae slightly raised and with low but broad serrations, posteroventral carina minutely dentate. All carinae of mid and hind legs quite regularly set with small, pointed teeth. Posterodorsal carina with two slightly enlarged teeth sub-basally and with a ± prominent, triangular lobe-like tooth at the apex. Posteroventral carina sub-basally furnished with a prominent, rounded and tri-dentate lobe (Fig. 325). Metafemora without any enlarged teeth. Posterodorsal carina of meso- and metatibiae apically protruded into a rounded, tri-dentate lobe. Posteroventral carina slightly dilated and forming a triangular tooth apically. Probasitarsus with all carinae moderately elevated and smooth; as long as remaining tarsomeres except claw combined. Meso- and metabasitarsi with a slightly raised dorsal carina and all carinae minutely dentate; about as long as following three tarsomeres combined. Comments: Redtenbacher (1908: 452) described Pharnacia semilunaris from single & in NHMW not aware that it was identical with the species that Brunner v. Wattenwyl (1907: 186, pl. 8: 1) had already described and illustrated as Eucarcharus inversus based on a & in MNHN. Examination of both type specimens has clearly shown them to be the same species. Consequently, Ph. semilunaris Redtenbacher falls as a junior synonym of E. inversus Brunner v. Wattenwyl, 1907 (syn. nov.). The strikingly rounded, almost spherical but unarmed vertex, long tergite VIII, rounded dorsal carina of the meso- and metabasitarsi and armature of legs clearly place this species in the genus Tirachoidea Brunner v. Wattenwyl , 1893 (comb. nov.). TABLE 38. Measurements [mm] of Tirachoidea inversa (Brunner v. Wattenwyl, 1907) comb. nov. &, HT (MNHN)

&, HT of semilunaris (NHMW)

Body

180.6

200.0

Head

8.0

9.0

Pronotum

6.4

7.9

Mesonotum

39.4

40.0

Metanotum

16.2

16.8

Median segment

11.3

12.6

Profemora

45.3

49.0

Mesofemora

35.3

38.5

Metafemora

43.3

49.0

Protibiae

54.0

55.0

Mesotibiae

37.6

38.6

Metatibiae

44.7

50.0

Antennae

34.2

> 16.0

The HT of E. inversus Brunner v. Wattenwyl in MNHN is remarkably slender, which is due to the specimen was a newly adult & not in egg-production when captured. Hence, the body has strongly shrunken during a provisional preservation in ethanol and subsequent dehydration. The measurements given by Brunner v. Wattenwyl (1907: 186) match almost perfectly with the actual measurements taken, except for the body length which Brunner v. Wattenwyl stated to be 157 mm. This is most certainly a printing mistake and should read



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177 mm. The actual body length taken is 180.6 mm (→ Table 38). The prominent lateral lobes of tergite VII made Günther (1935b: 138) erroneously transfer E .inversus Brunner v. Wattenwyl to his newly described genus Lobophasma Günther, 1935 (Type species: Eucarcharus rex Günther, 1928) which is here shown to be a synonym of Phobaeticus Brunner v. Wattenwyl , 1907 (syn. nov.). % and egg unknown. Distribution (Fig. 404): Java (Tugu & Sukabami 2000 m). Number of specimens examined: 3

Tirachoidea jianfenglingensis (Bi, 1994) (Figs. 8–10, 23, 28, 48–49, 143–144, 159–160, 224–226, 296–297, 356, 365, 405, 424) Pharnacia jianfenglingensis Bi, 1994: 10. HT, &: Hainan Province, Ledong County, Mt. Jianfengling, 13.X.1992, leg. Liu Zuyao, Wang Tianqi & Yin Haisheng (SIES). Chen, Xu et al., 1994: 189. Chen, 1999a: 49, figs. Otte & Brock, 2005: 265. Chen & He, 2007: 349, figs. 316 a-c. Tirachoidea jianfenglingensis, Chen et al., 2002: 108. Hennemann et al., 2008: 40. Pharnacia maxima, Redtenbacher, 1908: 450 (in part - only % from Tonking [= N-Vietnam] in NHMW). Tirachoidea westwoodi, Qi & Liu, 1992: 250. [Description of % from Yunnan Province, China. Misidentification] Further material: [29 &&, 29 %%, eggs] N-VIETNAM: 1 %: Tonkin, Than Moi, 6./7.1899, leg. Fruhstorfer, Coll. Br. v. W., No. 24.321 (NHMW, No. 858); 1 %, 1 &, 1 egg: ex Zucht, F. Hennemann, 1998–2001, S-Vietnam, Cuc Phuong, Nho Quam District, Ninh Binh Province (NHMW, ex coll. FH);, 1 %: Tonkin, Than Moi, 6./7., H. Fruhstorfer (ETHZ); 1 &: ex Zucht P. Heusi, IV. 1998, N-Vietnam, Cuc Phuong, Nho Quam District, Ninh Binh Province (coll. FH, No. 0405-1); 15 %%, 17 &&, 300 eggs: ex Zucht, F. Hennemann, 1998–2001, N-Vietnam, Cuc Phuong, Nho Quam District, Ninh Binh Province (coll. FH, No’s 0405-2 to 33 & ED); 1 %, 1 &, 1 egg: ex Zucht, F. Hennemann, 1998–2001, N-Vietnam, Cuc Phuong, Nho Quam District, Ninh Binh Province (SMFM, ex coll. FH); 1 %: Museum Paris, Cho-Moi, Tonkin, m. Roget 111-96, alcool, determined as T. biceps Redt (MNHN).; 2 %%: Museum Paris, Tonkin, Reg. De Hoa-Binh, A. De Cooman 1930 (MNHN); 1 &: Museum Paris, Tonkin, Hoa-Binh, A. Idoeman 1926 (MNHN); 1 &: Hoa-Binh, Tonkin, A. De Cooman, 37e envoi (MNHN); 1 &: Tonkin, Region de Hoa-Binh, Museum Paris, 1934 A. De Cooman (MNHN); 1 %, 1 &, 2 eggs: ex Zucht, F. Hennemann, 1998–2001, S-Vietnam, Cuc Phuong, Nho Quam District, Ninh Binh Province (ZSMC, ex coll. FH); 1 %, 1 &, 2 eggs: ex Zucht, F. Hennemann, 1998–2001, N-Vietnam, Cuc Phuong, Nho Quam District, Ninh Binh Province (coll. OZ, ex coll. FH); 4 %%, 4 &&, 10 eggs: ex Zucht O. Conle VI. 2000, N-Vietnam, Cuc Phuong, Nho Quam District, Ninh Binh Province (coll. OC). N-THAILAND: 1 %: Doi Chieng Dao, 1.7–21.´33, N Siam, Alt 4000 ft., de Schauensee Sinkler Exped. (ANSP).

Diagnosis: Closely related to T. siamensis spec. nov. and T. westwoodii (Wood-Mason, 1875) but well characterized and distinguished from other members of the genus by the conspicuous dark transverse stripes in the posterior portion of the vertex of && and distinct diagonal white stripe on the tegmina of %%. The eggs distinguish T. jianfenglingensis from all other members of the genus by the elongate capsule which is conspicuously longer than high (Figs. 159–160). From T. siamensis spec. nov. it is distinguished by: the more slender and elongate body; relatively longer basitarsi; less prominent spination of the legs and more conically raised vertex of both sexes, which bears two rounded humps (Figs. 296–297), as well as the less deeply excavated posterior margin of the anal segment of


HENNEMANN & CONLE

&& (Fig. 225). %% clearly differ by: the typical white diagonal stripe of the tegmina; more prominent black serrations on the anterodorsal carina of the profemora; shorter probasitarsus and shorter, broader semi-tergites of the anal segment (Fig. 226). Furthermore, the meso- and metasternum of && are not considerably darker coloured than the remaining body. From T. westwoodii (Wood-Mason, 1875) && differ by the more slender body and relatively more elongate body segments; lack of a posteromedian tubercle on abdominal tergite VII; smaller lateral lobes of tergite VII and shorter subgenital plate (Fig. 224). %% differ by: the more robust body; relatively shorter mesothorax, abdominal segments and legs; more prominently swollen abdominal tergite VIII; less distinctly in-curving cerci; less truncate, apically constricted semi-tergites of the anal segment (Fig. 226); smaller number of triangular serrations on the anterodorsal carina of the profemora; less numerous but distinct teeth of the mid and hind legs and more prominently rounded dorsal carina of the meso- and metabasitarsi. In addition to these morphological features T. jianfenglingensis is well separated by its geographical pattern, being distributed more eastward than T. westwoodi and restricted to Northern Vietnam, South China and NE-Thailand (Fig. 405). Etymology: The name refers to the type-locality of this magnificent species, Mount Jianfengling in the Ledong County of Hainan Province, S-China. Description: && (Figs. 143, 415): Large to very large (body length 200.0–280.0 mm) and rather slender for the genus (maximum body width 10.0–13.0 mm). General colouration of body and legs yellowish or greyish pale to mid brown, occasionally dark greenish Entire body surface furnished with numerous paler and darker speckles and legs with irregular and ± distinct transverse whitish bands. Back of head with two irregular dark brown transverse stripes. Pronotum in posterior half with a dark brown, roundly M-shaped marking. Metanotum with a ± distinct C-shaped posterolateral marking. Border between meso- and metasternum with a broad white and beige transverse band. Meso- and metasternum coloured like rest of body. Abdominal sternites with distinct irregular white speckles and markings, tergite II almost entirely white. Eyes yellowish brown. Antennae brown with all antennomeres except scapus and pedicellus dark brown ventrally. Leg armature brown with black points. Head (Fig. 296): Oval, 1.5x longer than wide with the vertex strongly conically swollen and armed a pair of rounded humps; the sinistral one smaller than the dextral. A fine, slightly curved transverse impression behind the bases of the antennae and an oval, smooth and slightly raised area between the eyes. Eyes large and moderately convex; their length contained almost 3.5x in that of cheek. Antennae almost reaching posterior margin of mesonotum and consisting of 34–35 antennomeres; these first increasing then decreasing in length towards apex of antenna. Scapus dorsoventrally flattened, 2.5x longer than wide and strongly constricted basally. Pedicellus cylindrical and less than 1/3 the length of scapus. III longer than pedicellus, IV strongly shortened, V–XII increasing in length. Following antennomeres elongate. All finely bristled. Thorax: Pronotum slightly shorter and narrower than head and about 1.5x longer than wide, posterior margin broader than anterior margin. Anterior margin distinctly raised and followd by a rather deep transverse furrow. Median transverse depression slightly curved and very short, not reaching lateral margins of segment. Mesothorax about 2x longer than head and pronotum combined, very slightly constricted at anterior margin and gently widened at the posterior. Mesonotum with a fine, pale brown longitudinal median line and roughly parallel-sided. Mesosternum with a very indistinct longitudinal median carina. Metanotum a little less than 2/ 5 the length of mesonotum, about 2.2x longer than wide, rectangular and with a minute posteromedian tubercle; very gently constricted medially. Metasternum smooth. Abdomen: Median segment about 2/3 the length of metanotum and gently constricted medially. Segments II–V increasing in length, II about 1.5x, V 2.5x longer than wide, VI as long as III. Tergites IV–VI with a small, transverse posteromedian tubercle, most decided on V. Tergite VII about ¾ the length of VI, posterolaterally dilated into a broad rounded lobe, which extends by as much as ¾ the width of segment. Praeopercular organ rather distinct and formed by two converging carinae near posterior margin of sternum VII (Fig. 365).



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Tergite VIII very slightly longer than VII and longer than IX and X combined, distinctly narrower than previous, constricted medially and almost 4x longer than wide. IX 1.5x longer than wide, parallel-sided. Anal segment as long as IX, with a faint longitudinal median carina and a broad, roundly triangular excavation posteromedially; posterolateral angles broadly rounded (Fig. 225). Supraanal plate broad, rounded and with a fine median carina. Cerci small, oval in cross-section tapered towards a pointed apex and at best very slightly projecting over posterior margin of anal segment. Gonapophyses dark reddish brown, elongate, up-curving and ± distinctly projecting over anal segment. Subgenital plate strongly keeled and boat-shaped, posterior margin rounded and reaching to or slightly projecting over apex of anal segment (Fig. 224). Legs: All long and moderately robust with all carinae of mid and hind legs acutely spinose, meso- and metafemora very slightly down-curving. Profemora slightly longer than mesonotum, mesofemora about as long as mesothorax, metafemora roughly reaching to posterior margin of abdominal tergite IV and hind tarsi reaching posterior of tergite IX. Anterodorsal carina of profemora armed with 13–17 large, triangular serrations; posterodorsal carina set with 8–11 small teeth. Posteroventral carina with 16–20 small, pointed teeth; anteroventral carina unarmed. Anterodorsal carina of protibiae sparsely but rather regularly serrate, posteroventral carina densely armed with minute spines. All carinae of meso- and metafemora armed with pointed spines, which increase in size towards the base of femur; less numerous and slightly smaller on dorsal carinae. Ventral carinae of mesofemora very gently widened basally. Posterodorsal carina occasionally with 1–2 slightly enlarged triangular teeth about ¼ off the base. Medioventral carina of meso- and metafemora armed with 5–7 strong and rather long, gently hooked spines which increase in size towards base of femur. All carinae of meso- and metatibiae densely armed with minute, pointed spines; more sparsely on dorsal carinae. Posterodorsal carina usually with a ± prominent triangular lobe pre-medially and forming a small, rounded and dentate lobe apically. Probasitarsus with all carinae smooth but distinctly elevated, as long as remaining segments combined except claw. Dorsal carina of meso- and metabasitarsi distinctly raised and rounded apically, all carinae minutely dentate; slightly longer than following three tarsomeres combined. %% (Fig. 144): Large to very large (body length 139.5–183.0 mm) and slender for the genus with long alae (74.5–90.0 mm) and characteristic white markings on tegmina. General colouration of body, legs, tegmina and costal region of alae yellowish or greyish mid brown; the abdomen furnished with irregular, small pale and dark brown speckles. Occasionally a few white spots are present on mesonotum. Back of head with two ± distinct dark brown transverse stripes, sometimes interrupted medially. Abdominal tergite IX with a bold white marking anterolaterally. Pro, meso- and metasternum each with a distinct white marking posteriorly. Abdominal sternites II–V set with a few whitish spots and markings and the posterior margin broadly white. Anterior margin of tegmina broadly white and basal portion of anterior margin of alae with a white longitudinal band. Tegmina with a broad white diagonal band. Anal region of alae transparent greyish brown with brown veins. Spination of legs black, ventral surfaces ± distinctly marbled with pale grey or white. Eyes dark reddish brown. Antennae brown with ventral surfaces of all antennomeres except scapus and pedicellus blackish brown. Head (Fig. 297): Generally as in &&, but depression behind bases of antennae very indistinct. Eyes much more prominent and projecting hemispherically; their length contained only a little more than 2x in that of cheek. Antennae projecting over posterior margin of tergite III, otherwise as in &&. Thorax: Pronotum as in && but about as head, 1.8x longer than wide. Mesothorax almost 2.5x longer than head and pronotum combined. Mesonotum parallel-sided, very slightly widening at posterior margin and occasionally covered with a few very minute whitish granules. Mesosternum with a fine longitudinal median carina. Metasternum smooth and with an indistinct pale longitudinal median line. Tegmina oval, rounded apically, tapered towards the base and with a distinct, roundly conical central hump. Alae reaching to or projecting over posterior margin of abdominal tergite V. Abdomen: Median segment distinctly longer than metanotum, smooth and gently narrowing towards the posterior. Segments II–V very slightly increasing in length and in average about 6x longer than wide; VI


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slightly shorter. Tergites V and VI with a small, transverse posteromedian tubercle (sometimes lacking on VI). Tergite VII about 3/5 the length of VI and about 3x longer than wide; lateral margins in posterior portion slightly dilated and forming a ± distinct, rounded lobe, which extends by at best 1/3 the width of segment. Tergite VIII slightly longer than VII, prominently swollen and widening towards the posterior, about 2.5x longer than wide. IX a little shorter but considerably narrower than VIII, widened anteriorly, distinctly constricted medially and about 3x longer than wide. Anal segment shorter than IX, strongly laterally compressed and tectiform. Semi-tergites slightly tapered in apical portion and apex broadly triangular (Fig. 226). Interior surfaces apically swollen and armed with numerous minute spines (Figs. 23 & 28). Cerci almost reaching apex of anal segment, cylindrical basally and compressed apically, gently in-curving (Fig. 28). Poculum bulgy, strongly convex and cup-like and with a a blunt but distinct central spine; reaching about 1/3 the way along tergite IX. Legs: All very long and moderately slender. Profemora longer, the mesofemora about as long as head, pro- and mesonotum combined, metafemora projecting over posterior margin of abdominal segment IV and metatibiae roughly reaching apex of abdomen. Anterodorsal carina of profemora armed with 15–20 prominent and acute, triangular serrations. Posterodorsal carina set with 2–8 very minute teeth. Posteroventral carina with 14–20 small, pointed teeth. Anterodorsal and posteroventral carinae of protibiae densely but minutely serrate. Armature of mid and hind legs generally as in && but less distinct. The enlarged, pre-medial tooth or lobe of the posterodorsal carina may be lacking on the metatibiae. Probasitarsus longer than remaining tarsomeres combined, dorsal carina distinctly raised and ledge-like. Meso- and metabasitarsus with dorsal carina distinctly raised and rounded, all carinae minutely dentate; as long as remaining tarsomeres combined. Eggs (Figs. 159–160 & 356): Large, capsule oval and 1.5x longer than high. General colouration of capsule and micropylar plate creamish beige, operculum greyish mid brown. Lateroventrally with a very faint, slightly darker marking or several smaller markings. Capsule surface glossy and very unevenly granulose (70x). Micropylar plate more than ¾ the length of capsule, shaped like a bold inverted “Y”. Apices of all extensions rounded, outer margin mid brown. Micropylar cup placed in posteromedial gap of plate and posteriorly bordered by a slightly curved ridge. Median line indistinct and gently raised, reaching to polar-area. Polar-area with an oval blackish marking and a small, blunt tubercle in the centre. Operculum oval and slightly convex. Capitulum large and bowl-shaped with the outer margin folded up-wards, orange brown. Stalk distinct, pale brown. Measurements [mm]: Length including capitulum 6.0–6.2, length 5.3–5.4, width 3.6–3.7, height 4.0–4.1, length of micropylar plate 3.9. Comments: Bi (1994: 10) described Pharnacia jianfenglingensis from a single & collected at the Jianfengling Natural Protective Region in the Hainan Province of South China. The % was described and illustrated by Qi & Liu (1992: 250) based on specimens erroneously identified as Tirachoidea westwoodii (WoodMason, 1875) and originating in the Jinping County, Yunnan Province (900–1000 m) in Southern China. The specimens described and illustrated clearly relate to T. jianfenglingensis (Bi, 1994), which can apart from the distribution and shape of the semi-tergites of the anal segment be seen in the conspicuously large size. Qi & Liu (1992: 250) cited body lengths of 154.0–180.0 mm for %% and lengths of up to 280.0 mm for && (→ Table 39). The % from Northern Vietnam (Than Moi) in NHMW, erroneously identified and described as “Pharnacia maxima Bates” by Redtenbacher (1908: 450), is a typical specimen of T. jianfenglingensis. Bi (1994: 11) stated T. jianfenglingensis to feed on Terminalia catappa (Combretaceae) and Anthocephalus chinensis (Rubiaceae) in the Hainan Province and Qi & Liu (1992: 252) reported it to feed on Eucalyptus spp. (Myrtaceae) in Yunnan. According to Chen et al. (1994) T. jianfenglingensis has two generations per year in the latter locality. Since the late 1990’s T. jianfenglingensis is being cultured in Europe and was included on the Phasmid Study Group culture-list as culture No. 208 “Pharnacia sp. Vietnam”. The original eggs were purchased from a Nature Reserve Project in Cuc Phuong, Nho Quam District, Ninh Binh Province (some 100 km south of



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Hanoi) by P. Heusi (Uster, Switzerland) in 1996. In captivity in Europe it readily accepts oaks (Quercus robur, Q. petraea and Q. rubra, Fagaceae) and bramble (Rubus fruticosus, Rosaceae) as alternative food plants. Distribution (Fig. 405): Southern China (Hainan Province: Ledong County, Jianfengling & Yunnan Province: Jinping County 900–1000 m), Northern Vietnam (Ninh Binh Province: Nho Quam District, Cuc Phuong; Cho-Moi & Hoabinh) and Northeast Thailand (Doi Chieng Dao 4000 ft.). Number of specimens examined: 57 TABLE 39. Measurements [mm] of Tirachoidea jianfenglingensis (Bi, 1994). &, HT (after Bi)

%%

&&

Body

243.0

200.0–260.0 (280.0**)

139.5–183.0

Head

-

11.0–13.1

6.1–7.3

Pronotum

-

8.8–10.0

4.3–6.5

Mesonotum

71.0

38.5–47.5

23.7–31.0

Metanotum

34.0*

14.8–20.5

7.1–9.5

Median segment

-

11.6–14.0

14.5–16.5

Tegmina

-

-

11.1–15.2

Alae

-

-

74.5–90.0

Profemora

56.0

45.6–61.5

41.5–51.0

Mesofemora

45.0

35.5–48.0

34.5–45.0

Metafemora

53.0

41.5–58.5

44.0–54.0

Protibiae

-

47.0–71.0

49.0–62.0

Mesotibiae

-

34.5–46.0

34.0–46.0

Metatibiae

-

42.5–62.0

44.5–68.5

Subgenital pate

27.0

24.8–32.5

-

Antennae

46.0–56.5

46.0–56.5

82.0–90.0

* metanotum including median segment ** according to Qui & Liu (1992: 250)

Tirachoidea siamensis spec. nov. (Figs. 139, 142, 161–162, 218–220, 273, 298, 324, 357, 364, 406, 415, 423) HT, &: N. Thailand, Maetha, Lamphun, X.1998, Coll. Phimpisarn; BMNH(E) 2005-98 (BMNH). PT, 2 %%, 1 &: N. Thailand, Maetha, Lamphun, X.1998, Coll. Phimpisarn; BMNH(E) 2005-98 (BMNH). PT, 1 %, 1 &: N. Thailand, Maetha, Lamphun, X.1998, Coll. Phimpisarn (coll. PDB). PT, %: Thailand, Lomgao, Petchabun, Phimpisarn, 21.X.1985; BMNH(E) 2005-98 (BMNH). PT, 3 %%: N-Thailand, Dansai Loei, leg. 14.–15.IX.1986 (coll. FH, No. 0142-1 to 3). PT, 2 &&: N-Thailand, Chiang Mai, leg. Lehmann IV. 1994 (coll. FH, No’s 0142-4 & 7). PT, 6 &&: Central Thailand, Lemgao, Petchabun, X. 1985 (coll. FH, No’s 0142-5 to 6 & 8 to 11). PT, 1 %, 1 &: ex Zucht: F. Hennemann I. 1997, urspr.: N-Thailand, Chiang Mai (coll. FH, No’s 0142-12 & 13). PT, 2 %%, 60 eggs: ex Zucht F. Hennemann 1997, urspr.: N-Thailand, Chiang Mai (coll. FH, No’s 0142-14, 15 & ED). PT, 12 &&: Thailand, Lemgao, Petchabun, leg. Lehmann (coll. OC, No’s 00136 to 146). PT, &: Thailand, Prov. Kon Kaen, Bez. Phu Wiang, Tam Phra, 250m, Dry Dipterocarpus Forest, 2.11.1984, H. Felten leg. (SMFM).


