Studies on Neotropical Phasmatodea XVI

Zootaxa 4128 (1): 001–211 http://www.mapress.com/j/zt/ Copyright © 2016 Magnolia Press

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ZOOTAXA

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http://doi.org/10.11646/zootaxa.4128.1.1 http://zoobank.org/urn:lsid:zoobank.org:pub:B4D2CD84-8994-4CEF-B647-3539C16B6502

ZOOTAXA 4128

Studies on Neotropical Phasmatodea XVI: Revision of Haplopodini Günther, 1953 (rev. stat.), with notes on the subfamily Cladomorphinae Bradley & Galil, 1977 and the descriptions of a new tribe, four new genera and nine new species (Phasmatodea: “Anareolatae”: Phasmatidae: Cladomorphinae) FRANK H. HENNEMANN¹*, OSKAR V. CONLE² & DANIEL E. PEREZ-GELABERT³ ¹*Reiboldstrasse 11, 67251 Freinsheim, Germany. E-mail: [email protected] ²Am Freischütz 16, 47058 Duisburg, Germany. E-mail: [email protected] ³Integrated Taxonomic Information System (ITIS) and Department of Entomology, National Museum of Natural History, Smithsonian Institution, Washington, DC 20013-7012, USA. E-mail:[email protected] Website: www.phasmatodea.com * corresponding author

Magnolia Press Auckland, New Zealand

Accepted by B. Mantovani: 19 Feb. 2016; published: 27 Jun. 2016

FRANK H. HENNEMANN, OSKAR V. CONLE & DANIEL E. PEREZ-GELABERT Studies on Neotropical Phasmatodea XVI: Revision of Haplopodini Günther, 1953 (rev. stat.), with notes on the subfamily Cladomorphinae Bradley & Galil, 1977 and the descriptions of a new tribe, four new genera and nine new species (Phasmatodea: “Anareolatae”: Phasmatidae: Cladomorphinae) (Zootaxa 4128) 211 pp.; 30 cm. 27 Jun. 2016 ISBN 978-1-77557-971-7 (paperback) ISBN 978-1-77557-972-4 (Online edition)

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HENNEMANN ET AL.

Table of contents 1. 2. 3. 3.1 4. 4.1 4.2 4.2.1 4.2.2 4.2.3 4.2.4 4.2.5 5. 5.1 5.2 5.3 5.4 5.4.1 5.4.2 5.5 5.6 5.7 5.8 5.9 6. 7. 8. 8.1 8.2 8.3 8.4 9. 10.

Abstract . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5 Material & methods. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6 Abbreviations . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6 Subfamily Cladomorphinae Bradley & Galil, 1977 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7 Keys to the tribes of Cladomorphinae Bradley & Galil, 1977 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11 Discussion of the tribes of Cladomorphinae Bradley & Galil, 1977 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12 Tribe Cladoxerini Karny, 1923 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12 Tribe Cladomorphini Bradley & Galil, 1977 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15 Tribe Cranidiini Günther, 1953 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17 Tribe Hesperohasmatini Bradley & Galil, 1977 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18 Tribe Pterinoxylini n. trib. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 28 Revision of Haplopodini Günther, 1953 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 33 Genus Aploploides Rehn & Hebard, 1938 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 36 Genus Apteroplopus n. gen. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 40 Genus Cephaloplopus n. gen. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 42 Genus Diapherodes Gray, 1835 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 59 The gigantea-group . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 61 The jamaicensis-group . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 62 Genus Haplopus Burmeister, 1838 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 105 Genus Paracranidium Brock, 1998 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 148 Genus Parhaplopus n. gen. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 150 Genus Venupherodes n. gen. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 165 Species incertus sedis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 173 Biogeography . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 173 Biology & ecology . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 179 Summary . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 180 Taxonomic checklist of Haplopodini Günther, 1953 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 182 List of new taxa . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 184 List of taxonomic changes. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 184 List of designated neotypes and lectotypes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 186 Acknowledgements . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 186 References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 187