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Further material: [6 &&, 1 %]: THAILAND: 1 &: Thailand, Chiang Mai, IV. 1994 (coll. OZ, No. 0245-1); 1 &: Thailand, Potvin vend. (ISNB); 1 &, 1 %: Thailand, Lemgao, Petchabun, don N. Mal (ISNB). 1 &: ex Zucht F. Hennemann 1997, urspr.: N-Thailand, Chiang Mai (coll. FH, No 0142-15). VIETNAM: 1 &: Tonkin, Hoabinh, 8.1918, R.V. de Salvaza leg. (BMNH); 1 &: Museum Paris, Cochinchina, Cap St. Jaques, com, Fuquet, 1913; Pharnacia westwoodi Wood-Mason L. Chopard & L. Berland det. (MNHN).

Diagnosis: Closely related to T. cantori (Westwood, 1859) but differing by: the relatively longer mesothorax; more globose vertex; more distinct tubercles of the vertex and entirely brown antennae of both sexes. && furthermore differ by: the slightly more slender body (maximum body width 12.0–14.0 mm); more shallow posteromedian excavation of the anal segment (Fig. 219); broader and rounded lateral lobes of abdominal tergite VII (Fig. 273); more distinct ridges of the praeopercular organ (Fig. 364) and orange or red armature of the legs. %% differ by the rounded apex of the semi-tergites of the anal segment (Fig. 220); longer meso- and metabasitarsi; more sparse armature of the legs and presence of a ± distinct, triangular median tooth on the posterodorsal carina of the meso- and metatibiae. Apart from these morphological features the geographical pattern of T. siamensis spec. nov. (Fig. 406) well separates it from T. cantori, which is distributed more southward and restricted to Peninsular Malaysia and very southern Myanmar (Mergui District). Also similar to T. jianfenglingensis (Bi, 1994) but easily distinguished by: the generally smaller size; more robust body and legs; less conical vertex; uniform colouration of the head and shorter basitarsi of both sexes. && furthermore differ by: the considerably more prominent armature of the mid and hind legs (Fig. 324); basally elevated outer ventral carinae of the mesofemora; less distinct praeopercular organ (Fig. 364); more distinctly excavated posterior margin of the anal segment (Fig. 219) and dark brown meso- and metasternum. %% clearly differ from those of T. jianfenglingensis by: lacking the white diagonal line of the tegmina; less prominent and less numerous serrations of the anterodorsal carina of the profemora as well as the much more slender and elongate semi-tergites of the anal segment (Fig. 220). T. jianfenglingensis is generally distributed more north and eastward than T. siamensis spec. nov., being restricted to SW-China, Northern Vietnam and the very northern portions Thailand (Fig. 405). The eggs readily differ from all other known eggs of the genus by the creamish grey colour and very defined black lateroventral marking of the capsule, slender micropylar plate and narrowed polar-area as well as the very large, conical capitulum (Figs. 161–162). Etymology: The name refers to the distribution of this new species and is derived from Thailand’s old name “Siam”. Description: The colouration is described from both dried and live specimens. && (Fig. 142): Of medium size (body length 194.0–229.0 mm) and moderately robust for the genus (12.0–14.0 mm) with prominent leg armature. General colouration of body and legs varying from pale yellowish or greyish over mid to almost blackish brown, the body usually all over set with numerous paler and darker brown speckles. Occasionally the dorsal surface of the head and body may be furnished with several large but irregular pale yellowish markings. Cheeks with two faint and weakly defined longitudinal streeks in the lower portion. Meso- and metasternum dark brown with a few whitish spots roughly arranged in two longitudinal rows. Abdominal sternites with ± distinct whitish mottling. Legs with irregular often very indistinct whitish or pale grey transverse bands, which are more defined on the ventral surfaces. Complete armature of legs dark red with black points; Pre-median and apical dorsal lobes of the meso- and metatibiae blackish brown to black. Eyes dark reddish brown. Antennae reddish mid to dark brown. Head (Fig. 298): About 1.5x longer than wide, vertex strongly rounded and set with a pair of ± distinct forward-pointing spiniform tubercles; the dextral one considerably larger than the sinistral. Between the bases of antennae with a deep oval impression. Eyes large, circular and strongly convex, the anterior margin slightly



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triangular; length contained a little more than 3x in that of cheek. Antennae reaching about 3/4 the way along mesonotum and consisting of about 30 antennomeres. Scapus dorsoventrally flattened, 2.3x longer than wide and very gently narrowed towards the base. Pedicellus a little more than 1/3 the length of scapus, indistinctly longer than wide and cylindrical. III slightly longer than pedicellus, IV strongly shortened, V–XV increasing in length. All finely bristled. Thorax: Pronotum slightly narrower and shorter than head, almost 1.3 x longer than wide and conspicuously widened towards the posterior. Anterior margin raised, followed by a well defined transverse furrow and with a small rounded impression at the lateral angles. Median transverse depression rather distinct but short and covering less than half the width of segment. Mesothorax about 1.8x longer than head and pronotum combined, very slightly constricted at anterior margin and broadened at the posterior. Mesosternum with a blunt longitudinal carina along lateral margins. Metanotum distinctly shorter than mesonotum, about 1.2x longer than wide, rectangular and with a very minute spine posteromedially. Very minute, scale-like traces of tegmina and alae present (ca. 1.0 mm) Abdomen: Median segment almost 4/5 the length of metanotum, slightly trapezoidal and with the anterior margin rounded. Segments II–VI increasing in length, II 1.2x, VI longest segment and about 2x longer than wide. Tergites II–VII with a transverse, scale-like tubercle which is most decided on V; sometimes lacking on II and III. VII about 2/3 the length of VI, posterolaterally dilated into a broad rounded lobe, which extends by about half the width of segment (Fig. 273). Praeopercular organ formed by two short, but rather acute converging carinae, which terminate in a minute spine posteriorly, near posterior margin of sternum VII (Fig. 364). Tergite VIII a little longer than VII and longer than IX and X combined, distinctly narrower than previous, slightly constricted medially and about 3x longer than wide. IX about half the length of previous, 1.3x longer than wide and strongly convex. Anal segment a little longer than IX, with a very indistinct median carina and a broad triangular excavation posteromedially; posterolateral angles roundly triangular (Fig. 219). Supraanal plate broad, rounded, with a fine median carina and a small posteromedian notch. Cerci small, oval in cross-section, gradually tapered towards a pointed tip and reaching apex of anal segment. Gonapophyses dull yellowish brown, elongate, up-curving and slightly projecting over anal segment. Subgenital plate strongly keeled, boat-shaped and projecting over apex of abdomen by about half the length of anal segment (Fig. 218). Legs: All of moderate length and rather robust, the meso- and metafemora very gently down-curving and slightly swollen basally. Profemora as long as pro- and mesonotum combined, mesofemora slightly shorter than mesothorax, metafemora slightly projecting over posterior margin of abdominal segment IV and hind tarsi reaching about half way along tergite VIII. Anterodorsal carina of profemora armed with 10–14 pointed, triangular serrations. Posterodorsal carina set with 7–11 small triangular teeth. Posteroventral carina with 12– 14 pointed serrations. Anterodorsal carina of protibiae sparingly serrate, posterodorsal carina with 6–8 minute teeth, posteroventral sparingly serrate. All carinae of meso- and metafemora armed with strongly pointed spines and teeth of various sizes; these become more increasingly larger towards the base of femur and are in general more minute on the dorsal carinae. Two outer ventral carinae of mesofemora gently elevated subbasally, the posteroventral carina bearing 2–3 conspicuously enlarged and broadened lobe-like triangular teeth in that portion. Posterodorsal carina with a ± large, triangular sub-basal tooth and a slightly enlarged tooth apically (Fig. 324). Medioventral carina of meso- and metafemora armed with 5–7 prominent, hooked spines which increase in size towards base of femur; the 2nd and 3rd being largest. All carinae of meso- and metatibiae densely armed with pointed spines; more sparingly and minutely on dorsal carinae. Posterodorsal carina armed with a very prominent, foliaceous triangular lobe pre-medially and forming a conspicuous rounded and dentate lobe apically. Spines of the medioventral carina considerably longer and more slender than those of the two outer carinae. Probasitarsus with all carinae distinctly but uniformly elevated and unarmed; as long as remaining tarsomeres combined except claw. Meso- and metabasitarsi with a strongly raised and apically rounded dorsal carina, all carinae serrate; a little longer than following three tarsomeres combined.


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%% (Fig. 139, 423): Of moderate size (body length 128.5–139.0 mm) with long alae (66.5–71.0 mm). General colouration of body and legs yellowish to greyish cream-brown, occasionally with a few small white spots on mesonotum. Cheeks sometimes with a very indistinct and poorly defined brown marking close to ventral margin of head capsule. Abdominal sternites with irregular white mottling and single spots which become increasingly distinct towards the hinder segments. Meso- and metasternum greyish mid brown and irregularly set with conspicuous white spots. Lateral margins of abdominal tergite VIII whitish, IX with a longitudinal white marking along the lateral margins. Tegmina creamish dark brown with the central hump very dark brown to black. Anterobasal portion of tegmina broadly white. Costal region of alae plain yellowish or greenish pale to mid brown, with a broad white longitudinal stripe along the anterior margin which terminates about ¼ off the base. Anal region transparent greyish brown with brown veins. Ventral surfaces of legs with ± distinct white mottling; apices of all femora and tibiae whitish. Complete armature of the legs black. Antennae plain orange mid brown. Eyes reddish brown. Head: Generally as in &&, but eyes much more prominent and projecting hemispherically; their length contained less than 2x in that of cheek. Antennae projecting over posterior margin of abdominal tergite II, otherwise as in &&. Thorax: Pronotum narrower and shorter than head, slightly widened towards the posterior and about 1.5x longer than wide, otherwise as in &&. Mesothorax almost 2x longer than wide and slightly widened at posterior margin. Mesonotum parallel-sided and with a very indistinct longitudinal median carina. Mesosternum with a shallow longitudinal median carina. Metasternum smooth. Tegmina oval, narrowed towards the base and broadly rounded posteriorly; central hump prominent and conical. Alae reaching about half way along abdominal tergite VI. Abdomen: Median segment almost 2x longer than metanotum, gently narrowing towards the posterior. Segments II–VI ± 3.6x longer than wide, parallel-sided. VII about 2/3 the length of VI, constricted medially and slightly dilated posterolaterally. VIII slightly longer than VII, strongly swollen and broadening towards the posterior, almost 2x wider than VI. IX as long as VIII, distinctly constricted medially and anterior margin almost 2x broader than posterior margin. Anal segment a little shorter than IX, strongly laterally compressed and tectiform. Semi-tergites tapered and gently down-curving in their apical half; apex narrow and rounded (Fig. 220). Interior surfaces of semi-tergites apically densely covered with minute black teeth. Cerci cylindrical, gradually tapered towards a rather acute apex and slightly in-curving; almost reaching apex of anal segment. Poculum very bulgy, strongly convex and cup-like, marginated posteriorly and with a blunt backward pointing central spine. Legs: All moderately long and slender. Profemora longer than head, pro- and mesonotum combined, mesofemora slightly longer than pro- and mesonotum combined, metafemora reaching about half way along abdominal segment V and metatibiae reaching to posterior of tergite IX. Anterodorsal carina of profemora with 7–17 distinct triangular teeth, posteroventral carina with a somewhat smaller number of slightly smaller and more slender teeth. The posterodorsal carina of profemora occasionally set with at best 6 minute teeth. Mid and hind legs with all carinae sparingly and rather minutely dentate; the teeth more numerous on the ventral carinae. Medioventral carina of meso- and metafemora with 5–12 small but pointed spines. Posterodorsal carina of meso- and metatibia forming a conspicuous, rounded and dentate lobe apically and often with a ± enlarged and broadly triangular tooth pre-medially. Probasitarsus distinctly longer than remaining tarsomeres combined, all carinae gently and uniformly elevated but unarmed. Mesobasitarsus slightly shorter, metabasitarsus a little longer than combined length of remaining tarsomeres; Dorsal carinae gently rounded and smooth or minutely dentate; ventral carinae dentate. Variation: && in particular show considerable variation concerning to the size, colouration, size of the two spiniform tubercles of the vertex and degree of leg armature. A captive reared & in coll. FH (No. 0142-15) is remarkably small, measuring a body length of only 151.0 mm. As the small size may have been caused by insufficient breeding conditions or alternative foodplants used it is not designated a type and excluded from REVISION OF PHARNACIINI


247

the table of measurements below. One & PT in the first author’s collection (coll. FH, No. 0142-5) has the complete dorsal body surface furnished with irregular, large pale yellowish markings. Another PT in the same collection (coll. FH, No. 0142-7) has the tubercles of the vertex very weakly developed and only represented by pair of shallow and blunt humps. %% show variation only concerning to the number and degree of leg armure and presence or absence of a enlarged pre-median teeth on the posterodorsal carina of the meso- and metatibiae. Eggs (Figs. 161–162, 357, 415): Medium-sized to large, capsule strongly rounded but slightly longer than high and conspicuously narrowed posteriorly. General colouration of capsule and micropylar plate creamish grey, operculum black, capitulum dull reddish brown (Fig. 415). Lateroventrally with a very prominent and well defined black marking. Capsule surface shiny and very minutely granulose (40x). Micropylar plate more than ¾ the length of capsule, shaped like an elongate inverted “Y”. Apices of all extensions rounded, outer margin blackish brown. Micropylar cup placed in posteromedial gap of plate, indistinct and posteriorly bordered by a slightly curved ridge. Median line indistinctly raised, very slender and almost reaching polar-area. Polar-area with an oval blackish marking and a small, blunt spine in the centre. Operculum slightly oval, almost flat. Capitulum very large, at least 2/3 the width of operculum, irregularly conical and with numerous carinae and impressions. Stalk short, white. Measurements [mm]: Length including capitulum 4.8–5.6, length 3.8–4.8, width 3.1–3.8, height 3.5–4.2, length of micropylar plate 3.0–3.8. Comments: Seen from the numerous specimens available T. siamensis spec. nov. appears to be quite common in Northern and Central Thailand. The record “Cochinchina” based on a & in MNHN appears doubtful although the specimen clearly represents T. siamensis spec. nov. In the early 1990’s culture-stock was imported to Europe originating in Chiang Mai (N-Thailand) and subsequently included on the Phasmid Study Group culture-list as culture No. 197. Oaks (Quercus robur, Q. petraea, Q. rubra, Fagaceae) and bramble (Rubus fruticosus, Rosaceae) are readily accepted as alternative foodplants, but the species has proven rather difficult to maintain in culture. TABLE 40. Measurements [mm] of Tirachoidea siamensis spec. nov. HT, & (BMNH)

PT, && (coll. FH)

PT, %% (coll. FH)

Body

229.0

194.0–225.0

128.5–139.0

Head

14.0

11.9–14.0

5.8–6.6

Pronotum

11.0

8.5–9.5

4.3–4.9

Mesonotum

43.0

34.0–43.5

20.8–23.2

Metanotum

18.5

14.7–18.0

18.5–23.0*

Median segment

12.0

10.3–12.5

-

Tegmina

-

-

9.3–11.3

Alae

-

-

66.5–71.0

Profemora

48.0

44.5–53.0

34.2–37.0

Mesofemora

40.0

32.0–41.0

27.1–30.0

Metafemora

47.0

39.0–47.5

34.5–37.0

Protibiae

50.0

45.8–59.5

43.6–44.7

Mesotibiae

38.0

31.0–41.0

28.6–30.0

Metatibiae

46.5

39.8–52.8

37.0–40.0

Antennae

> 27.0

43.0

62.5–68.0

* including median segment


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Distribution (Fig. 406): North and Central Thailand (Dansai Loei; Lamphun: Maetha; Petchabun: Lemgao; Chiang-Mai & Province Khon Kaen: Phu Wiang, Tam Phra 250 m) and N-Vietnam (Hoabinh). S-Vietnam (Cochinchina) [with doubt]. Number of specimens examined: 42

Tirachoidea westwoodii (Wood-Mason, 1875) stat. rev. (Figs. 140, 211, 303, 403, 407–411) Phibalosoma westwoodii Wood-Mason, 1875: 216. LT [by present designation], &: Assam, Samaguting in Assam, leg. J. Butler (depository unknown); PLT, &: Assam, Nazeerah, leg. Foster (depository unknown). Wood-Mason, 1877a: 161. Phibalosoma westwoodi, Wood-Mason, 1877b: 347, pl. 3: 1a–d (& terminal abdominal segments & head). [Unjustified emendation]. Tirachoidea westwoodi, Kirby, 1904a: 359. Pharnacia westwoodi, Redtenbacher, 1908: 451. Brock, 1996: 27. Otte & Brock, 2005: 265. [Not: Pharnacia westwoodi, Giglio-Tos, 1910: 43 (& from Java). Misidentification = Pharnacia sumatrana (Brunner v. Wattenwyl, 1907)] [Not: Tirachoidea westwoodi, Qi & Liu, 1992: 250—description of presumed % from Yunnan Province, S-China. Misidentification = T. jianfenglingensis (Bi, 1994)] [Not: Pharnacia westwoodi, Chen & He, 2007: 351, figs. 317 a-d. Misidentification relating either to T. jianfenglingensis (Bi, 1994) or T. siamensis spec. nov.] Further material: [4 %%]: 2 %%: N-Myanmar, Tenasserim Highland, 18.III.–19.IV.1996, leg. Lehmann (coll. FH, No’s 0211-1 & 2); 2 %%: NMyanmar, Tenasserim 5.1995 –18.3.1996 (coll. OC).

Diagnosis: This species is closely related to T. jianfenglingensis (Bi, 1994) && differing by: the lack of dark transverse stripes in the posterior portion of the vertex; slightly shorter mesothorax; posteromedian tubercle of abdominal tergite VII (Fig. 407); shape of the anal segment (Fig. 408) and longer subgenital plate, which slightly extends over the apex of the abdomen (Fig. 407). %% are easily distinguished from those of T. jianfenglingensis by: more elongate body and legs; relatively longer body segments; less convex vertex with the pair of humps decidedly less distinct (Fig. 303); more slender tergite VIII; slightly shorter and sharply truncate semi-tergites of the anal segment (Fig. 228); more elongate and distinctly in-curving cerci; more numerous triangular serrations of the anterodorsal carina of the profemora; greater number of serrations of the anterodorsal carina of the protibiae; more numerous and minute spines of the mid- and hind legs as well as the not rounded but uniformly raised dorsal carina of the meso- and metabasitarsi. Etymology: Dedicated to Dr. John Obadiah Westwood, one of the greatest English entomologist at that time. Description, &&: These are only known from the very brief and insufficient original diagnosis (WoodMason, 1875: 216) and a set of very accurate drawings of the head and apex of the abdomen provided by Wood-Mason subsequently (1877b, pl. 3: 1a–d, here reproduced as Figs. 407–411). The body length cited in the original description is ca. 240.0 mm. Wood-Mason (1875: 216) stated T. westwoodii to be closely related to T. cantori (Westwood, 1859) and to differ from this species by the larger lateral lobes of abdominal tergite VII and shape of the head, which has the vertex convex and with two rounded humps of unequal size. Since Wood-Mason made no mention of any conspicuous leg armature, this must be presumed to be rather regular and similar to T. cantori, lacking any enlarged teeth or lobes. The following brief diagnosis is based on the set of measurements attached to the original descriptions and



249

illustrations of the head and apex of the abdomen provided by Wood-Mason (1877b). Head about 1.4x longer than wide with the vertex distinctly conically raised and set with two unequal blunt humps; the dextral one being decidedly larger than the sinistral (Figs. 409–411). Eyes circular and moderately sized, their length contained about 3.5x in that of the cheek. Antennae reaching about two thirds the way along the mesothorax. Scapus about 2.3x longer than wide and flattened dorsoventrally. Pedicellus cylindrical and about 1/3 the length of scapus. Mesothorax about 1.7x longer than the head and pronotum combined. Abdominal tergite VII with a distinct, blunt median tubercle at posterior margin (Fig. 407). Lateral margins of tergite VII greatly elevated and in posterior half forming a large rounded lobe, which extends laterally by about half of body width (Fig. 408). Tergite VIII slightly longer than VII and longer than IX and X combined, gently constricted medially and about 2.3x longer than wide. Anal segment indistinctly longer than wide and dorsally with four conspicuous rounded swellings. Posterior margin with a shallow and widely triangular excavation. Supraanal plate very small and transverse (Fig. 408). Subgenital plate strongly keeled and boat-shaped, strongly convex in the median portion and projecting over apex of abdomen by about half the length of the anal segment (Fig. 407). %% (Fig. 140): Very long (body length 164.0–165.0 mm) and very slender for the genus with long alae (70.0–70.5 mm), body of uniform width (± 2.5 mm). General colouration of body and legs more or less uniformly greyish to greenish mid brown, mesonotum mid to dark brown. Metasternum with a slight greenish wash. Complete spination of the legs ochre with black points. Tegmina greenish brown, with an very faint pale diagonal band and a longitudinal white marking in the basal half of the anterior margin. Costal region of alae brown with a slight greenish wash and dull greenish brown veins, anterior margin with a bold white longitudinal line which terminates about 1/3 off the base. Anal region transparent greyish brown with dark brown veins. Antennae mid to dark brown. Eyes reddish mid brown irregularly marbled with darker brown. Head (Fig. 303): Oval, 1.5x longer than wide, vertex slightly convex and with two very indistinct low humps. Between the eyes with an oval, slightly raised and smooth area. A very fine impressed transverse line between the bases of the antennae. Eyes prominent, circular and projecting hemispherically from head capsule; their length contained about 2.3x in that of cheek. Antennae with all segments densely bristled and at least projecting over posterior margin of metanotum (broken in all examined specimens). Scapus flattened dorsoventrally, about 3x longer than wide, very gently constricting towards the base and with a distinct longitudinal depression ventrally. Pedicellus less than ¼ the length of scapus, cylindrical. III longer than pedicellus, IV strongly shortened, V to X increasing in length. Following antennomeres very elongate. Thorax: Pronotum about 1.5x longer than wide, rectangular, shorter and narrower than head. Median line distinct and impressed. Median transverse depression distinct, curved but very short and not reaching lateral margins of segment. Mesonotum 3x longer than head and pronotum combined, smooth. Mesosternum with a distinct and acute longitudinal median carina. Metasternum very slightly tectiform. Tegmina oval, constricted basally and with an acute, conical hump in the centre. Alae reaching about half way along abdominal tergite V. Abdomen: Median segment 2.5x longer than wide and distinctly longer than metanotum, slightly narrowing towards the posterior. Segments II–VI 6.5–7x longer than wide, all slightly constricted medially. Tergites V and VI with a small, scale-like posteromedian tubercle. Segment VII slightly more than 2/3 the length of previous and gently widening towards the posterior; posterior margin about 1.5x broader than anterior margin. Tergite VIII slightly longer than wide, strongly swollen and gradually broadening towards the posterior; posterior margin 2x broader than posterior margin of VII. IX about 1.5x longer but narrower than VIII, 3x longer than wide and strongly constricted medially. Anal segment as long as VIII, laterally compressed and strongly tectiform. Semi-tergites short with the apex acutely truncate and the lower corner slightly elevated (Fig. 211). Interior surfaces densely covered with minute blackish brown teeth. Cerci elongate, oval in cross-section, gradually tapered and with the apical portion distinctly in-curving; apex pointed. Poculum strongly convex, reaching about half way along tergite IX and with a blunt, transverse central spine (Fig. 211). Legs: All very long and slender, with all carinae of mid and hind legs minutely but densely serrate, but


HENNEMANN & CONLE

lacking any enlarged teeth or lobes. Profemora longer than head, pro- and mesonotum combined, mesofemora about as long as combined length of pro- and mesonotum, metafemora reaching half way along abdominal segment V and metatibiae slightly projecting over apex of abdomen. Anterodorsal carina of profemora with 25–30 distinct and triangular black serrations; posteroventral carina with a similar number of small but acute teeth. Anterodorsal and posteroventral carinae of protibiae densely but minutely serrate, other carinae smooth. Medioventral carina of meso- and metafemora indistinct and armed with 4–6 distinct but elongate and needlelike spines. Occasionally a slightly enlarged, triangular median tooth is present on the posterodorsal carina of meso- and metatibiae. Probasitarsus distinctly longer than remaining segments combined, with all carinae slightly elevated and unarmed. Meso- and metabasitarsi slightly longer than remaining segments combined, dorsal carina gently raised and almost of uniform height, all carinae minutely dentate. Comments: Wood-Mason (1875: 216) described Phibalosoma westwoodii as being closely related to T. cantori (Westwood, 1859). He based his description on two && collected in the Naga Hills of NE-India (close to the border to Myanmar), one from Nazeerah and one from Samaguting. Although extensive research in collections world-wide has not traced Wood-Mason’s types, the latter specimen is here selected as the lectotype in order to fix Wood-Mason’s species. The type specimens were presumed in NZSI by most subsequent authors, but research has revealed there are no types of Phasmatodea in Calcutta at all (personal communication Dr. M. S. Shishodia, NZSI). Inquiries to the museum in New Dehli have received no reply. Should they not be in New Dehli, both specimens must be presumed lost and a neotype designated. Although the original diagnosis is very brief and insufficient, Wood-Mason’s very accurate illustrations of the head and last abdominal segments of the & (1877b, pl. 3: 1a–d) well identify this species. The presumed % was described and illustrated by Qi & Liu (1992: 250) based on specimens collected in the Yunnan Province of China. These are however clearly misidentified and represent T. jianfenglingensis (Bi, 1994). Hence China can be regarded an erroneous record.This presumably also concerns to the specimens referred to as "Pharnacia westwoodi" by Chen & He (2007: 350). Four %% from the Highland of Tenasserim (N-Myanmar) in the author’s collections appear to represent the so far unknown opposite sex of T. westwoodii. This is indicated by the large size, elongate body and shape of the head. Eggs unknown. TABLE 41. Measurements [mm] of Tirachoidea westwoodii (Wood-Mason, 1875) stat. rev. %% (coll. FH)

Body

164.0–165.0

Head

6.1–6.2

Pronotum

4.7–5.0

Mesonotum

32.5–33.0

Metanotum

7.5–7.8

Median segment

14.0–14.3

Tegmina

11.0–11.4

Alae

70.0–70.5

Profemora

47.3–48.8

Mesofemora

37.3–39.5

Metafemora

46.0–50.0

Protibiae

64.0–64.5

Mesotibiae

42.6–43.1

Metatibiae

56.9–58.0

Antennae

> 55.0



251

Distribution (Fig. 403): NE-India (Naga Hills: Nazeerah & Samaguting) and N-Myanmar (Highland of Tenasserim). Number of specimens examined: 4

7 Species incertus sedis Phobaeticus sp. 1 %: no data, 198, f. Pharnacia acanthopus Burm. (RMNH).

Comments: A large (body length 154.2 mm) and slender % which clearly belongs in Phobaeticus Brunner v. Wattenwyl and appears to be closely related to Ph. magnus nom. nov. and Ph. pinnipes (Redtenbacher, 1908). The specimen does not bear any locality and is rather incomplete, lacking both fore legs and the left mid leg. The general colour is a buff greyish brown, the legs being slightly greenish and the meso- and metapleurae greyish white. The head is indistinctly longer than wide, smooth and strongly constricted towards the posterior margin which is distinctly triangular. The alae are well developed and slightly project over the posterior margin of tergite IV (61.5 mm). There is a bold white longitudinal line along the anterior margin of both the alae and tegmina, the latter having an additional white central spot. The genitalia are typical for Phobaeticus, the cerci being very slender, elongate, strongly in-curving and tapered towards the apex. The semi-tergites of the anal segment are elongate and constricted towards the apex. The poculum is strongly convex, has a distinct transverse central hump and reaches about half way along tergite IX. The meso- and metatibiae each have three conspicuously enlarged, triangular lobes on their posterodorsal carina.

8 Discussion of Pharnaciini Günther, 1953 The five genera here revised in detail, Baculonistria gen. nov., Pharnacia Stål, 1877, Phobaeticus Brunner v. Wattenwyl, 1907, Phryganistria Stål, 1875 and Tirachoidea Brunner v. Wattenwyl, 1893, together form a monophyletic taxon, the tribe Pharnaciini Günther, 1953. Pharnaciini shows an evolution from moderately basal to more derived forms, which can be seen in several evolutionary trends like the length reduction of the median segment, length reduction or complete loss of the tegmina and alae, increasing elongation of the mesothorax and distributional pattern (→ 8.1). Due to morphological features of the genital exosceleton and the profemora as well as a different copulation habits discussed above (→ 4.) Pharnaciini is here removed from the subfamily Phasmatinae and family Phasmatidae s. str., and transferred to the newly introduced subfamily Clitumninae (→ 5.). Apart from Pharnaciini, Clitumninae at present only contains one further tribe, the Clitumnini. These two tribes show a rather clear evolution from basal, often strikingly large forms which often have alate or brachypterous %% (Pharnaciini), to more derived and specialized often remarkably slender and completely apterous forms with a strongly shortened median segment (Clitumnini). Both have similar geographical distributions, although Clitumnini appears to have mostly radiated on the Asian mainland, while Pharnaciini has its centre of distribution in Sundaland (Fig. 82). Those genera of Pharnaciini in which the median segment is equal in length or longer than the metanotum and %% have fully developed alae, this is Pharnacia and Tirachoidea, may be interpreted as the evolutionary most basal taxa of the tribe. On the other side, Phryganistria and Baculonistria gen. nov. in which the median segment may be as short as only 1/8 the length of the metanotum and both sexes are completely apterous, can be regarded the most derived and evolutionary modern taxa amongst Pharnaciini. Phobaeticus shows consid-


HENNEMANN & CONLE

erable polymorphy concerning to numerous features of the insects and eggs and even shows a rather clear evolution intragenerically. This can be seen in several steps of length reduction of the median segment as well as an reduction and final loss of the tegmina and alae. Evolutionary, these steps can be interpreted as a transitory from the primitive to the more derived forms of Pharnaciini. Principally, the more derived forms (Phobaeticus, Phryganistria and Baculonistria gen. nov.) tend to show much more intrageneric polymorphy concerning to numerous features of the insects and eggs than do the presumed evolutionary older forms (Pharnacia and Tirachoidea). The more derived genera of Pharnaciini, Phryganistria and Baculonistria gen. nov., show striking affinity to certain genera of Clitumnini. This is for instance seen in the very slender and elongate body, complete loss of tegmina and alae in both sexes and conspicuously shortened median segment. Baculonistria gen. nov. has the antennae conspicuously shortened, with the antennomeres IV–XII of && being only indistinctly longer than wide, and && bear two very minute spines between the eyes. Both features are unique in Pharnaciini but frequently in most genera of Clitumnini. A prominent praeopercular organ like in && of Phryganistria and Phobaeticus is also present in most representatives of e.g. the genus Ramulus Saussure, 1862. The shape of the semi-tergites of the %%’s anal segments is also very similar in certain taxa of both tribes, having the apex more or less strongly elevated and finger-like. However, the strongly dentate carinae of the extremities and longer antennae readily distinguish Phryganistria and Baculonistria gen. nov. from Clitumnini. The most prolific distinguishing feature is presented by the eggs, which have a distinctly stalked, knob- or hat-like capitulum in Pharnaciini (→ 5.2.4). Although not a member of Phasmatidae s. str. some representatives of Pharnaciini strikingly resemble certain genera of the tribe Phasmatini (Phasmatidae s. str.: Phasmatinae, → 4.3.1). For instance, Tirachoidea and Peloriana Uvarov, 1940 exhibit obvious similarity concerning to the shape and structure of the head and thorax as well as the armature of the legs. Except for differing by the key features of Phasmatinae: Phasmatini, such as well developed tegmina and alae, a median segment which is distinctly longer than the metanotum, ocelli, lack of a praeopercular organ and foliaceous cerci, && of Peloriana appear like winged versions of Tirachoidea. Another striking example is represented by Phobaeticus and Ctenomorpha Gray, 1833. Both genera are typical for containing the most slender and elongate taxa within their tribes, and as in the other examples it is only the key features that distinguish these two genera. Ctenomorpha shows the same conspicuous prominent, black dorsal serrations on the profemora typical for most species of Phobaeticus. For instance, the % of Ctenomorpha gargantua Hasenpusch & Brock, 2006 looks remarkably alike the % of e.g. Phobaeticus magnus nom. nov. Similarity not only concerns to the insects but to the eggs as well. The eggs of both tribes are typical for being ovoid or lens-shaped and having a conspicuous knob or hat-like, stalked capitulum. The lack of a median line distinguishes the eggs of Phasmatini from those of Pharnaciini, which always have a median line. Fellenberg (1993) described the egg of what appears to be Paronchestus pasimachus (Westwood, 1859) but erroneously stated it to be a species of Onchestus Stål, 1875 (Brock & Hasenpusch, 2005: 17). This shows a similar polar keel to that seen in eggs of certain Phobaeticus. The main characteristics, variation and intergeneric relationships between the five genera of Pharnaciini are provided below. The possible relationship between the genera here suggested is shown in the dendrogram illustrated in figure 367, but a full phylogenetic analysis of this tribe would be inappropriate at present for several reasons. Two species of Baculonistria gen. nov. are only known from the original descriptions, numerous species are only known from a single sex and the eggs of 19 species have not been recognized until now. Resolving the phylogeny of Pharnaciini would certainly require more material than is currently available. A) Pharnacia and Tirachoidea: These two genera are obviously very closely related to each other and can with high probability be interpreted as adelphotaxa, which together form the sister-group of the remaining Pharnaciini, namely Phobaeticus + Phryganistria + Baculonistria gen. nov.. Both genera are characterized by the rather massive and robust body (&& in particular); the more or less prominently laterally expanded abdominal tergite VII of both sexes; long median segment which is slightly longer or at least equal in length



253

(&&), or considerably longer than the metanotum (%%); alate %%; externally lacking (Pharnacia) or indistinct (Tirachoidea) praeopercular organ of &&, as well as the typical micropylar plate of the eggs, which is shaped like a bold inverted “Y”. In neither genus the subgenital plate of && is strongly elongated or lanceolate and never projects considerably over the anal segment, as is the case in certain taxa of the other three genera. The Y-shaped micropylar plate has been developed by extension of the posterolateral angles of the plate on both sides of the posteromedian gap. The apices of these two posterior “arms” are broadened and truncate in Pharnacia or slender and rounded in Tirachoidea. Furthermore, the eggs of both genera differ by the different structure of the capsule surface which is rough, minutely punctated or granulated and not glossy in Pharnacia, but almost completely plain and strongly glabrous in Tirachoidea. The almost spherical capsule of Tirachoidea does not possess any indentions or impressions near the operculum and polar-area (Figs. 157–166), whereas the capsule shows indentions and may even be slightly laterally compressed in Pharnacia (Figs. 145– 156). Insects of Tirachoidea are furthermore distinguished from Pharnacia by the spherical or conical vertex of both sexes, which usually bears a pair of minute granules, humps or blunt spines (Figs. 296–304), as well as the strongly raised and rounded dorsal carina of the basitarsi. The Y-shaped micropylar plate may be regarded a synapomorphy of Pharnacia + Tirachoidea, but other apomorphies of Pharnacia and Tirachoidea are difficult to define. An intrageneric systematization of these two genera would be inappropriate at this point due to certain sexes and eggs are still unknown. B) Phobaeticus: This genus shows strong intrageneric variability and polymorphy concerning to numerous external features of the insects and eggs. It is apparently closely related to Phryganistria and several features, like the prominent praeopercular organ of && and structures of the genitalia of %%, indicate it may be regarded the adelphotaxon of Phryganistria (Fig. 367). It is easily distinguished from Pharnacia and Tirachoidea by the presence of a distinct praeopercular organ in &&, which is usually formed by a pair of spines or lobes at the posterior margin of abdominal sternum VII, the very elongate, finger-like semi-tergites of the anal segment and elongate, apically constricted, in-curving cerci of %%. In general, the adults are considerably more slender and elongate than members of either Pharnacia or Tirachoidea, many species being extremely long and stick-like insects. The genus contains some of the world’s longest known insect and && of several species easily exceed body lengths of 25 cm with the body no wider than 5 mm. Phobaeticus chani Bragg spec. nov., for instance, is the world’s longest know extant insect with a maximum recorded body length of almost 35.7 cm and an overall length with the fore legs stretched out straight of almost 60 cm. Many external features of the insects, e.g. the length relation of the median segment and metanotum, shape of the head, leg armature, length of the tegmina and alae of %%, length of the subgenital plate of && and shape of the %% anal segment are subject to considerable intrageneric variation. The median segment is either slightly longer, equal in length or distinctly shorter than the metanotum. Correlating to the length reduction of the median segment, %% either have fully developed alae and are capable of active flight, may be brachypterous or even entirely apterous. && of species with apterous %% have a median segment shorter than the metanotum, whereas it is equal or longer than the metanotum in species with winged %%. The head is either elongate and distinctly longer than wide with the vertex flat, to gently oval, or may even be almost spherical with the vertex conspicuously rounded. The subgenital plate of && varies from being very short and staying behind the posterior margin of the anal segment to strongly elongated and lanceolate; in some species it may project over the anal segment by as much as the combined length of the terminal 3–4 abdominal tergites. The mesonotum is usually smooth and destitute of spines or granules. Only one species, Phobaeticus foliatus (Bragg, 1995) from Borneo, has the mesonotum in both sexes armed with several prominent spines. Tergite VII of && is either slender with the lateral margins not obviously dilated or has the lateral margins expanded into a rounded lobe. As an extreme Ph. rex (Günther, 1928) has this lobe irregularly dentate and projecting laterally by as much as the complete length of tergite VII. Ph. foliatus (Bragg) even has a more or less prominentm, rounded and dentate lateral appendage on each abdominal tergite, again a feature unique in Pharnaciini. The legs are very long and slender and, if compared with Pharnacia and Tirachoidea, are longer in relation to the body.


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The armature shows extreme polymorphy and may even vary considerably within single species. Ph. foliatus (Bragg) as an extreme has all femora and tibiae covered with several very large and foliaceous appendages. Not only do the insects show considerable variation, but the eggs as well. This concerns to the shape and structure of the capsule surface as well as to the shape and size of the micropylar plate (Figs. 169–185). The capsule is more or less strongly laterally compressed and surrounded by a ± prominent dorsoventral keel, which gives the egg a lens-like general appearance. In Ph. chani Bragg spec. nov. the dorsoventral keel is protruded into large, foliaceous and clockwisely curved extensions. Seen laterally the polar area and the micropylar region are impressed to a variable degree (depending on height of keel). The capsule surface is either almost smooth and glabrous, very slightly rugulose, distinctly verrucose or wrinkled. The micropylar plate is remarkable for being less than 2/3 the length of the capsule (longer in all other genera of Pharnaciini). It is often transverse and bilobed or kidney-shaped and has a prominent posteromedian impression; rarely longer than wide. The short, often transverse micropylar plate and distinct dorsoventral keel of the egg capsule may be interpreted as autapomorphies of Phobaeticus. C) Phryganistria: This genus comprises four very long and slender, evolutionary rather derived species, characteristic for the shortened median segment, which is at best 2/5 the length of the metanotum. Both sexes are completely apterous and resemble certain of the larger representatives of Clitumnini, and the sexual dimorphism is much less developed than in Pharnacia, Tirachoidea and Phobaeticus. The shortened median segment may be interpreted as apomorphic for Phryganistria. Just as in the closely related Phobaeticus, which can be regarded the adelphotaxon (Fig. 367), && possess a prominent, very similarly structured praeopercular organ, which is formed by a pair of spines or pointed lobes close to the posterior margin of abdominal sternum VII. As in Phobaeticus several morphological features of the insects are subject to strong intrageneric variation. These include: the size of the %% in relation to &&; length of the subgenital plate of &&; structure and armature of the meso- and metafemora of %% and the structure of the abdominal tergites of &&. %% are principally much more slender than && but may either be much shorter than the corresponding && or of almost equal length, reaching body lengths of up to 250 mm. The subgenital plate of && is either short and does not project over the apex of the anal segment, or is distinctly elongated, lanceolate and exceeds the anal segment by as much as the combined length of the three terminal abdominal segments. The abdominal tergites II–VI of && are usually parallel-sided and rectangular but in one case (Phryganistria virgea (Westwood, 1848)) have the lateral margins gently convex and rounded. The meso- and metafemora of %% are either slender and of the usual proportions for Pharnaciini, or remarkably broadened and armed with a prominent, dagger-like sub-apical spine on the two outer ventral carinae. The egg capsule is distinctly longer than wide, slightly laterally compressed and exhibits a faint keel on the dorsal and ventral surfaces as well as on the polar-area (Figs. 186–191). This keel is however much less developed than in Phobaeticus. The micropylar plate is rather large, covers more than 2/3 of the capsule length, and is of a simple shape, being elongate, tapered anteriorly and with a conspicuous median gap posteriorly. The general shape is similar to the plates seen in certain genera of Clitumnini (e.g. Cuniculina Brunner v. Wattenwyl, 1907 and Entoria Stål, 1875). Although Phryganistria shows striking similarity to certain taxa of Clitumnini, representatives are easily distinguished by the key features of Pharnaciini: the distinct armature of the extremities; much longer, filiform antennae, which are longer than the head, pro- and mesonotum combined; elongated antennomeres, which are much longer than wide; slender and more or less parallel-sided scapus and the presence of a distinct, stalked capitulum in the eggs (→ 5.2.4). D) Baculonistria gen. nov.: The evolutionary most derived forms of Pharnaciini are represented by Baculonistria gen. nov.. The genus contains only three known species all endemic to southern China and is at once distinguished from all other genera of the tribe by: the very prominently shortened, transverse median segment which is only about 1/8 the length of the metanotum; prominent sub-apical spine of the medioventral carina of the meso- and metafemora and shortened antennae of both sexes, which are distinctly shorter than the head, pro- and mesonotum combined. && furthermore differ by the presence of two small spines between



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the eyes and prominent but simple praeopercular organ, which is formed by a single transverse, semi-circular lobe at posterior margin of abdominal sternum VII. Eggs of Baculonistria gen. nov. are strongly globose and oval in cross-section, have the entire capsule surface covered with prominent, irregular impressions and possess a prominent opercular collar and large, roughly heart-shaped micropylar plate. The polar-area bears a rounded, keel-like extension dorsally and ventrally, but the capsule lacks a conspicuous longitudinal keel on the dorsal and ventral surfaces as in Phryganistria (and Phobaeticus). The short antennae, spines between the eyes of &&, very short median segment, simple praeopercular organ and prominent sub-apical spine of the medioventral carina of the meso and metafemora may be regarded as autapomorphies of Baculonistria gen. nov. These indicate Baculonistria gen. nov. to be the sister-group of the remainder of Pharnaciini (Pharnacia + Tirachoidea + Phobaeticus + Phryganistria, Fig. 367), but this certainly needs confirmation. Several of the typical and apomorphic features of Baculonistria gen. nov., e.g. the presence of a pair of spines between the eyes of &&, the shortened antennae and strongly shortened median segment, emphasize the close relation between Pharnaciini and Clitumnini here suggested. Although Baculonistria gen. nov. shows striking similarity to certain genera of Clitumnini (e.g. Entoria Stål, 1875), it is readily distinguished from members of this tribe by key features of Pharnaciini, e.g. the distinct armature of the extremities; antennae which are distinctly longer than the head and pronotum combined; slender scapus and presence of a distinct, stalked capitulum in the eggs (→ 6.1). The world’s longest extant insects Pharnaciini represents the bulge of the so-called Oriental “giant stick-insects” and contains several of the world’s longest known insects. Phobaeticus chani Bragg spec. nov. is the longest known extant insect, the longest specimen, the HT &, having a body length of 357.0 mm and an overall length with the fore legs stretched out straight of more than 566.0 mm (would be greater by 7–10 mm if the HT was complete). The second longest known species in means of body-length is Phobaeticus kirbyi Brunner v. Wattenwyl, the HT measuring 317.0 mm. A & of the same species from the Temburong District of Brunei (body length 283.0 mm) measured an overall length of 546.0 mm. Concerning to the overall length, the second longest species is Phobaeticus serratipes (Gray) with a & from Tasek Chini (Peninsular Malaysia) measuring 555.0 mm from the tip of the protarsi to the apex of the abdomen. The body length of this specimen however only was 278.0 mm and thus remarkably shorter than the greatest recorded body length for Ph. kirbyi (317.0 mm). There are several other species among Pharnaciini for which body lengths close to 30 cm have been recorded for &&, e.g. Phobaeticus magnus nom. nov. 315.0 mm, Phobaeticus redtenbacheri (Dohrn) 300.0 mm, Phryganistria heusii (Hennemann & Conle) 293.0 mm or Tirachoidea jianfenglingensis (Bi) 280.0 mm. Pharnaciini also contains the world’s longest known extant % insect, the HT % of Phryganistria heusii (Hennemann & Conle) measuring a body length of 250.5 mm and an overall length of over 470.0 mm with the legs stretched out. Similar body lengths are only achieved by a very few species elsewhere in the world, examples include: 1) Otocrania aurita (Burmeister, 1838; Phasmatidae s. l.: Cladomorphinae) from Brazil: 245.0 mm; 2) Bactrododema hippotaurum (Karsch, 1896; Diapheromeridae: Palophinae) from Malawi, East Africa: 263.0 mm; 3) Achrioptera punctipes punctipes (Audinet-Serville, 1838; Phasmatidae s. l.: Achriopterini) from Madagascar: 258.3 mm; 4) Nesiophasma plateni (Dohrn, 1910; Phasmatidae s. l.: Stephanacridini) from the Talaud Islands: 263.0 mm; 5) Macrophasma lyrata (Redtenbacher, 1908; Phasmatidae s. l.: Stephanacridini) from New Guinea: 268.0 mm; 6) Ctenomorpha gargantua Hasenpusch & Brock, 2006 (Phasmatinae: Phasmatini) from Queensland, Northern Australia: c. 300.0 mm; 7) Acrophylla titan (MacLeay, 1826; Phasmatinae: Phasmatini) from Australia: 270.0 mm, and 8) Hermarchus pythonius (Westwood, 1859; Phasmatidae s. l.: Stephanacridini) from Fiji: 242.0 mm.. Certainly, up to date only fractions of the actually existing Pharnaciini are discovered and regularly new species of these strikingly large insects are found. Thus, still several new species or perhaps even genera


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belonging to Pharnaciini are expected to be discovered in the future, once further research and extensive new collections are carried out in so far unprospected parts of the Oriental Region (e.g. wide parts of Kalimantan (Borneo), Sumatra, northern India, northern Myanmar, China, the Philippines or certain island faunas). But even fairly well studied regions still reveal new taxa.

8.1 Biogeography Pharnaciini are a typical faunal element of the Orientalis and are the largest and perhaps most striking representatives of the phasmid fauna of that region, representing the large bulge of the Oriental so-called giant stick-insects. Most of the taxa are distributed throughout Sundaland (Peninsular Malaysia, Sumatra, Java, Borneo and Palawan) but representatives are also found in the Philippines, India, Sri Lanka, southern flanks of the Himalaya (Bhutan, Sikkim and Darjeeling), as well as in the mountainous and low-land regions of Myanmar, Thailand, Kampuchea and Vietnam, and as far north as southern and eastern China. The Oriental Region comprises continental Asia south of the Palearctic Region, the Large Sunda Islands (Sumatra, Java and Borneo), the Philippines, Sulawesi and the Lesser Sunda Islands as far east as Timor. The eastern regions of the Orientalis form a zone of transition to the Australian Region, the Wallacea, which is in the west bounded by Wallace’s line. The Oriental and Australian Regions are separated by Weber’s line, which marks the western boundary of the Sahul shelf (see Briggs, 1987). The northern regions of continental Asia as well as Japan are included in the Paleartic Region which comprises all portions north of a line running along the Himalaya, through central China and between Japan and the Ryukyu Islands. On the basis of its geological history and physiogeography the Oriental Region is split into several geographical subregions or units some of which are still controversely defined by different zoologists (see Inger, 1999). Some of these subregions are helpful for understanding the distributional pattern of Pharnaciini and can be summarized as follows (see Inger, 1999): a) India and Sri Lanka (except the Southern Himalayan Flanks of India); b) the Himalaya and northeastern montane regions comprising Nepal, Bhutan, Sikkim, Darjeeling, Pradesh, Ladakh, Kashmir, Manipur, Nagaland and the mountainous regions of northern Myanmar, Thailand and Vietnam as well as soutern China; c) the Southeast Asian lowlands which comprise the central valleys of Myanmar, Thailand, Kampuchea as well as southern and coastal Vietnam; d) Sundaland; e) Sulawesi; f) the Philippine Islands excluding the Palawan chain; g) the Lesser Sunda Islands from Bali on the west to Timor on the east. Sundaland or former “Malaya” (see Mell, 1930; Johnson, 1964), the main distributional area of Pharnaciini, is one of those subregions which are still controversely defined, mainly due to Peninsular Malaysia shows strong Burmese elements among its fauna (see Eliot, 1992). Followingly, some zoologists regard Peninsular Malaysia as part of Sundaland while others regard it as a separate subregion, which additionally comprises the peninsular regions of Myanmar and Thailand. Herein, Sundaland is defined to cover the zoogeographical unit of the land masses on the Sunda shelf, that is: Peninsular Malaysia south of the Isthmus of Krah (or south of Latitude 7–10°N depending on authors), including the islands Sumatra, Java, Borneo, Bali, Balabac, the Palawan chain, the Calamaines and Cuyos Islands, all the smaller intervening islands, as well as the islands off the west coast of Sumatra (Simeuluë, Nias, the Batu Islands, Kepulauan, the Mentawei Islands and Engano) and the Christmas and Cocos-Keeling Islands (see Eliot, 1992; Nässig & Treadaway, 1998: 231). Within the defined area the sea is quite shallow but depth increases rapidly outside. The eastern boundary of Sundaland is formed by Wallace’s line and the lands east of it, e.g. the Lesser Sunda Islands from Lombok to Timor, Sulawesi and the Moluccas as well as the Philippines (except Palawan and the very eastern islands of the Sulu Archipelago). Due to they are not situated on the Sunda shelf these islands do not belong to Sundaland (see Johnson, 1964; Vane-Wright, 1990) and are all included in the Wallacea. Some background information on the geology of Sundaland may help to understand the distributional pat-



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tern of the taxa and resulting taxonomic arrangement established in the present work. It is generally believed that as late as the Pleistocene the glaciation caused a decrease of the world-wide sea-level and developed land or island chain bridges between the Asian continent and the Large Sunda Islands (see Briggs, 1987). During this time the present Peninsular Malaysia, the Large Sunda Islands, Borneo and almost certainly Palawan and Balabac were united to form a large more or less united land mass which received the term Sundaland (see Mell, 1930, Johnson, 1964). These land bridges are presumed to have allowed an interchange of the faunas of the Asian continent and Sundaland, although the position regarding the land connection between the mainland of Asia and Sundaland is fairly obscure (see Cox & Moore, 1992). The present fragmentation was brought about by the conditions prevailing in the last Ice Age, since the world-wide post-glacial increase of the sealevel large areas were submerged which caused an isolation of the islands. Java is believed to have been the first major island to be isolated from the rest of Sundaland and exhibits a less rich but stronger differentiated fauna from that of Peninsular Malaysia, Sumatra or Borneo, which together are also termed Neomalaya (see Moulton, 1915; Nässig & Treadaway, 1998). The small islands off the west coast of Sumatra (Simeuluë, Nias and Engano) are not on the Sunda Shelf and the Batu and Mentawei Islands are separated from the Sumatran mainland by deep channels and possess quite some degree of endemicity, why they are often termed Paramalaya (Toxopeus, 1926). In general however they should best be included into Sundaland (Nässig & Treadaway, 1998). The subfamily Clitumninae has predominantly radiated in the Oriental Region but exhibits a wide dispersal in the whole Indo-Malayan Archipelago. Diversification and specialization, especially on the Asian continent, becomes obvious in the very speciose and derived Clitumnini. This tribe only has a few representatives in the Wallacea but has dispersed as far north as to the southern parts of the Palaearctic Region (China, Korea and Japan) (→ Figs. 1 & 82). The rather small tribe Medaurini trib. nov. in contrast is restricted to continental SE-Asia (southern Indo-Chinese subregion, or Northeastern Montane Region according to Inger, 1999) and lacks any representatives in Sundaland. The more basal Pharnaciini has the centre of its dispersal in Sundaland and settled that complete subregion of the Orientalis. Representatives are furthermore also present in India and Sri Lanka as well as the Philippines, but have radiated not any further northward than NE-India and S-China. The highest densities of known species are found in the biogeographic subdivision termed Neomalaya: Borneo (8 species), Sumatra and surrounding islands (7 species) and the Malayan Peninsula (7 species). The genera Pharnacia Stål, 1877 and Tirachoidea Brunner v. Wattenwyl, 1893 are here recognized as the evolutionary most basal forms amongst Pharnaciini (→ 8.) both showing wide distributional patterns. A great majority of the known species is restricted to Sundaland but single representatives are found as far southeast as Java and Lombok, and as far northeast as the Philippine Islands (the latter islands belong to Wallacea), but both genera are missing in mainland India and Sri Lanka. Pharnacia is restricted to Sundaland but has two species in the Philippines and one in Palawan (→ Fig. 85), whereas Tirachoidea is absent in both Palawan and the Philippine Islands (→ Fig. 86). On the Asian mainland however the distribution of Tirachoidea extends as far north as NE-India, Bangladesh, Myanmar, South China, Thailand and Vietnam (southern Indo-Chinese subregion, or Northeastern Montane Region according to Inger, 1999). The presence of species of both genera in Java and continental Asia possibly indicates Pharnacia and Tirachoidea have dispersed as early as during the Pleistocene when the glaciation caused a decrease of the world-wide sea-level and developed land or island chain bridges between the Asian continent and the Sunda Islands (see Briggs, 1987). The climate divide between the perhumid equatorial tropical climate and the monsoonal (seasonal tropical to subtropical) climate in the northern parts of Peninsular Malaysia and southern Thailand, which is the present natural northern borderline of Sundaland (see Nässig & Treadaway, 1998), does not seem to have been a barrier at least for Tirachoidea. Phobaeticus Brunner v. Wattenwyl, 1907 as well is widely distributed throughout Sundaland and the southern portions of continental Asia (→ Fig. 83). It is however missing in Java, the first island believed to be


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isolated from the rest of Sundaland (see Moulton, 1915; Nässig & Treadaway, 1998), but also represented in the Philippines and Palawan (Wallacea), the southern Indo-Chinese subregion (Thailand, Myanmar, Laos & Kampuchea) and as far west as southern India and Sri Lanka (Indian subregion). No species have so far been recorded from the very eastern regions of the Indo-Chinese subregion (Vietnam). The highest densities of known species are found in Borneo (6 species), India (4 species) and Sumatra (4 species). Possible explanations for the distribution of Phobaeticus as presently seen may be similar to those mentioned for Tirachoidea above. Phryganistria Stål, 1875 contains rather derived representatives of Pharnaciini (→ 8.) and is distributed more northward than either Pharnacia, Tirachoidea or Phobaeticus, being a typical faunal element of the Indo-Chinese subregion and northeastern portions of the Indian subregion. It is missing in Sundaland, the Philippines, Sri Lanka or mainland India but commonly found in the highlands and valleys of northern Vietnam, Laos, Myanmar, Bangladesh, the southern flanks of the Himalaya (Bhutan, Darjeeling and Sikkim), NEIndia and as far north as the Guangxi Province of S-China. (→ Fig. 85). The most derived Pharnaciini are represented by Baculonistria gen. nov. (→ 8.), which is known from only three species that appear to be endemic to regions along the Yangtze River in southern and eastern China (Chongqing municipality, Henan Province & Anhui Province). The latter two provinces are attributed to the Palearctic Region. This genus contains the northernmost distributed representatives of Pharnaciini, which are the only species of the entire tribe known to occur in the Palaearctic Region. The presence of a separate genus in this particular region is supported by the proportionally high degree of endemism in the central mountainous regions of China. The eastern regions of China (e.g. Anhui Province) is a transitional zone between the faunas of both realms (Wu & Yuan, 1997: 100), which furthermore serves to explain the occurence of Pharnaciini as far north on the Asian mainland. The large distributional pattern of Pharnaciini may be explained by their high mobility and ability of dispersal. During the glaciation periods the sea-level decreased and increased with land or island chain connections developing or being submerged. Species inhabiting swamp- or lowland forests were principally able to disperse from one present-day land to the next during the times of low sea-level, while species inhabiting mountainous regions most likely were not, except for are very mobile e.g. well flying species (Nässig & Treadaway, 1998). As most Pharnaciini are inhabitants of lowland or at best hilly regions, and do not show any particular specialization to mountainous biotopes, they perhaps were principally in advantage to disperse more rapidly and widely in the Oriental Region. As a result, Pharnaciini can be regarded a typical faunistic element of the Oriental Region which derives to a quite early settlement most presumably during the Miocene via Australia, Wallacea and the southern Sunda Islands. They represent the largest and most striking Phasmatodea in that region and principally show a distribution which took (or still takes) place from the southern to the more northern parts of the Oriental Region towards the Palaearctic Region. The distributional pattern shows corellation to the presumed evolution and relationships of Pharnaciini and supports the allotment and hypotheses concerning to its position in the Phasmatidae s. l. suggested in the present work (→ 8. & Fig. 367). The following is a summary which lists all known species known from certain countries / islands / geographical regions. The parentheses state the number of recorded species. Doubtful or erroneous records are included in parentheses at the end of each section. China [7]: Baculonistria alba (Chen & He, 1990); Baculonistria longicornis (Bi & Wang, 1998); Baculonistria yuexiensis (Chen & He, 1993); Phryganistria guangxiensis Chen & He, 2007; Phryganistria longzhouensis Chen & He, 2007 Phryganistria grandis Rehn, 1906 (?); Tirachoidea jianfenglingensis (Bi, 1994). Sri Lanka [2]:



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Phobaeticus hypharpax (Westwood, 1859); Phobaeticus lobulatus (Carl, 1913). India [6]: Phobaeticus ingens (Redtenbacher, 1908); Phobaeticus annamallayanus (Wood-Mason, 1877); Phobaeticus sinetyi Brunner v. Wattenwyl, 1907; Phryganistria virgea (Westwood, 1848); Phobaeticus hypharpax (Westwood, 1859); Tirachoidea westwoodii (Wood-Mason, 1875). Myanmar [6]: Phobaeticus ingens (Redtenbacher, 1908); Phobaeticus magnus nom. nov.; Phryganistria fruhstorferi (Brunner v. Wattenwyl, 1907); Phryganistria grandis Rehn, 1906; Tirachoidea cantori (Westwood, 1859); Tirachoidea westwoodii (Wood-Mason, 1875). [Tirachoidea biceps (Redtenbacher, 1908) & Phobaeticus serratipes (Gray, 1839) – in error ?]. Vietnam [4]: Phryganistria grandis Rehn, 1906; Phryganistria heusii (Hennemann & Conle, 1997); Tirachoidea jianfenglingensis (Bi, 1994); Tirachoidea siamensis spec. nov.. Thailand [4]: Phobaeticus magnus nom. nov.; Tirachoidea cantori (Westwood, 1859); Tirachoidea jianfenglingensis (Bi, 1994); Tirachoidea siamensis spec. nov.. Peninsular Malaysia & Singapore [7]: Pharnacia sumatrana (Brunner v. Wattenwyl, 1907); Pharnacia tirachus (Westwood, 1859); Phobaeticus (?) incertus Brunner v. Wattenwyl, 1907; Phobaeticus pinnipes (Redtenbacher, 1908); Phobaeticus serratipes (Gray, 1835); Tirachoidea biceps (Redtenbacher, 1908); Tirachoidea cantori (Westwood, 1859). [Phryganistria virgea (Westwood, 1848) – Pulau Penang, in error ?] Sumatra and surrounding islands [7]: Pharnacia sumatrana (Brunner v. Wattenwyl, 1907); Phobaeticus (?) incertus Brunner v. Wattenwyl, 1907; Phobaeticus mucrospinosus spec. nov.; Phobaeticus rex (Günther, 1928); Phobaeticus serratipes (Gray, 1835); Phobaeticus sobrinus Brunner v. Wattenwyl, 1907; Tirachoidea biceps (Redtenbacher, 1908). Java [5]: Pharnacia sumatrana (Brunner v. Wattenwyl, 1907); Pharnacia heros Redtenbacher, 1908; Pharnacia tirachus (Westwood, 1859); Tirachoidea biceps (Redtenbacher, 1908); Tirachoidea inversa (Brunner v. Wattenwyl, 1907). [Phryganistria virgea (Westwood, 1848) – in error] Borneo [8]: Pharnacia borneensis spec. nov.; Phobaeticus chani Bragg spec. nov.; Phobaeticus foliatus (Bragg, 1995); Phobaeticus kirbyi Brunner v. Wattenwyl, 1907; Phobaeticus mjoebergi (Günther, 1935); Phobaeticus redtenbacheri (Dohrn, 1910); Phobaeticus rex (Günther, 1928); Tirachoidea herberti spec. nov. Palawan [2]: Pharnacia palawanica spec. nov.; Phobaeticus palawanensis spec. nov.. Philippines [4]: Pharnacia kalag Zompro, 2005; Pharnacia ponderosa Stål, 1877; Phobaeticus lumawigi Brock, 1997;


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Phobaeticus philippinicus (Hennemann & Conle, 1997). 8.2 Habitats & ecology The roughly 700 examined specimens were taken in a wide range of habitats, such as primary forest, secondary forest, montane and lowland dipterocarp forest, swamp forest, agricultural or even urban areas such as public parks, private gardens or small areas of secondary forest towards the outskirts of cities. The elevations of recorded localities range from sea-level to over 1500 m, but only a few species are found in altitudes above 1000 m. Examples include Phobaeticus redtenbacheri (Dohrn, 1910) from Sabah (Mt. Kinabalu), Phobaeticus philippinicus (Hennemann & Conle, 1997) from the Philippines (Mindoro Island, Mt. Halcon) or Tirachoidea biceps (Redtenbacher, 1908) from Java (Mt. Tengger). All three species are however also found at much lower elevations. At elevations above 2000 m members of the tribe appear to be completely absent and followingly, Pharnaciini can predominantly be regarded as lowland forms which do not show any significant specialization to mountainous biotopes. This advanced a high mobility and great ability of dispersal during the glaciation, which de- and increased the sea-level and developed land or island chain connections between the different islands and may explain the wide distribution of the tribe in Southeast Asia (→ 8.1.). In general these insects occur in heights between 1–6 metres off the ground, usually on low growing shrubs or small trees. Nymphs may also be found closer to the ground on low-growing vegetation. Principally the possibility to find specimens more close to ground-level increases in secondary forest or mountainous dipterocarp forest as the trees are typically smaller than in primary forest and the undergrowth is denser. In primary forest specimens tend to occur much higher, which is due to the high canopy blocks most of the sunlight why there is little vegetation at ground-level. Natural foodplants include several large-leaved jungle trees (see Tay & Seow-Choen, 1996, Lit & Eusebio, 2008) e.g. Canarium ovatum (Buseraceae), Macaranga conifera, Macaranga trilobata (Euphorbiaceae), Terminalia catappa (Combretaceae), Anthocephalus chinensis (Rubiaceae), Eucalyptus spp. (Myrtaceae), Tamarindus indica (Fagaceae), Terminalia microcarpa (Combretaceae), Psidium guajava (Myrtaceae) and Mangifera indica (Anacardiaceae) or low growing plants like Rubus moluccanus (Rosaceae). When no insectizides are applicated, the insects may also occur on single, cultivated guava (Psidium guajava, Myrtaceae) or mango-trees (Mangifera indica, Anacardiaceae) in private gardens or plantations. Alternative foodplants usually being accepted in captivity in Europe include bramble (Rubus fruticosus, Rosaceae), roses (Rosa spp., Rosaceae), oaks (Quercus robur & Q. ilex, Fagaceae), beech (Fagus sylvaticus, Fagaceae), eucalyptus (Eucalyptus spp., Myrtaceae), Salal (Gaultheria shallon, Ericaceae) and Japanese cherry (Prunus serrulata, Rosaceae). Principally most species of Pharnaciini are not considerably abundant although certain species are frequently found in smaller numbers. Almost none have so far been mentioned to occur as pests due to defoliation on jungle trees or in agricultural areas. Only Baculonistria alba (Chen & He, 1990) has been reported to have caused considerable damage (Chen & He, 1993; Chen, 1994; Hennemann, Conle & Zhang, 2008: 59) on Vernicia fordii (Euphorbiaceae), Cupressus funebris (Cupressaceae), Platycarya strobilacea, P. orientalis (Juglandaceae), Sapium sebiferum (Euphorbiaceae), Cyclobalanopsis sp. (Fagaceae), Salix cheilophila (Salicaceae), Zea mays (Poaceae) and Solanum tuberosum (Solanacea) e.g. in the Sichuan Province (now Chongqing) of China during 1986 (Cai & Liu, 1993: 471; reported as Phobaeticus sichuanensis Cai & Liu, 1993). Furthermore, these authors reported 20 square kilometres of Cupressus funebris to have been considerably defoliated by Baculonistria alba (Chen & He) in the same region (Fig. 414). Some species are quite abundant in certain localities, e.g. Phobaeticus kirbyi Brunner v. Wattenwyl, 1907 in Niah National Park (NE-Sarawak). Bragg (1995a: 27) reported this species to be exceptionally common along the path leading from the Park Head Quarters to the Great Cave in Niah National Park, with more than 40 adult specimens and numerous mating pairs encountered in only one night in August 1992. The authors encountered similar numbers in the same locality in August 1996 but the number of specimens in this biotope appears to be not large enough for



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causing any considerable damage on the jungle foliage. Pharnacia ponderosa Stål, 1877a is commonly encountered throughout the Philippines and is widely distributed on these islands. On certain islands, e.g. Panay or Marinduque it is very common but again the numbers of specimens are not large enough to cause any significant damage.

9 Taxonomic summary 9.1 New taxa and new names 1. Gigantophasmatini trib. nov. (subafmily Tropidoderinae) 2. Medaurini trib. nov. (subfamily Clitumninae) 3. Neohiraseini trib. nov. (subfamily Lonchodinae) 4. Baculonistria gen. nov. 5. Pharnacia borneensis spec. nov. 6. Pharnacia palawanica spec. nov. 7. Phobaeticus chani Bragg spec. nov. 8. Phobaeticus magnus nom. nov. 9. Phobaeticus mucrospinosus spec. nov. 10. Phobaeticus palawanensis spec. nov. 11. Tirachoidea herberti spec. nov. 12. Tirachoidea siamensis spec. nov.

[Loyalty Islands] [Continental SE-Asia] [Continental SE-Asia] [Central China] [N-Borneo] [Palawan: Brookes Point] [NE-Borneo: Sabah] [Thailand, Laos & Myanmar] [Western Sumatra: Solok] [Palawan: Salakot Pass] [Northern Borneo: Brunei & Sarawak] [Thailand & S-Vietnam]

9.2 List of taxonomic changes 1. Achriopterini Günther, 1953 (Type genus: Achrioptera Coquerel, 1861), here removed from the subfamily Phasmatinae and Phasmatidae s. str. [→ Phasmatidae s. l.: Achriopterini, 4.7.1]. 2. Andropromachus Carl, 1913: 48 (Type species: Andropromachus scutatus Carl, 1913: 49, pl. 1: 1 & 3), here transferred to Lonchodinae: Neohiraseini trib. nov. [→ 5.4.2]. 3. Anophelepis Westwood, 1859: 68 (Type species: Anophelepis telesphorus Westwood, 1859: 69, pl. 8: 3, 7, 7a) here removed from the subfamily “Platycraninae” and transferred to Phasmatinae: Acanthomimini [→ 4.3.2]. 4. Arphax Stål, 1875: 37 (Type species: Bacillus australis Charpentier, 1845, pl. 57), here removed from Phasmatinae: Acanthoxylini and provisionally transferred to Phasmatinae: Acanthomimini [→ 4.3.2]. 5. Bactridium grande Rehn, 1920: 242, pl. 10: 12, synonymised with Cladoxerus serratipes Gray, 1835 syn. nov. [Valid name = Phobaeticus serratipes (Gray, 1835), 6.3] 6. Baculum album Chen & He, 1990: 54, figs. 1a-c, here removed from Phobaeticus Brunner v. Wattenwyl, 1907 and selected as the type species of Baculonistria gen. nov. [Valid name = Baculonistria alba (Chen & He, 1990) comb. nov., 6.1] 7. Cladomimus Carl, 1915: 192 (Type species: Cladomimus griseus Carl, 1915: 192), here removed from the tribe Pharnaciini and transferred to Lonchodinae: Lonchodini [→ 5.4.1]. 8. Clitumnini Brunner v. Wattenwyl, 1893 (Type genus: Clitumnus Stål, 1875), here removed from Phasmatidae s. str. and transferred to the subfamily Clitumninae (Phasmatidae sensu lato) [→ 5.] 9. Clitumninae Brunner v. Wattenwyl, 1893 (Type genus: Clitumnus Stål, 1875), here introduced for a subfamily of Phasmatidae s. l. which contains the tribes Clitumnini and Pharnaciini [→ 5.]. 10. Clitumnus irregularis Brunner v. Wattenwyl , 1907: 195, synonymised with Phibalosoma tirachus West-


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wood, 1859 syn. nov. [Valid name = Pharnacia tirachus (Westwood, 1859) comb. nov., 6.2] 11. Cnipsomorpha Hennemann et al., 2008: 54 (Type species: Cnipsomorpha erinacea Hennemann, Conle, Zhang & Liu, 2007: 58), here removed from Pachymorphinae: Gratidiini and transferred to Clitumninae: Medaurini trib. nov. [→ 5.3]. 12. Diagoras Stål, 1877b: 66 (Type species: Diagoras ephialtes Stål, 1877b: 66), here removed from the tribe Pharnaciini and transferred to Phasmatidae sensu lato: Stephanacridini [→ 4.7.2] 13. Eucarcharus Brunner v. Wattenwyl , 1907: 185 (Type species: Lonchodes feruloides Westwood, 1859: 45, pl. 6: 5a, b), here removed from the tribe Pharnaciini and transferred to Phasmatidae sensu lato: Stephanacridini [→ 4.7.2]. 14. Eucarcharus inversus Brunner v. Wattenwyl , 1907: 186, here removed from Eucarcharus Brunner v. Wattenwyl, 1907 (→ Phasmatidae sensu lato: Stephanacridini) and transferred to the genus Tirachoidea Brunner v. Wattenwyl, 1893 [Valid name = Tirachoidea inversa (Brunner v. Wattenwyl, 1907) comb. nov., 6.5] 15. Eucarcharus lobulatus Carl, 1913: 12, here removed from Eucarcharus Brunner v. Wattenwyl, 1907 (Phasmatidae sensu lato: Stephanacridini) and transferred to the genus Phobaeticus Brunner v. Wattenwyl, 1907 [Valid name = Phobaeticus lobulatus (Carl, 1913) comb. nov., 6.3] 16. Extatosomatini Clark-Sellick, 1997 (Type genus Extatosoma Gray, 1833) here raised to subfamily level [→ Phasmatidae sensu stricto: Extatosomatinae, 4.5] 17. Eucarcharus rex Günther, 1928: 218, here automatically transferred to the genus Phobaeticus Brunner v. Wattenwyl, 1907 since being the type species of Lobophasma Günther, 1935, a junior synonym of Phobaeticus (syn. nov.) [Valid name = Phobaeticus rex (Günther, 1928) comb. nov., 6.3] 18. Gigantophasma Sharp, 1898: 87 (Type species: Gigantophasma bicolor Sharp, 1898: 87, pl. 7: 6), here removed from the tribe Pharnaciini and subfamily Phasmatinae, and selected as the type genus of the tribe Gigantophasmatini trib. nov. (Phasmatidae sensu stricto: Tropidoderinae) [→ 4.4.3]. 19. Gongylopus Brunner v. Wattenwyl, 1907: 233 (Type species: Gongylopus adiposus Brunner v. Wattenwyl, 1907: 233, pl. 9: 7), here removed from Pachymorphinae: Gratidiini and transferred to Clitumninae: Clitumnini [→ 5.2]. 20: Interphasma Chen & He, 2007: 328 (Type-species: Interphasma lushanense Chen & He, 2007: 329, figs. 297 a-e), here removed from Clitumnini and transferred to the tribe Medaurini trib. nov. [→ 5.3]. 21. Lanceocercata Bradler, 2001, here shown to be a synonym of Phasmatidae Gray, 1835 (sensu stricto) [→ 4.]. 22. Leiophasma Chen & He, 2007: 320 (Type-species: Leiophasma yunnanense Chen & He, 2007: 321, figs. 290 a-c), here synonymised with Paraleiophasma Chen & He, 2007: 322 syn. nov. [◊ addendum] 23. Leprocaulinus Uvarov, 1940: 176 (Type species: Leprocaulus alte-cornutus Redtenbacher, 1908: 473), here removed from the subfamily Necrosciinae and transferred to Lonchodinae: Lonchodini [→ 5.4.1]. 24. Lobophasma Günther, 1935b: 138 (Type species: Eucarcharus rex Günther, 1928: 218), here synonymised with Phobaeticus Brunner v. Wattenwyl , 1907 syn. nov. [→ 6.3]. 25. Lonchodinae Brunner v. Wattenwyl, 1893 (Type genus: Lonchodes Gray, 1835), here removed from the New World family Diapheromeridae and transferred to the Old World family Phasmatidae sensu lato [→ 5.4]. 26. Medaura Stål, 1875: 69 (Type species: Medaura brunneri Stål, 1875: 69), here removed from Clitumnini and transferred to the tribe Medaurini trib. nov. [→ 5.3]. 27. Medauroidea Zompro, 2000: 68 (Type species: Clitumnus extradentatus Brunner v. Wattenwyl, 1907: 193), here removed from Clitumnini and transferred to the tribe Medaurini trib. nov. [→ 5.3]. 28. Nearchus Redtenbacher, 1908: 448 (Type species: Nearchus maximus Redtenbacher, 1908: 448), here synonymised with Phobaeticus Brunner v. Wattenwyl , 1907 syn. nov. [→ 6.3]. 29. Nearchus foliatus Bragg, 1995b: 274, here transferred to the genus Phobaeticus Brunner v. Wattenwyl, 1907, since the original genus Nearchus Redtenbacher, 1908 is a junior synonym [Valid name = Phobaeti-



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cus foliatus (Bragg, 1995) comb. nov., 6.3] 30. Nearchus grubaueri Redtenbacher, 1908: 448, here synonymised with Phobaeticus incertus Brunner v. Wattenwyl, 1907 syn. nov. and removed from the tribe Pharnaciini [Systematic position uncertain but may belong in Nesiophasma Günther, 1934 (Phasmatidae sensu lato: Stephanacridini), → Nesiophasma incertus (Brunner v. Wattenwyl, 1907)?, 6.3] 31. Nearchus maximus Redtenbacher, 1908: 448, here automatically transferred to the genus Phobaeticus Brunner v. Wattenwyl, 1907 since it is the type species of Nearchus Redtenbacher, 1908, a junior synonym of Phobaeticus (syn. nov.). Renamed Phobaeticus magnus nom. nov. due to being preoccupied by Phobaeticus maximus (Bates, 1865), a junior synonym of Phobaeticus serratipes (Gray, 1835) syn. nov. [Valid name = Phobaeticus magnus nom. nov., 6.3] 32. Nearchus maximus mjöbergi Günther, 1935a: 10, here treated as a valid species and transferred to the genus Phobaeticus Brunner v. Wattenwyl , 1907 [Valid name = Phobaeticus mjoebergi (Günther, 1935) stat. nov., comb. nov., 6.3] 33. Nearchus redtenbacheri Dohrn, 1910: 409, here transferred to the genus Phobaeticus Brunner v. Wattenwyl, 1907, since the original genus Nearchus Redtenbacher, 1908 is a junior synonym [Valid name = Phobaeticus redtenbacheri (Dohrn) comb. nov., 6.3] 34. Neohirasea Rehn, 1904: 84 (Type species: Phasma (Acanthoderus) japonicum de Haan, 1842: 135, pl. 12: 4), here transferred to Lonchodinae: Neohiraseini trib. nov. [→ 5.4.1]. 35. Nesiophasma Günther, 1934c: 5 (Type species: Nesiophasma eremothocus Günther, 1934: 1934c: 6, fig. 1), here removed from the tribe Clitumnini and transferred to Phasmatidae sensu lato: Stephanacridini [→ 4.7.2]. 36. Parapachymorpha Brunner v. Wattenwyl, 1893: 95 (Type species: Parapachymorpha nigra 1893: 96, pl. 4: 35a & b), here removed from Pachymorphinae: Gratidiini and transferred to Clitumninae: Medaurini trib. nov. [→ 5.3]. 37. Pharnacia annamallayanum (Wood-Mason, 1877a: 161), here removed from Pharnacia Stål, 1877 and transferred to the genus Phobaeticus Brunner v. Wattenwyl, 1907 [Valid name: Phobaeticus annamallayanus (Wood-Mason, 1877) comb. nov.] 38. Pharnacia annulata Redtenbacher, 1908: 451, here removed from Phobaeticus Brunner v. Wattenwyl, 1907 and transferred to the genus Sadyattes Stål, 1875 (Phasmatidae sensu lato: Stephanacridini). [Valid name = Sadyattes annulatus (Redtenbacher, 1908) comb. nov., 4.7.2 and 6.3] 39. Pharnacia biceps Redtenbacher, 1908: 451, here removed from Pharnacia Stål, 1877 and transferred to the genus Tirachoidea Brunner v. Wattenwyl, 1893 [Valid name = Tirachoidea biceps (Redtenbacher) stat. rev., 6.5.] 40. Pharnacia chiniensis Seow-Choen, 1998c: 184, here synonymised with Pharnacia biceps Redtenbacher, 1908 syn. nov. [Valid name = Tirachoidea biceps (Redtenbacher), 6.5] 41. Pharnacia enganensis Redtenbacher, 1908: 451, here removed from Phobaeticus Brunner v. Wattenwyl, 1907 and transferred to the genus Sadyattes Stål, 1875 (Phasmatidae sensu lato: Stephanacridini) [Valid name = Sadyattes enganensis (Redtenbacher, 1908) comb. nov. , 4.7.2 and 6.3] 42. Pharnacia heros Redtenbacher, 1908: 453, here removed from Phobaeticus Brunner v. Wattenwyl, 1907 and re-transferred to the genus Pharnacia Stål, 1877 [Valid name = Pharnacia heros Redtenbacher, 1908 stat. rev., 6.2] 43. Pharnacia magdiwang Lit & Eusebio, 2008, here synonymised with Pharnacia ponderosa Stål, 1877 syn. nov. [Valid name: Pharnacia ponderosa Stål, 1877, 6.2] 44. Pharnacia nigricornis Redtenbacher, 1908: 452, here removed from Pharnacia Stål, 1877 and transferred to the genus Sadyattes Stål, 1875 (Phasmatidae sensu lato: Stephanacridini) [Valid name = Sadyattes nigricornis (Redtenbacher, 1908) comb. nov. , 4.7.2 and 6.3] 45. Pharnacia rigida Redtenbacher, 1908: 453, synonymised with Phobaeticus sumatranus Brunner v. Wat-


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tenwyl, 1907 syn. nov. [Valid name = Pharnacia sumatrana (Brunner v. Wattenwyl, 1907), 6.2] 46. Pharnacia semilunaris Redtenbacher, 1908: 452, here synonymised with Eucarcharus inversus Brunner v. Wattenwyl, 1907 syn. nov. [Valid name = Tirachoidea inversa (Brunner v. Wattenwyl, 1907) comb. nov., 6.5] 47. Pharnacia spectabilis Redtenbacher, 1908: 453, here synonymised with Phibalosoma hypharpax Westwood, 1859 syn. nov. [Valid name = Phobaeticus hypharpax (Westwood, 1859), 6.3] 48. Pharnaciini Günther, 1953 (Type genus: Pharnacia Stål, 1877), here removed from Phasmatidae sensu stricto and transferred to the subfamily Clitumninae (Phasmatidae sensu lato) [→ 5.] 49. Phasmatotaenia Navás, 1907: 10 (Type species: Taeniosoma sanchezi Bolivar, 1895: 31, fig. 1), here removed from the tribe Pharnaciini and transferred to Phasmatidae sensu lato: Stephanacridini [→ 4.7.2]. 50. Phenacocephalus Werner, 1930: 179 (Type species: Phenacocephalus coronatus Werner, 1930: 179), here removed from the subfamily Necrosciinae and transferred to Lonchodinae: Lonchodini [→ 5.4.1]. 51. Phibalosoma cantori Westwood, 1859: 74, pl. 37, fig. 1, 1a–b, pl. 38: 1, 1a–b, here removed from Pharnacia Stål, 1877 and automatically re-transferred to the re-established genus Tirachoidea Brunner v. Wattenwyl, 1893, since it is the type species of Tirachoidea [Valid name = Tirachoidea cantori (Westwood) stat. rev., 6.5]. 52. Phibalosoma maximum Bates, 1865: 341, here synonymised with Cladoxerus serratipes Gray, 1835 syn. nov. [Valid name = Phobaeticus serratipes (Gray, 1835), 6.3]. 53. Phibalosoma tirachus Westwood, 1859: 75, pl. 37: 3, here removed from Phobaeticus Brunner v. Wattenwyl, 1907 and re-transferred to the genus Pharnacia Stål, 1877 [Valid name = Pharnacia tirachus (Westwood, 1859) stat. rev., 6.3] 54. Phibalosoma westwoodii Wood-Mason, 1875: 216, here removed from Pharnacia Stål, 1877 and transferred to the genus Tirachoidea Brunner v. Wattenwyl, 1893 [Valid name = Tirachoidea westwoodii (Wood-Mason) comb. nov., 6.5]. 55. Phobaeticus beccarianus Brunner v. Wattenwyl, 1907: 185, here synonymised with Phobaeticus sobrinus Brunner v. Wattenwyl, 1907 syn. nov. [Valid name = Phobaeticus sobrinus Brunner v. Wattenwyl, 1907, 6.3] 56. Phobaeticus fruhstorferi Brunner v. Wattenwyl , 1907: 184, here removed from Phobaeticus Brunner v. Wattenwyl, 1907 and transferred to the genus Phryganistria Stål, 1875 [Valid name = Phryganistria fruhstorferi (Brunner v. Wattenwyl, 1907) comb. nov., 6.4] 57. Phobaeticus heusii Hennemann & Conle, 1997b: 505, figs. 1–5, here removed from Phobaeticus Brunner v. Wattenwyl, 1907 and transferred to the genus Phryganistria Stål, 1875 [Valid name = Phryganistria heusii (Hennemann & Conle, 1997) comb. nov., 6.4] 58. Phobaeticus lambirica Seow-Choen, 1998a: 40, fig. 1, synonymised with Eucarcharus rex Günther, 1928 syn. nov. [Valid name = Phobaeticus rex (Günther, 1928) comb. nov., 6.3] 59. Phobaeticus longicornis Bi & Wang, 1998: 10, figs. 5 & 6, here removed from Phobaeticus Brunner v. Wattenwyl, 1907 and transferred to Baculonistria gen. nov. [Valid name = Baculonistria longicornis (Bi & Wang, 1998), 6.1] 60. Phobaeticus sichuanensis Cai & Liu, 1993: 469, here synonymised with Baculum album Chen & He, 1990 syn. nov. [Valid name = Baculonistria alba (Chen, 1990), 6.1] 61. Phobaeticus yuexiensis Chen & He, 1993: 54, figs. 1 & 2, here removed from Phobaeticus Brunner v. Wattenwyl, 1907 and transferred to Baculonistria gen. nov. [Valid name = Baculonistria yuexiensis (Chen & He, 1993), 6.1] 62. Phryganistria Stål, 1875: 6 (Type species: Bacteria sarmentosa Westwood, 1848: 65, pl. 36: 1), here removed from the tribe Clitumnini and transferred to Pharnaciini [→ 6.4]. 63. Phryganistria grandis Rehn, 1906: 279, here removed from Phobaeticus Brunner v. Wattenwyl, 1907 and re-transferred to Phryganistria Stål, 1875 [Valid name = Phryganistria grandis Rehn, 1906 stat. rev., 6.4]



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64. Pseudocentema Chen, He & Li, 2002 : 106 (Type species: Pseudocentema bispinatum Chen & He (in: Chen, He & Li), 2002: 107, figs. 8a & b), here transferred to Lonchodinae: Neohiraseini trib. nov. [→ 5.4.2]. 65. Qiongphasma Chen, He & Li, 2002: 106 (Type species: Qiongphasma jianfengense Chen & He (in: Chen, He & Li), 2002: 106, figs7a & b), here transferred to Lonchodinae: Neohiraseini trib. nov. [→ 5.4.2]. 66. Sadyattes Stål, 1875: 44, 88 (Type species: Sadyattes borrii Stål, 1875: 88), here removed from the tribe Pharnaciini and transferred to Phasmatidae sensu lato: Stephanacridini [→ 4.7.2] 67. Spinohirasea Zompro, 2002b: 67 (Type species: Spinohirasea crassithorax Zompro, 2002b : 68, figs. 1–7 (= Menexenus bengalensis Brunner v. Wattenwyl, 1907), here transferred to Lonchodinae: Neohiraseini trib. nov. [→ 5.4.2]. 68. Stephanacridini Günther, 1953 (Type genus: Stephanacris Redtenbacher, 1908), here removed from the subfamily Phasmatinae and Phasmatidae sensu stricto [→ Phasmatidae sensu lato: Stephanacridini, 4.7.2]. 69. Tirachoidea Brunner v. Wattenwyl , 1893: 83 (Type species: Phibalosoma cantori Westwood, 1859), here removed from synonymy with Pharnacia Stål, 1877 and re-established as a valid genus (stat. rev.) [→ 6.5]. 70. Vasilissa Kirby, 1896: 468 (Type species: Vasilissa walkeri Kirby, 1896: 469), here removed from Phasmatidae: Acanthoxylini and provisionally transferred to Phasmatinae: Acanthomimini [→ 4.3.2].

9.3 Taxonomic catalogue of Pharnaciini Günther, 1953 Baculonistria gen. nov. Type species: Baculum album Chen & He, 1990: 54, by present designation.

Species included: 1. Baculonistria alba (Chen & He, 1990: 54) [Baculum]. = Phobaeticus sichuanensis (Cai & Liu, 1993: 469, figs. 1–4). syn. nov. 2. Baculonistria longicornis (Bi & Wang, 1998: 10, figs. 5–6) [Phobaeticus]. 3. Baculonistria yuexiensis (Chen & He, 1993: 54, figs. 1–2) [Phobaeticus]

Pharnacia Stål, 1877 Type species: Pharnacia ponderosa Stål, 1877a: 40, by subsequent designation of Kirby, 1904a: 359.

Species included: 1. Pharnacia borneensis spec. nov. 2. Pharnacia heros Redtenbacher, 1908: 453. 3. Pharnacia kalag Zompro, 2005: 73. 4. Pharnacia palawanica spec. nov. 5. Pharnacia ponderosa Stål, 1877a: 40. = Pharnacia magdiwang Lit & Eusebio, 2008: 117, figs, 2c, 2f, 3a, 4, 5. syn. nov. 6. Pharnacia sumatrana (Brunner v. Wattenwyl, 1907: 184) [Phobaeticus]. = Pharnacia rigida Redtenbacher, 1908: 453. syn. nov. 7. Pharnacia tirachus (Westwood, 1859: 75, pl. 37: 3) [Phibalosoma]. = Clitumnus irregularis Brunner v. Wattenwyl, 1907: 195. syn. nov.


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Phobaeticus Brunner v. Wattenwyl, 1907 Type species: Phobaeticus sobrinus Brunner v. Wattenwyl, 1907: 184, pl. 7: 1a & b, by subsequent designation of Brock, 1996: 28. = Baculolonga Hennemann & Conle, 1997a: 347. = Lobophasma Günther, 1935b: 138. syn. nov. = Nearchus Redtenbacher, 1908: 448. syn. nov.

Species included: 1. Phobaeticus annamallayanus (Wood-Mason, 1877a: 161) [Phibalosoma]. 2. Phobaeticus chani Bragg spec. nov. 3. Phobaeticus foliatus (Bragg, 1995b: 274) [Nearchus]. 4. Phobaeticus hypharpax (Westwood, 1859: 75, pl. 13: 6) [Phibalosoma]. = Pharnacia spectabilis Redtenbacher, 1908: 453. syn. nov. 5. Phobaeticus ingens (Redtenbacher, 1908: 453) [Pharnacia]. 6. Phobaeticus kirbyi Brunner v. Wattenwyl, 1907: 185. = Pharnacia sagitta Redtenbacher, 1908: 454, pl. 22: 2. = Pharnacia pilicornis Redtenbacher, 1908: 455. 7. Phobaeticus lobulatus (Carl, 1913: 12) [Eucarcharus]. 8. Phobaeticus lumawigi Brock, 1997: 43. 9. Phobaeticus magnus nom. nov. [= replacement name for Nearchus maximus Redtenbacher, 1908: 448]. 10. Phobaeticus mjoebergi (Günther, 1935a: 10) [Nearchus maximus mjöbergi]. 11. Phobaeticus mucrospinosus spec. nov. 12. Phobaeticus palawanensis spec. nov. 13. Phobaeticus philippinicus (Hennemann & Conle, 1997a: 349, figs. 10–16) [Baculolonga]. 14. Phobaticus pinnipes (Redtenbacher, 1908: 452) [Pharnacia]. 15. Phobaeticus redtenbacheri (Dohrn, 1910: 409) [Nearchus]. 16. Phobaeticus rex (Günther, 1928: 218) [Eucarcharus]. = Phobaeticus lambirica Seow-Choen, 1998a: 40, fig. 1. syn. nov. 17. Phobaeticus serratipes (Gray, 1835: 42) [Cladoxerus]. = Phasma (Bacteria) acanthopus Burmeister, 1839: 565. = Bactridium grande Rehn, 1920: 242, pl. 10: 12. syn. nov. = Phibalosoma maximum Bates, 1865: 341. syn. nov. 18. Phobaeticus sinetyi Brunner v. Wattenwyl, 1907: 184. 19. Phobaeticus sobrinus Brunner v. Wattenwyl, 1907: 184, pl. 7: 1. = Phobaeticus beccarianus Brunner v. Wattenwyl, 1907: 185. syn. nov. ? Phobaeticus incertus Brunner v. Wattenwyl, 1907: 185 [Possibly belonging in Nesiophasma Günther, 1934]. = Nearchus grubaueri Redtenbacher, 1908: 448. syn. nov.

Phryganistria Stål, 1875 Type species: Bacteria sarmentosa Westwood, 1848: 66, pl. 32: 1 & 2, by subsequent designation of Kirby, 1904a: 358.

Species included: 1. Phryganistria fruhstorferi (Brunner v. Wattenwyl, 1907: 184) [Phobaeticus].



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2. Phryganistria grandis Rehn, 1906: 279. 3. Phryganistria guangxiensis Chen & He, 2007: 337, figs. 308 a-b. [S-China (Guangxi)] 4. Phryganistria heusii (Hennemann & Conle, 1997b: 505 figs. 1–5) [Phobaeticus]. 5. Phryganistria longzhouensis Chen & He, 2007: 338, figs. 309 a-d. [S-China (Guangxi)] 6. Phryganistria virgea (Westwood, 1848: 65, pl. 32: 2) [Bacteria]. = Bacteria sarmentosa Westwood, 1848: 66, pl. 32: 1.

Tirachoidea Brunner v. Wattenwyl, 1893 stat. rev. Type species: Phibalosoma cantori Westwood, 1859: 74, pl. 37: 1, 1a–b & pl. 38: 1, 1a–b, by subsequent designation of Kirby, 1904a: 359.

Species included: 1. Tirachoidea biceps (Redtenbacher, 1908: 451) [Pharnacia]. = Pharnacia chiniensis Seow-Choen, 1998c: 184 syn. nov. 2. Tirachoidea cantori (Westwood, 1859: 74, pl. 37: 1 & 38: 1) [Phibalosoma]. 3. Tirachoidea herberti spec. nov. 4. Tirachoidea inversa (Brunner v. Wattenwyl, 1907: 186, pl. 8: 1) [Eucarcharus]. = Pharnacia semilunaris Redtenbacher, 1908: 452. syn. nov. 5. Tirachoidea jianfenglingensis (Bi, 1994: 10) [Pharnacia]. 6. Tirachoidea siamensis spec. nov. 7. Tirachoidea westwoodii (Wood-Mason, 1875: 216) [Phibalosoma].

10 Conclusion The tribe Pharnaciini Günther, 1953 was revised at the species level and shown to be polyphyletic. Splitting of the tribe and transferring formerly included genera to other tribes or subfamilies has shown the entire subfamily Phasmatinae to be not a natural group. Further studies on the systematic position and relationships of Pharnaciini have revealed a re-arrangement of the entire family Phasmatidae to be badly needed. This was already suggested by Bradler (2001) who established the supposedly monophyletic group “Lanceocercata” for predominantly Australian taxa, some of which belong to the subfamily Phasmatinae sensu Bradley & Galil, 1977. The monophyly of “Lanceocercata” was here confirmed but shown to be a synonym of what should actually be regarded the family Phasmatidae, here referred to as Phasmatidae sensu stricto. As already pointed out by Bradler (2001: 183) a monophyly of Phasmatidae sensu stricto would cause fundamental changes in the current allotment of the concerned subfamilies and tribes. Based on a detailed study of the genital exosceleton of the insects, but the genitalia in particular, and egg-morphology new hypotheses of the relationships within former Phasmatidae (= Phasmatidae sensu lato) and the newly defined Phasmatidae sensu stricto were presented and a new arrangement with keys to all concerned subfamilies and tribes provided. The Phasmatidae sensu stricto (= Lanceocercata) are characterised by several supposedly apomorphic features and contain six distinct subfamilies: Phasmatinae, Tropidoderinae, Extatosomatinae, Pachymorphinae, Xeroderinae and “Platycraninae”. The latter three subfamilies are most certainly polyphyletic and deserve further detailed clarification. The tribe Extatosomatini was removed from Tropidoderinae and raised to subfamily level. Phasmatinae contains the three tribes Phasmatini, Acanthomimini and Acanthoxylini. The three tribes Tropidoderini, Monandropterini and Gigantophasmatini trib. nov. together form the Tropidoderinae. The tribes Clitumnini and Pharnaciini were shown to be closely related to each other but are no members of Phasmatidae sensu


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stricto, genital features indicating close relationship to Lonchodinae. The new tribe Medaurini trib. nov. was established to contain those taxa of Clitumnini which have a well developed vomer in %% and ovoid eggs, as well as two genera formerly misplaced in Pachymorphinae: Gratiidini. All other subfamilies formerly attributed to Phasmatinae, this is Eurycanthinae and the New World Cladomorphinae, are not closely related and must provisionally be referred to as members of Phasmatidae sensu lato. The subfamily Lonchodinae is shown to be closely related to Clitumninae and hence transferred from Diapheromeridae to Phasmatidae sensu lato. Those genera of Lonchodinae formerly referred to as the “Neohirasea-complex” are transferred to the new tribe Neohiraseini trib. nov., which is doubtful to belong in Lonchodinae. Lonchodinae and Clitumninae are likely to form an own monophyletic family within Phasmatidae sensu lato, for which Lonchodidae is an available name. The position of Gratidiini (subfamily Pachymorphinae) remains uncertain. In combination with the more detailed study of the genitalia and eggs here presented, the new hypotheses concerning to the relationships within Phasmatidae sensu lato, diagnoses and keys to the concerned subfamilies and tribes are hoped to result in more comprehensive cladistic or genetic studies in the future. Certainly, further fundamental taxonomic changes are necessary to approach a natural classification of the Old World anareolate Phasmatodea. The Oriental tribe Pharnaciini contains five genera: Baculonistria gen. nov., Pharnacia Stål, 1877, Phobaeticus Brunner v. Wattenwyl, 1907 (= Baculolonga, = Lobophasma, = Nearchus), Phryganistria Stål, 1875 and Tirachoidea Brunner v. Wattenwyl, 1893. The latter genus was removed from synonym with Pharnacia. New diagnoses of all five genera, as well as keys, descriptions and illustrations were provided of all 40 known species. Several new synonymies were detected and seven new species described. Phobaeticus chani Bragg spec. nov. is the longest known extant insect in the world with an overall length of almost 60 cm in the &&. The Pharnaciini are a superb example to emphazise our still rather fractional knowledge of this fascinating order of insects world-wide. Although, they contain the world’s longest known insects, even up to date by far not all of the actually existing species are discovered and regularly new species of these strikingly large Phasmatodeans are found. The world’s longest extant insect, Phobaeticus chani Bragg spec. nov., is described herein although Borneo is amongst the more sufficiently studied areas of Southeast Asia. With certainty, several new species or perhaps even new genera of Pharnaciini are awaiting discovery and description, especially once collections are made in so far lesser prospected regions such as wide parts of Kalimantan (Borneo), Sumatra, the Philippines or northern India. In some cases the authors are aware of single specimens of undescribed taxa, which it is not yet appropriate to describe. Several species are still only known from a single sex or the original type-specimen(s) and the eggs of 19 species have not yet become known. Future taxonomic studies should therefore not only aim the description of new taxa but also the recognition of so far unknown sexes, eggs or older poorly known taxa in the more comprehensive representation now available. It is hoped, that this work will be a prolific basis for future studies on these fascinating insect-giants and a useful tool for the identification of taxa. Furthermore, the hypotheses and re-arrangement of the family Phasmatidae Gray, 1835 here presented are hoped to be helpful for future studies on the phylogenetics of Old World anareolate Phasmatodea.

11 Acknowledgements The authors would like to express their thanks to everybody who supported the present work. Thanks are due to the curators and staff of the following museums and institutions for access to the collections, providing information and the loan of specimens: Dr. Christiane Amedegnato, Simon Poulain & Emmanuel Delfosse (MNHN, Paris); Dr. Ulrike Aspöck & late Dr. A. P. Kaltenbach (NHMW, Vienna), Dr. Martin Brancucci, Dr. D. Burckhardt & Armin Coray (NHMB, Basel), Dr. Jacques Cools (ISNB, Bruxelles),



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Dr. Mauro Daccordi & Dr. P. L. Scaramozzino (MRSN, Torino), Dr. Andrei Gorokhov (ZMAS, St. Petersburg); Dr. M. Dohrn (MLUH, Halle), Dr. Roland Emmrich (SMTD, Dresden), Dr. Isabel Izquierdo & Mercedes París (MNCN, Madrid); Dr. Dimitri Logunov (MUME), Dr. George Beccaloni & Judith Marshall (BMNH, London), Dr. M. Müller (ETHZ, Zürich), Dr. George McGavin & Dr. Darren Mann (OXUM, Oxford), Dr. Wolfgang A. Nässig (SMFM, Frankfurt a. M.), Dr. Michael Ohl, Dr. Kurt K. Günther & Mrs. Isolde Dorandt (MNHU, Berlin), Dr. Roberto Poggi (MCSN, Genova), Dr. Wolfgang Schawaller (SMNS, Stuttgart), Dr. Wolfgang Schneider (HLDH, Darmstadt), Prof. Klaus Schönitzer & Mrs. Tanja Kothe (ZSMC, Munich), Dr. P. Schwendinger, Prof. Volker Mahnert & Dr. Bernd Hauser (MHNG, Genève), Dr. M. S. Shishodia (NZSI, Calcutta), Prof. Hans Strümpel (ZMUH, Hamburg), Dr. Jan van Tol, Dr. Caroline Pepermans & Rob de Vries (RMNH, Leiden), and Dr. Daniel Otte, Dr. Jason Weintraub & Dr. John Gelhaus (ANSP, Philadelphia). Thanks are due to the following persons for loaning specimens from their private collections, providing culture material, photos, literature or other helpful information and the good contact during the past fifteen years: Dr. John T. Clark-Sellick (Kettering, England), Paul D. Brock (Slough, England), Peter Heusi (Uster, Switzerland), Heinz van Herwaarden (Netherlands), Allan J. E. Harman (Essex, England), Kristien Rabaey & Rob Simoens (Veurne, Belgium), Arnaud and Christoph Bauduin (Flers en Escrebieux, France) and Thorsten Thron (Otzberg, Germany). Special thanks are due to Dr. Phil E. Bragg (Nottinghamshire, England) for carefully reading sections of the manuscript, making constructive comments, loaning specimens from his collection, taking photos as well as providing data and measurements. Furthermore, Phil Bragg has gratfully agreed to include his description of Phobaeticus chani spec. nov., the world’s longest known extant insect, in the present work. Mel Herbert and his family (Hütschenhausen, Germany) shall additionally be thanked for accomodation and assistance during night-time collecting in Brunei and loaning specimens from his collection. With his vast local knowledge Dr. Francis Seow-Choen (Singapore) provided assistance during night-time collecting in Singapore and generously gave away specimens from his collection. Michael K. P. Yeh (Ipoh, Malaysia) kindly assisted during day-time collecting in West Malaysia and supplied preserved and live specimens as well as eggs of local species. Lerdchai Kwantalae (Nong-Khai, Thailand) provided information on the natural habitats and foodplants as well as photos and live eggs of Phobaeticus magnus. Weiwei Zhang (Chongqing, China) kindly presented specimens of Baculonistria alba as well as information on the natural habitats of this species, Chinese literature and translations of Chinese descriptions. Mr. Xiang Guowei (Chongqing Forestry Bureau, China) generously allowed to use a photo documenting the damage caused by Baculonistria alba in the Yunyang County of Chongqing in 2005. Tet Fatt Wong (Tanah Rata, Malaysia), Ismael O. Lumawig (Luzon, Philippines), Noel Mohagan (Mindoro, Philippines), Sabine Steinke and Hilmar Lehmann (both Germany) supplied specimens. N. Cliquennois (Madagascar) shall be thanked for constructive discussion on Monandropterini and Acanthomimini, and Wim, Jacques and Rita Potvin (St. Pieters-Leeuw, Belgium) generously rendered hospitality during a visit to ISNB. Joachim Bresseel (Meise, Belgium) kindly provided photos of live Phobaeticus magnus, Rob Krijns (Maastricht, Netherlands) provided photos of a live male Pharnacia kalag and the mating habits of various species. Rainer Galunder (Nümbrecht, Germany) provided photos, specimens and culture stock of Baculonistria alba and Mieke Duytschaever (Belgium) provided a picture of the copulation habit of Extatosoma tiaratum. Gisela Hennemann (Freinsheim, Germany) and Thorsten O. Seifert (Grünstadt, Germany) kindly took care for the first author’s live insects when on expedition or visiting museum collections. Marina Friede (Bolsterlang, Germany) spent much time on digitalizing literature and photos and looked after the second author’s live insects when on occasion. The anonymous referees shall be thanked for their helpful and constructive comments on the manuscript. Finally we want to thank our parents Gisela & Hans-Peter Hennemann (Freinsheim & Altwarmbüchen, Germany) and Xenia & Matthäus Conle (Obermühlegg, Germany) to whom this work is dedicated. It would have never been possible to realize it without their patience, understanding for the author’s “strange hobby”and support in many ways.


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PLATE I. Eggs [scale = 1 mm] 145–146. Pharnacia ponderosa Stål, 1877, 145. dorsal view, 146. lateral view 147–148. Pharnacia kalag Zompro, 2005, 147. dorsal view, 148. lateral view 149–150. Pharnacia palawanica spec. nov., 149. dorsal view, 150. lateral view 151–152. Pharnacia sumatrana (Brunner v. Wattenwyl , 1907), 151. dorsal view, 152. lateral view 153–154. Pharnacia borneensis spec. nov., 153. dorsal view, 154. lateral view 155–156. Pharnacia tirachus (Westwood, 1859), 1877, 155. dorsal view, 156. lateral view 157–158. Tirachoidea cantori (Westwood, 1859), 157. dorsal view, 158. lateral view 159–160. Tirachoidea jianfenglingensis (Bi, 1994), 159. dorsal view, 160. lateral view



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PLATE II. Eggs [scale = 1 mm] 161–162. Tirachoidea siamensis spec. nov., 161. dorsal view, 162. lateral view 163–164. Tirachoidea herberti spec. nov., 163. dorsal view, 164. lateral view 165–166. Tirachoidea biceps (Redtenbacher, 1908), 165. dorsal view, 166. lateral view 167–168. Phobaeticus (?) incertus Brunner v. Wattenwyl , 1907, 167. dorsal view, 168. lateral view 169–170. Phobaeticus serratipes (Gray, 1835), 169. dorsal view, 170. lateral view 171–172. Phobaeticus kirbyi Brunner v. Wattenwyl , 1907, 171. dorsal view, 172. lateral view 173–174. Phobaeticus redtenbacheri (Dohrn, 1910), 173. dorsal view, 174. lateral view 175–176. Phobaeticus philippinicus (Hennemann & Conle, 1997), 175. dorsal view, 176. lateral view


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PLATE III. Eggs [scale = 1 mm] 177–178. Phobaeticus hypharpax (Westwood, 1859), 177. dorsal view, 178. lateral view 179–180. Phobaeticus sinetyi Brunner v. Wattenwyl , 1907, 179. dorsal view, 180. lateral view 181–183. Phobaeticus chani Bragg spec. nov., 181. dorsal view, 182. lateral view, 183. caudal view of operculum 184–185. Phobaeticus magnus nom. nov., 184. dorsal view, 185. lateral view 186–187. Phryganistria virgea (Westwood, 1848), 186. dorsal view, 187. lateral view 188–189. Phryganistria grandis Rehn, 1906, 188. dorsal view, 189. lateral view 190–191. Phryganistria heusii (Hennemann & Conle, 1997), 190. dorsal view, 191. lateral view 192–193. Baculonistria gen. nov. alba (Chen & He, 1990), 192. dorsal view, 193. lateral view REVISION OF PHARNACIINI


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PLATE IV. [scale = 10 mm] 194. Pharnacia ponderosa Stål, 1877, & apex of abdomen 195. Pharnacia ponderosa Stål, 1877, & anal segment & supraanal plate 196. Pharnacia ponderosa Stål, 1877, % apex of abdomen 197. Pharnacia borneensis spec. nov., & apex of abdomen 198. Pharnacia borneensis spec. nov., & anal segment & supraanal plate 199. Pharnacia borneensis spec. nov., 1908, % apex of abdomen 200. Pharnacia sumatrana (Brunner v. Wattenwyl , 1907), & apex of abdomen 201. Pharnacia sumatrana (Brunner v. Wattenwyl , 1907), & anal segment & supraanal plate 202. Pharnacia sumatrana (Brunner v. Wattenwyl , 1907), % apex of abdomen 203. Pharnacia heros Redtenbacher, 1908, & PLT apex of abdomen 204. Pharnacia heros Redtenbacher, 1908, % apex of abdomen 205. Pharnacia tirachus (Westwood, 1859), & apex of abdomen 206. Pharnacia tirachus (Westwood, 1859), & anal segment & supraanal plate 207. Pharnacia tirachus (Westwood, 1859), % apex of abdomen


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PLATE V. [scale = 10 mm] 208. Pharnacia kalag Zompro, 2005, & PT apex of abdomen 209. Pharnacia palawanica spec. nov., & PT apex of abdomen 210. Tirachoidea inversa (Brunner v. Wattenwyl , 1907), & HT apex of abdomen 211. Tirachoidea westwoodii (Wood-Mason, 1875), % apex of abdomen 212. Tirachoidea cantori (Westwood, 1859), & apex of abdomen 213. Tirachoidea cantori (Westwood, 1859), & anal segment & supraanal plate 214. Tirachoidea cantori (Westwood, 1859), % apex of abdomen 215. Tirachoidea herberti spec. nov., & PT apex of abdomen 216. Tirachoidea herberti spec. nov., & PT anal segment & supraanal plate 217. Tirachoidea herberti spec. nov., % HT apex of abdomen 218. Tirachoidea siamensis spec. nov., & PT apex of abdomen 219. Tirachoidea siamensis spec. nov., & PT anal segment & supraanal plate 220. Tirachoidea siamensis spec. nov., % PT apex of abdomen



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PLATE VI. [scale = 10 mm] 221. Tirachoidea biceps Redtenbacher, 1908, & apex of abdomen 222. Tirachoidea biceps Redtenbacher, 1908, & anal segment & supraanal plate 223. Tirachoidea biceps Redtenbacher, 1908, % apex of abdomen 224. Tirachoidea jianfenglingensis (Bi, 1994), & apex of abdomen 225. Tirachoidea jianfenglingensis (Bi, 1994), & anal segment & supraanal plate 226. Tirachoidea jianfenglingensis (Bi, 1994), % apex of abdomen 227. Phobaeticus ingens (Redtenbacher, 1908), & LT apex of abdomen 228. Phobaeticus ingens (Redtenbacher, 1908), % apex of abdomen 229. Phobaeticus foliatus (Bragg, 1995), & apex of abdomen 230. Phobaeticus foliatus (Bragg, 1995), & anal segment & supraanal plate 231. Phobaeticus foliatus (Bragg, 1995), % apex of abdomen 232. Phobaeticus redtenbacheri (Dohrn, 1910), & apex of abdomen 233. Phobaeticus redtenbacheri (Dohrn, 1910), % apex of abdomen


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PLATE VII. [scale = 10 mm] 234. Phobaeticus magnus nom. nov., & apex of abdomen 235. Phobaeticus magnus nom. nov., & anal segment & supraanal plate 236. Phobaeticus magnus nom. nov. (Redtenbacher, 1908), % apex of abdomen 237. Phobaeticus kirbyi Brunner v. Wattenwyl , 1907, & apex of abdomen 238. Phobaeticus kirbyi Brunner v. Wattenwyl , 1907, & anal segment & supraanal plate 239. Phobaeticus kirbyi Brunner v. Wattenwyl , 1907, % apex of abdomen 240. Phobaeticus philippinicus (Hennemann & Conle, 1997), & AT apex of abdomen 241. Phobaeticus philippinicus (Hennemann & Conle, 1997), & AT anal segment &supraanal plate 242. Phobaeticus philippinicus (Hennemann & Conle, 1997), % HT apex of abdomen 243. Phobaeticus serratipes (Gray, 1835), & apex of abdomen 244. Phobaeticus serratipes (Gray, 1835), & anal segment & supraanal plate 245. Phobaeticus serratipes (Gray, 1835), % apex of abdomen 246. Phobaeticus hypharpax (Westwood, 1859), & apex of abdomen 247. Phobaeticus hypharpax (Westwood, 1859), & anal segment & supraanal plate 248. Phobaeticus hypharpax (Westwood, 1859), % apex of abdomen 249. Phobaeticus sinetyi Brunner v. Wattenwyl , 1907, & PLT apex of abdomen 250. Phobaeticus sinetyi Brunner v. Wattenwyl , 1907, & PLT anal segment & supraanal plate 251. Phobaeticus sinetyi Brunner v. Wattenwyl , 1907, % LT apex of abdomen REVISION OF PHARNACIINI


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PLATE VIII. [scale = 10 mm] 252. Phobaeticus (?) incertus Brunner v. Wattenwyl , 1907, & HT apex of abdomen 253. Phobaeticus lumawigi Brock, 1997, & HT apex of abdomen 254. Phobaeticus lumawigi Brock, 1997, & HT anal segment & supraanal plate 255. Phobaeticus mucrospinosus spec. nov., & HT apex of abdomen 256. Phobaeticus lobulatus (Carl, 1913), & apex of abdomen 257. Phobaeticus chani Bragg spec. nov., & anal segment & supraanal plate 258. Phobaeticus chani Bragg spec. nov., & apex of abdomen (lateral view) 259. Phobaeticus chani Bragg spec. nov., & apex of abdomen (ventral view) 260. Phobaeticus chani Bragg spec. nov., % apex of abdomen (dorsal view) 261. Phobaeticus chani Bragg spec. nov., % apex of abdomen (lateral view) 262. Phobaeticus chani Bragg spec. nov., % apex of abdomen (ventral view)


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PLATE IX. [scale = 10 mm] 263. Phobaeticus palawanensis spec. nov., % apex of abdomen 264. Phobaeticus sobrinus Brunner v. Wattenwyl , 1907, % apex of abdomen 265. Phobaeticus rex (Günther, 1928), & HT apex of abdomen 266. Baculonistria gen. nov. alba (Chen & He, 1990), & apex of abdomen 267. Baculonistria gen. nov. alba (Chen & He, 1990), % apex of abdomen 268. Baculonistria gen. nov. alba (Chen & He, 1990), & anal segment & supraanal plate 269. Pharnacia ponderosa Stål, 1877, & abdominal tergite VII (dorsal view) 270. Phobaeticus rex (Günther, 1928), & HT abdominal tergite VII (dorsal view) 271. Pharnacia ingens Redtenbacher, 1908, & abdominal tergite VII (dorsal view) 272. Tirachoidea cantori (Westwood, 1859), & abdominal tergite VII (dorsal view) 273. Tirachoidea siamensis spec. nov., & abdominal tergite VII (dorsal view) 274. Phobaeticus lobulatus (Carl, 1913), & HT abdominal tergite VII (dorsal view) 275. Phobaeticus hypharpax (Westwood, 1859), & abdominal tergite VII (dorsal view) 276. Phobaeticus foliatus (Bragg, 1995), & HT abdominal tergite VII (dorsal view) REVISION OF PHARNACIINI


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PLATE X. [scale = 10 mm] 277. Phryganistria heusii (Hennemann & Conle, 1997), % left mid leg 278. Phryganistria heusii (Hennemann & Conle, 1997), & head (lateral view) 279. Phryganistria heusii (Hennemann & Conle, 1997), % head (lateral view) 280. Phryganistria grandis Rehn, 1906, & apex of abdomen 281. Phryganistria grandis Rehn, 1906, % apex of abdomen 282. Phryganistria heusii (Hennemann & Conle, 1997), & apex of abdomen 283. Phryganistria heusii (Hennemann & Conle, 1997), % apex of abdomen 284. Phryganistria fruhstorferi (Brunner v. Wattenwyl , 1907), & apex of abdomen 285. Phryganistria fruhstorferi (Brunner v. Wattenwyl , 1907), % apex of abdomen 286. Phryganistria virgea (Westwood, 1848), & apex of abdomen 287. Phryganistria virgea (Westwood, 1848), % apex of abdomen 288. Phryganistria heusii (Hennemann & Conle, 1997), & anal segment & supraanal plate 289. Phryganistria grandis Rehn, 1906, & anal segment & supraanal plate


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PLATE XI. [scale = 10 mm] 290. Pharnacia ponderosa Stål, 1877, & head (lateral view) 291. Pharnacia ponderosa Stål, 1877, % head (lateral view) 292. Pharnacia sumatrana (Brunner v. Wattenwyl , 1907), & head (lateral view) 293. Pharnacia sumatrana (Brunner v. Wattenwyl , 1907), % head (lateral view) 294. Pharnacia tirachus (Westwood, 1859), & head (lateral view) 295. Pharnacia tirachus (Westwood, 1859), % head (lateral view) 296. Tirachoidea jianfenglingensis (Bi, 1994), & head (lateral view) 297. Tirachoidea jianfenglingensis (Bi, 1994), % head (lateral view) 298. Tirachoidea siamensis spec. nov, & head (lateral view) 299. Tirachoidea biceps (Redtenbacher, 1908), & head (lateral view) 300. Tirachoidea biceps (Redtenbacher, 1908), & head (lateral view) 301. Tirachoidea biceps (Redtenbacher, 1908), % head (lateral view) 302. Tirachoidea inversa (Brunner v. Wattenwyl , 1907), & head (lateral view) 303. Tirachoidea westwoodii (Wood-Mason, 1875), % head (lateral view) 304. Tirachoidea herberti spec. nov., &HT head (lateral view) REVISION OF PHARNACIINI


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PLATE XII. [scale = 10 mm] 305. Phobaeticus serratipes (Gray, 1835), & head (lateral view) 306. Phobaeticus serratipes (Gray, 1835), % head (lateral view) 307. Phobaeticus kirbyi Brunner v. Wattenwyl , 1907, & head (lateral view) 308. Phobaeticus kirbyi Brunner v. Wattenwyl , 1907, % head (lateral view) 309. Phobaeticus redtenbacheri (Dohrn, 1910), % head (dorsal view) 310. Phobaeticus magnus nom. nov., % head (dorsal view) 311. Phobaeticus palawanensis spec. nov., % HT head (dorsal view) 312. Phobaeticus foliatus (Bragg, 1995), % head, pro and mesonotum (dorsal view) 313. Phobaeticus chani Bragg spec. nov., & HT head (dorsal view)


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PLATE XIII. [scale = 10 mm] 314. Pharnacia heros Redtenbacher, 1908, & left fore leg 315. Pharnacia sumatrana (Brunner v. Wattenwyl , 1907), & left fore leg 316. Pharnacia tirachus (Westwood, 1859), & (subadult nymph) left fore leg (MNHU) 317. Pharnacia tirachus (Westwood, 1859), & (subadult nymph) left fore leg (MNHU) 318. Pharnacia ponderosa Stål, 1877, & left mid leg 319. Pharnacia sumatrana (Brunner v. Wattenwyl , 1907), & left mid leg 320. Pharnacia kalag Zompro, 2005, & HT left mid leg 321. Pharnacia kalag Zompro, 2005, & PT left mid leg



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PLATE XIV. [scale = 10 mm] 322. Pharnacia tirachus (Westwood, 1859), & left mid leg 323. Tirachoidea biceps (Redtenbacher, 1908), & left mid leg 324. Tirachoidea siamensis spec. nov., & PT left mid leg 325. Tirachoidea inversa (Brunner v. Wattenwyl , 1907), & left mid leg (HT of Ph. semilunaris Redtenbacher, 1908) 326. Phobaeticus foliatus (Bragg, 1995), & left mid leg 327. Phobaeticus foliatus (Bragg, 1995), & left fore leg


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PLATE XV. [scale = 10 mm] 328. Phobaeticus mucrospinosus spec. nov., & HT left mid leg 329. Phobaeticus magnus nom. nov., & left mid leg 330. Phobaeticus lumawigi Brock, 1997, & HT left mid leg 331. Phobaeticus rex (Günther, 1928), & HT right mid leg 332. Phobaeticus serratipes (Gray, 1835), & left mid leg 333. Phobaeticus serratipes (Gray, 1835), & left mid leg



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PLATE XVI. [scale = 10 mm] 334. Phobaeticus hypharpax (Westwood, 1859), & left mid leg 335. Phobaeticus hypharpax (Westwood, 1859), & left mid leg 336. Phobaeticus sobrinus Brunner v. Wattenwyl, 1907, & left mid leg (HT of Ph. beccarianus Br. v. W., 1907) 337. Pharnacia tirachus (Westwood, 1859), % left profemur 338. Phobaeticus chani Bragg spec. nov., &, HT right mesofemur


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PLATE XVII. [scale = 10 mm] 339. Phobaeticus hypharpax (Westwood, 1859), & left profemur 340. Phobaeticus sinetyi Brunner v. Wattenwyl, 1907, & PLT left profemur 341. Phobaeticus sinetyi Brunner v. Wattenwyl, 1907, % LT left profemur 342. Phobaeticus sinetyi Brunner v. Wattenwyl, 1907, & left mesofemur 343. Phobaeticus serratipes (Gray, 1835), & left profemur 344. Phobaeticus serratipes (Gray, 1835), % left profemur 345. Phobaeticus kirbyi Brunner v. Wattenwyl , 1907, & left profemur 346. Phobaeticus kirbyi Brunner v. Wattenwyl , 1907, % left profemur 347. Phobaeticus palawanensis spec. nov., % HT left profemur 348. Phobaeticus philippinicus (Hennemann & Conle, 1997), & (nymph) PT basal third of left mesotibia 349. Phobaeticus philippinicus (Hennemann & Conle, 1997), & (nymph) PT base of left mesofemur 350. Phobaeticus philippinicus (Hennemann & Conle, 1997), & (nymph) PT base of left profemur 351. Baculonistria gen. nov. alba (Chen & He, 1990), %, right mid leg



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PLATE XVIII. Internal micropylar plate of eggs & praeopercular organ of && 352. Pharnacia borneensis spec. nov., internal micropylar plate 353. Pharnacia sumatrana (Brunner v. Wattenwyl , 1907), internal micropylar plate 354. Phobaeticus serratipes (Gray, 1835), internal micropylar plate 355. Phryganistria heusii (Hennemann & Conle, 1997), internal micropylar plate 356. Tirachoidea jianfenglingensis (Bi, 1904), internal micropylar plate 357. Tirachoidea siamensis spec. nov., internal micropylar plate 358. Baculonistria gen. nov. alba (Chen & He, 1990), praeopercular organ of & 359. Phryganistria heusii (Hennemann & Conle, 1997), praeopercular organ of & 360. Phobaeticus serratipes (Gray, 1835), praeopercular organ of & 361. Phobaeticus kirbyi Brunner v. Wattenwyl , 1907, praeopercular organ of & 362. Phobaeticus ingens (Redtenbacher, 1908), praeopercular organ of & 363. Tirachoidea biceps (Redtenbacher, 1908), praeopercular organ of & 364. Tirachoidea siamensis spec. nov., praeopercular organ of & PT 365. Tirachoidea jianfenglingensis (Bi, 1994), praeopercular organ of & 366. Tirachoidea herberti spec. nov., praeopercular organ of & PT


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PLATE XIX. Fig. 367. Possible relationships within the family Phasmatidae sensu lato The hypothetic tree above shows the relationships within the family Phasmatidae s. l. here suggested. It does not restrict to defining apomorphies for each clade or taxon, but also uses key-features helpful for the distinction of the group. It should hence not be interpreted as a strict phylogenetic cladogramm.



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PLATE XX. Distribution 368. Baculonistria gen. nov. alba (Chen & He, 1990) 369. Baculonistria gen. nov. longicornis (Bi & Wang, 1998) 370. Baculonistria gen. nov. yuexiensis (Chen & He, 1993) 371. Pharnacia borneensis spec. nov. 372. Pharnacia heros Redtenbacher, 1908 373. Phobaeticus ingens (Redtenbacher, 1908)


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PLATE XXI. Distribution 374. Pharnacia kalag Zompro, 2005 375. Pharnacia palawanica spec. nov. 376. Pharnacia ponderosa Stål, 1877 377. Pharnacia sumatrana (Brunner v. Wattenwyl , 1907) 378. Pharnacia tirachus (Westwood, 1859) 379. Phobaeticus chani Bragg spec. nov.



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PLATE XXII. Distribution 380. Phobaeticus foliatus (Bragg, 1995) 381. Phobaeticus hypharpax (Westwood, 1859) 382. Phobaeticus kirbyi Brunner v. Wattenwyl , 1907 383. Phobaeticus lobulatus (Carl, 1913) 384. Phobaeticus lumawigi Brock, 1997 385. Phobaeticus magnus nom. nov.


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PLATE XXIII. Distribution 386. Phobaeticus mjoebergi (Günther, 1935) 387. Phobaeticus mucrospinosus spec. nov. 388. Phobaeticus palawanensis spec. nov. 389. Phobaeticus philippinicus (Hennemann & Conle, 1997) 390. Phobaeticus redtenbacheri (Dohrn, 1910) 391. Phobaeticus rex (Günther, 1928)



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PLATE XXIV. Distribution 392. Phobaeticus serratipes (Gray, 1835) 393. Phobaeticus sinetyi Brunner v. Wattenwyl, 1907 394. Phobaeticus sobrinus Brunner v. Wattenwyl, 1907 395. Nesiophasma (?) incerta (Brunner v. Wattenwyl , 1907) 396. Phryganistria fruhstorferi (Brunner v. Wattenwyl , 1907) 397. Phryganistria grandis Rehn, 1906


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PLATE XXV. Distribution 398. Phryganistria heusii (Hennemann & Conle, 1997) 399. Phryganistria sarmentosa (Westwood, 1848) 400. Tirachoidea biceps (Redtenbacher, 1908) 401. Tirachoidea cantori (Westwood, 1859) 402. Tirachoidea herberti spec. nov. 403. Tirachoidea westwoodii (Wood-Mason, 1875)



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PLATE XXVI. Distribution & T. westwoodii (Wood-Mason, 1875) & 404. Tirachoidea inversa (Brunner v. Wattenwyl , 1907) 405. Tirachoidea jianfenglingensis (Bi, 1994) 406. Tirachoidea siamensis spec. nov. 407. Tirachoidea westwoodii (Wood-Mason, 1875), & apex of abdomen (lateral view) [reproduced from Wood-Mason, 1875, pl. 3: 1d]. 408. Tirachoidea westwoodii (Wood-Mason, 1875), & apex of abdomen (dorsal view) [reproduced from Wood-Mason, 1875, pl. 3: 1c]. 409. Tirachoidea westwoodii (Wood-Mason, 1875), & head (dorsal view) [reproduced from Wood-Mason, 1875, pl. 3: 1]. 410. Tirachoidea westwoodii (Wood-Mason, 1875), & head (lateral view, dextral) [reproduced from Wood-Mason, 1875, pl. 3: 1a]. 411. Tirachoidea westwoodii (Wood-Mason, 1875), & head (lateral view, sinistral) [reproduced from Wood-Mason, 1875, pl. 3: 1b].


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PLATE XXVII. 412. Plankwalk leading through moist lowland rainforest in Batu Niah National Park (NE-Sarawak, Malaysia), a habitat where Phobaeticus kirbyi Brunner v. Wattenwyl, 1907 is found in frequent numbers. 413. Montane forest near Mount Kinabalu (N-Sabah, Malaysia) at about 1500 metres, the type-locality of Pharnacia borneensis spec. nov. and habitat of Phobaeticus redtenbacheri (Dohrn, 1910), Phobaeticus foliatus (Bragg, 1995), Phobaeticus kirbyi Brunner v. Wattenwyl, 1907 and Phobaeticus chani Bragg spec. nov.. 414. Damage caused on Cupressus funebris (Cupressaceae) in Yunyang County, Chongqing municipality (China) by Baculonistria gen. nov. alba (Chen & He, 1990) in 2005. (© Xiang Guowei) 415. Eggs of Pharnacia Stål, 1877 and Tirachoidea Brunner v. Wattenwyl, 1907 showing the different surface structure of the capsule, being rough in Pharnacia (left, Ph. sumatrana (Brunner v. Wattenwyl, 1907)) and smooth and shiny in Tirachoidea (right, T. siamensis spec. nov.). 416. Baculonistria gen. nov. alba (Chen & He, 1990), mating couple (Chongqing: China, © Weiwei Zhang)



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PLATE XXVIII. 417. Pharnacia sumatrana (Brunner v. Wattenwyl, 1907), & (Peninsular Malaysia: Tapah Hills) 418. Phobaeticus kirbyi Brunner v. Wattenwyl, 1907, mating couple (NE-Sarawak: Batu Niah National Park) 419. Pharnacia sumatrana (Brunner v. Wattenwyl, 1907), & ventral view of thorax showing the conspicuous purplish spots characteristic for this species (Peninsular Malaysia: Tapah Hills) 420. Phobaeticus serratipes (Gray, 1835), & head and thorax showing the fine longitudinal orange lateral line typical for this species (captive reared from West Malaysia)


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PLATE XXIX. 421. Phobaeticus magnus nom. nov., & (captive reared from N-Thailand, © J. Bresseel) 422. Phobaeticus magnus nom. nov., % head and thorax (captive reared from N-Thailand, © J. Bresseel) 423. Tirachoidea siamensis spec. nov., % (captive reared from N-Thailand: Chiang Mai)



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PLATE XXX. 424. Tirachoidea jianfenglingensis (Bi, 1994), & (captive reared from N-Vietnam: Cuc Phuong) 425. Pharnacia kalag Zompro, 2005, % (captive reared from Philippines: Mindanao Id., © R. Krijns) 426. Pharnacia kalag Zompro, 2005, % head and thorax (captive reared from Philippines: Mindanao Id., © R. Krijns)


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PLATE XXXI. 427. Pharnacia ponderosa Stål, 1877, mating couple (captive reared from Philippines) 428. Tirachoidea biceps (Redtenbacher, 1908), subadult % showing the pretty white markings on the body rarely seen in this species (captive reared from Java) 429. Baculonistria gen. nov. alba (Chen & He, 1990), % closeup of head and pronotum, showing the typical white Colouration (captive reared from China: Chongqing, © R. Galunder) 430. Baculonistria gen. nov. alba (Chen & He, 1990), & (captive reared from China: Chongqing, © R. Galunder) REVISION OF PHARNACIINI


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PLATE XXXII. Figures 431-435 show the copulation position of Phasmatidae with the % grasping the longitudinal keel in the basal portion of the & subgenital plate (= sternum VIII). Figures 436–438 show the copulation position in Clitumninae with the % grasping the & abdomen at the posterior margin of sternum VII. 431. Acrophylla wuelfingi (Redtenbacher, 1908) [Phasmatinae: Phasmatini] lateral view (captive reared from NAustralia: Queensland, © R. Krijns) 432. Anchiale sp. , [Phasmatinae: Phasmatini] lateral view (captive reared from Solomon Islands: Malaita, © R. Krijns) 433. Extatosoma tiaratum tiaratum (Mac Leay, 1826) [Extatosomatinae] lateral view (captive reared from N-Australia: Queensland, © M. Duytschaever) 434. Eurycnema goliath (Gray, 1834) [Phasmatinae: Phasmatini] lateral view (captive reared from N-Australia) 435. Eurycnema goliath (Gray, 1834) [Phasmatinae: Phasmatini] ventral view (captive reared from N-Australia) 436. Pharnacia ponderosa Stål, 1877 [Clitumninae: Pharnaciini] lateral view (captive reared from Philippines) 437. Pharnacia ponderosa Stål, 1877 [Clitumninae: Pharnaciini] lateral view (captive reared from Philippines) 438. Pharnacia ponderosa Stål, 1877 [Clitumninae: Pharnaciini] ventral view of genitalia during copulation with a white spermatophore seen at the left (captive reared from Philippines, © Jérome Le Maistre)


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ADDENDUM Recently, after the present work was submitted for publication, the authors received the book “Phasmatodea of China” by Chen Shuchun and He Yunheng published in late 2007 (see citation below). Amongst several new genera, one of which is preoccupied (see below), this describes two new species of the genus Phryganistria Stål, 1875 from S-China (Guangxi Aut. Reg.). These could unfortunately not be fully integrated into the manuscript anymore. Both species are however cited in the list of species of Phryganistria and briefly discussed below. Chen, S. C. & He, Y. (2007) Phasmatodea of China. China Forestry Publishing House. 476 pp, 12 plates.

Phryganistria guangxiensis Chen & He, 2007 Phryganistria guangxiensis Chen & He, 2007: 337, figs. 308 a-b (&). HT, %: China, Guangxi Zhuang Aut. Reg., Napo Co., Defu 350 m, 19.VI.2000, leg. Li Wenzhu (IZCAS).



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Differentiation: %% of this species, the only sex known, are similar to the Vietnamese species Ph. grandis Rehn, 1906 and Ph. heusii (Hennemann & Conle, 1997) and the Chinese Ph. longzhouensis Chen & He, 2007 with which it shares the large sub-apical spine on the two outer ventral carinae of the meso- and metafemora. It however differs from the two Vietnamese species by: the smaller size; plain brown colouration of the body; unarmed posteroventral carina of the profemora, and more slender semi-tergites of the anal segment. According to Chen & He (2007: 338) it merely differs from the very similar Ph. longzhouensis, also from Guangxi autonomous region, by the more numerous serrations of the anterodorsal carina of the profemora (22-23 compared to 13–15 in longzhouensis) and differently shaped anal segment. Characterization: This species is known only from the unique % HT in IZCAS. The following brief description is based on the English translation of the Chinese original description and illustration of the HT presented by Chen & He (2007: 451, fig. 308). %: Moderately sized for the genus (body length 176.0 mm). Body surface smooth and glabrous except for a indistinct longitudinal median carina. General colouration of body and legs brown. Head: Oval, slightly narrowed posteriorly and unarmed. Antennae reaching about half way along metanotum. Scapus long and thickened apically, pedicellus cylindrical and 3rd antennomere about half the length of pedicellus. Thorax: Pronotum almost rectangular and about 1.5x longer than wide. Mesothorax, elongate and almost 5.5x longer than combined length of head and pronotum. Abdomen: Median segment less than 1/3 the length of metanotum. Abdominal segment II longer than median segment and about 2.5x longer than wide. II–VI increasing in length, VI more than 5x longer than wide, VII a little shorter than II; all gently constricted medially. Tergites VIII–X broader than previous, VIII slightly widened towards the posterior and less than 1/3 the length of VII. IX distinctly shorter than VIII and roughly quadrate. Anal segment a little longer than VIII, longitudinally split and consisting of two semi-tergites. These rather elongate and roundly triangular in lateral aspect with the anterodorsal portion rounded; apex narrowly rounded. Cerci of moderate length, just not reaching apex of anal segment and slightly in-curving. Poculum cup-like with a distinct longitudinal median carina and roughly reaching to posterior margin of tergite IX. Legs: All long and slender, the meso- and metafemora moderately thickened. Profemora considerably longer than mesothorax, mesofemora about as long as pro- and mesonotum combined, metafemora reaching posterior margin of abdominal tergite V and metatibiae distinctly projecting over apex of abdomen. Anterodorsal carina of profemora with 22–23 distinct serrations, all other carinae unarmed. Two outer ventral carinae of meso- and metafemora acutely dentate and with a long sub-apical spine, medioventral carina with only 2–3 small spines, and dorsal carinae smooth except for one or two minute teeth. Meso- and metatibiae with all three ventral carinae dentate and dorsal carinae smooth. Probasitarsus very elongate, slender and longer than remaining tarsomeres combined. Meso- and metabasitarsus about as long as following three tarsomeres combined, the ventral carinae with a few minute teeth. Measurements of the HT according to Chen & He (2007: 338) [in mm]: body 176.0, mesonotum 42.0, metanotum 26.0, median segment 7.0, profemora 59.0, mesofemora, 46.0, metafemora 55.0.

Phryganistria longzhouensis Chen & He, 2007 Phryganistria longzhouensis Chen & He, 2007: 338, figs. 309 a-d (%, &). HT, %: China, Guangxi Zhuang Aut. Reg., Longzhou Co., Sanlian 350 m, 13.VI.2000, leg. Chen Jun (IZCAS); PT, 1 %, 1 &: China, Guangxi Zhuang Aut. Reg., Longzhou Co., Sanlian 350 m, 13.VIII.2000, leg. Zhou Zhihong (CAU).


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Differentiation: According to Chen & He (2007: 339) %% are very similar to Ph. guangxiensis Chen & He, 2007 and merely differ by the less numerous serrations of the anterodorsal carina of the profemora (13–15 compared to 22–23 in guangxiensis) and differently shaped anal segment (see comments below). && appear to be similar to the two Vietnamese species Ph. grandis Rehn, 1906 and Ph. heusii (Hennemann & Conle, 1997) but differ by the shape of the anal segment. Any other differences are difficult to identify from the original description and illustrations provided by Chen & He (2007: 339). Description: The following brief description of both sexes is based on the English translation of the Chinese original description and illustrations presented by Chen & He (2007: 452, figs. 309 a–d). &: Of moderate size for the genus (body length 240.0 mm). Body surface smooth except for an indistinct, fine longitudinal median carina. General colouration of body and legs brown, the vertex with some darker brown longitudinal stripes. Head: Oval and longer than pronotum, the vertex gently rounded and smooth. Antennae reaching about ¾ the way along mesonotum. Scapus rectangular longer than wide and thickened apically. Pedicellus cylindrical and much shorter than scapus. Thorax: Pronotum with a longitudinal impressed line and the transverse median depression not reaching lateral margins of segment. Mesothorax elongate and about 3x longer than head and pronotum combined. Abdomen: Median segment trapezoidal and a little more than ¼ the length of metanotum. Segments II– VII roughly of equal width and rectangular, II–VI increasing in length, II about 1.8x, VI almost 3x longer than wide. II longer than median segment, VII about ¾ the length of VI. Praeopercular organ formed by a pair of tooth-like projections at posterior margin of sternite VII. Tergites VIII–X narrower than previous. VIII about 2/3 the length of VII, Ix much shorter than VIII and roughly quadrate in dorsal aspect. Anal segment a little longer than IX, longitudinally carinated and posterior margin with a wide and shallow median excavation. Supraanal plate small, rounded and not projecting over anal segment. Cerci small and tapered towards the tip. Subgenital plate longitudinally keeled, boat-shaped and almost reaching to apex of anal segment. Legs: All long and moderately slender. Profemora longer than pro- and mesonotum combined, mesofemora about as long as mesonotum, metafemora reaching about 2/3 the way along abdominal segment V and metatibiae reaching apex of abdomen. Anterodorsal carina of profemora with 13–16 prominent and acute serrations and posteroventral carinae less distinctly dentate. Meso- and metafemora with both outer ventral carinae distinctly but sparsely dentate, the medioventral carina with set with a few small spines, dorsal carinae smooth. Meso- and metatibiae with all three ventral carinae dentate. Posterodorsal carina very sparsely and minutely dentate and with a single enlarged tooth pre-medially. Probasitarsus longer, meso- and metabasitarsus about as long as following three tarsomeres combined. Ventral carinae of meso- and metatbasitarsus minutely dentate. Measurements according to Chen & He (2007: 338) [in mm]: body 240.0, mesonotum 48.0, metanotum 27.0, median segment 7.0, profemora 67.0, mesofemora 49.0, metafemora 58.0. %: Moderately sized for the genus (body length 142.0-175.0 mm). Body surface smooth and glabrous except for a indistinct longitudinal median carina. General colouration of body and legs brown, the vertex with a few darker brown maculations. Head: Oval, slightly narrowed posteriorly and unarmed, longer than pronotum. Antennae generally as in && but slightly longer. Thorax: Pronotum generally as in &&. Mesothorax elongate and > 5x longer than head and pronotum combined. Abdomen: Median segment longer than wide and less than ¼ the length of metanotum. Tergites VIII–X broader than II–VII. VIII slightly widened towards the posterior, IX shorter than VIII and indistinctly longer than wide. Anal segment about as long as VIII, longitudinally split and consisting of two semi-tergites. These rather elongate with the ventral margin almost straight and dorsal margin concavely excavated in the posterior



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portion; apex narrowed. Cerci small, not reaching tip of anal segment and tapered towards a narrow apex. Poculum small, cup-like and reaching to base of anal segment. Legs: All long and slender, the meso- and metafemora moderately thickened. Profemora considerably longer and mesofemora roughly equal in length to mesothorax. Anterodorsal carina of profemora with 13-15 distinct serrations and occasionally a single tooth on posteroventral carina. Armature of mid and hind legs similar to &&, but meso- and metafemora with a prominent long sub-apical spine on the two outer ventral carinae. Basitarsus longer than remaining tarsomeres combined. Measurements according to Chen & He (2007: 338) [in mm]: body 142.0–175.0, mesonotum 31.5–42.0, metanotum 21.4–26.0, median segment 5.6–7.0, profemora 40.0-63.0, mesofemora, 31.0–45.0, metafemora 36.0–55.0. Comments: As mentioned above, this species is very similar to Ph. guangxiensis Chen & He, 2007. The typelocalities of both species are very close to each other and even the measurements of the % PT match well with those of the HT of Ph. guangxiensis. Furthermore, the differences summarised by Chen & He (2007: 338) appear weakly supported and may lie well within the intraspecific variation of this species. It is therefore possible that Ph. longzhouensis is a synonym of Ph. guangxiensis, but any confirmed decision will require access to the type-specimens for detailed examination and comparison.

Leiophasma Chen & He, 2007 Leiophasma Chen & He, 2007: 320. Type-species: Leiophasma yunnanense Chen & He, 2007: 321, figs. 290 a–c, by original designation. [Preoccupied by Leiophasma Uvarov, 1940: 114]

Chen & He (2007: 320) established Leiophasma for a Chinese species of Clitumnini, obviously unaware that this generic name was already in use for a genus of the predominantly Madagascan family Anisacanthidae. Hence, Leiophasma Chen & He, 2007 is preoccupied by Leiophasma Uvarov, 1940 (Type-species: Orobia nigrolineata Stål, 1875: 17) and would deserve a replacement name. Chen & He (2007: 321) also described Paraleiophasma, an obviously very closely related genus, and distinguished it from their Leiophasma (not Uvarov, 1940) merely by the longer anal segment of &&, which is more decidedly incised posteromedially. However, as the && anal segment of Leiophasma Chen & He is very similar in structure to that of Paraleiophasma the length alone may well lie within the range of a single genus. Unfortunately, the %% of Leiophasma Chen & He and eggs of both genera are not yet know, but certainly are required for clarification. Therefore, and as Leiophasma Chen & He is preoccupied, the genus is here provisionally synonymised with Paraleiophasma Chen & He, 2007 (syn. nov.). If further studies reveal Paraleiophasma yunnanense (Chen & He, 2007, comb. nov.) to be generically different from Paraleiophasma Chen & He, a replacement will become necessary. Clarification by Chen & He is believed to be in progress.


HENNEMANN & CONLE

13 Index of scientific names The following is an index of tribal, generic and specific names in alphabetical order. Valid taxa are printed in bold letters, invalid and synonymous taxa in normal letters. In cases of valid species names, the valid genus is given in brackets. The genus mentioned for invalid species-names is the original genus in which the species was originally included. The index only contains generic and specific names which concern to the tribe Pharnaciini. acanthopus (Phasma (Bacteria)). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 182 Acanthomimini . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 32 Acanthoxylini . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 35 Achriopterini . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 54 alba (Baculonistria) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 87 annamallayanus (Phobaeticus). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 137 Baculolonga . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 129 Baculonistria . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 85 beccarianus (Phobaeticus) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 192 biceps (Tirachoidea). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .220 borneensis (Pharnacia) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 98 cantori (Tirachoidea) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 226 chani (Phobaeticus) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 138 chiniensis (Pharnacia) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 221 Cladomorphinae. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 21 Clitumninae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 62 Clitumnini . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 67 Eurycanthinae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 21 Extatosomatinae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 46 foliatus (Phobaeticus) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 141 fruhstorferi (Phryganistria). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 201 Gigantophasmatini . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 44 grande (Bactridium) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 82 grandis (Phryganistria) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 204 Gratidiini. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 49 grubaueri (Nearchus) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 195 guangxiensis (Phryganistria) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 312 herberti (Tirachoidea) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 233 heros (Pharnacia) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 103 heusii (Phryganistria). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 209 hypharpax (Phobaeticus) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 144 incertus (Nesiophasma ?) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 195 ingens (Phobaeticus) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 149 inversa (Tirachoidea). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 236 irregularis (Clitumnus) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 122 jianfenglingensis (Tirachoidea). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 240 kirbyi (Phobaeticus) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 153 lambirica (Pharnacia) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 180 Lanceocercata. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17 Lobophasma . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 129 lobulatus (Phobaeticus) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 158 Lonchodinae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 73 Lonchodini . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 75 longicornis (Baculonistria) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 91 longzhouensis (Phryganistria) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 312 lumawigi (Phobaeticus) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 160 magdiwang (Pharnacia) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 112 REVISION OF PHARNACIINI


315

magnus (Phobaeticus) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 162 maximum (Phibalosoma) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 182 maximus (Nearchus) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 162 Medaurini . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 72 mjoebergi (Phobaeticus) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 166 Monandropterini . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 43 mucrospinosus (Phobaeticus) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 167 Nearchus. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 129 Neohiraseini . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 78 operculatus (Clitumnus) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 162 Pachymorphinae. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 48 palawanensis (Phobaeticus) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 169 palawanica (Pharnacia) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 110 Pharnacia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 94 Pharnaciini . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 66, 80 Phasmatinae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 26 Phasmatini . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 28 Phobaeticus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 127 Phryganistria . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 198 philippinicus (Phobaeticus). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 171 pilicornis (Pharnacia) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 153 pinnipes (Phobaeticus) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 174 ponderosa (Pharnacia). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 112 redtenbacheri (Phobaeticus) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 175 rex (Phobaeticus) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 180 rigida (Pharnacia) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 117 sagitta (Pharnacia) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 153 sarmentosa (Phryganistria). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 212 semilunaris (Pharnacia) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 236 serratipes (Phobaeticus) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 182 siamensis (Tirachoidea) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 244 sichuanensis (Phryganistria). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 87 sinetyi (Phobaeticus) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 188 sobrinus (Phobaeticus) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 192 spectabilis (Phobaeticus) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 144 Stephanacridini. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 54 sumatrana (Pharnacia) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 117 Tirachoidea . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 217 tirachus (Pharnacia) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 122 Tropidoderinae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 37 Tropidoderini . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 39 virgea (Phryganistria) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 212 westwoodii (Tirachoidea) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 249 yuexiensis (Baculonistria) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 92 Xeroderinae. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 52


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