1. ABSTRACT The anareolate New World subfamily Cladomorphinae Bradley & Galil, 1977 is reviewed and keys to the six tribes currently included are presented; these are: Cladomorphini Bradley & Galil, 1977, Cladoxerini Karny, 1923, Cranidiini Günther, 1953, Pterinoxylini n. trib., Hesperophasmatini Bradley & Galil, 1977 and Haplopodini Günther, 1953 rev. stat.. New diagnoses are presented for all these tribes and possible relationships within Cladomorphinae are discusssed. Morphology of the genitalia and egg-structures indicate Cladomorphinae as presently treated to be polyphyletic. Two subordinate groups are recognized within present Cladomorphinae, which differ considerably in numerous morphological characters of the insects and eggs. The first group and here regarded as Cladomorphinae sensu stricto is formed by the mostly South American Cladomorphini + Cranidiini + Cladoxerini, while the second group is formed by the predominantly Caribbean Hesperophasmatini + Pterinoxylini n. trib. + Haplopodini. Members of the first group (= Cladomorphini sensu stricto) share the dorsally carinate basitarsus in which the two dorsal carinae are melted with another, increasingly elongated gonapophyses VIII of females which are noticeably longer than gonapophyses IX and lamellate as well as strongly displaced medioventral carina of the profemora. Cranidiini + Cladomorphini share the strongly elongated and filiform gonapophyses VIII and presence of gonoplacs in the females, specialized poculum of males and presence of a median line in the eggs. Cranidiini differs from all other tribes of Cladomorphinae by the entirely unarmed legs of both sexes, distinctly broadened and leaf-like body and prominent longitudinal keel of the mesosternum of females, prominently enlarged poculum and spinulose phallus of males as well as the conspicuous narrowing of the posteromedian gap of the internal micropylar plate of the eggs and noticeably separated median line. Cladomorphini is characteristic for the specialized vomer and poculum of males and distinct opercular structures of the eggs. Certain representatives of Cladomorphini indicate relationships to the “Phanoclesgroup” of Diapheromerinae: Diapheromerini, hence Cladomorphini as presently treated may be paraphyletic. The exclusively South American Cladoxerini (= Baculini n. syn.) differs from the other two tribes of Cladomorphinae sensu stricto by the distinctly serrate profemora of both sexes and conspicuously shortened antennae of females, which consist

REVISION OF HAPLOPODINI

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of less than 30 segments and are much shorter than the profemora in females. Genital morphology, such as the elongated gonapophyses VIII and presence of gonoplacs in females, as well as the lamellate medioventral carina of the profemora indicate close relation to Cladomorphini. Cranidiini appears to be the sister-taxon of Cladomorphini + Cladoxerini. The tribe Baculini Günther, 1953 is synonymised with Cladoxerini (n. syn.), on the basis that the type-genera of both tribes are congeneric, with Baculum Saussure, 1861 being a junior synonym of Cladoxerus St. Fargeau & Audinet-Serville, 1827 (n. syn.). The genus Tersomia Kirby, 1904 is removed from Hesperophasmatini and transferred to Cladoxerini. Wattenwylia Toledo Piza, 1938 is removed from Pachymorphinae: Gratidiini and transferred to Cladoxerini. A detailed new diagnosis is presented for Cranidiini along with a detailed differentiation and the tribe is shown to be monotypical, only containing its type-genus Cranidium Westwood, 1843. All Caribbean genera subsequently added to Cranidiini are removed and transferred to Haplopodini rev. stat.. The three tribes Hesperophasmatini + Pterinoxylini n. trib. + Haplopodini rev. stat. are closely related and might form a monophyletic clade within Cladomorphinae sensu lato. They differ from Cladomorphinae sensu stricto by the short gonapophyses VIII and reduced gonoplacs of females, unspecialized poculum of males and lack of a micropylar line in the eggs. Haplopodini Günther, 1953 is re-established (rev. stat.) and comprises almost exclusively Caribbean genera previously placed in Hesperophasmatini by Bradley & Galil (1977) or Cranidiini by Zompro, (2004). Aploploides Rehn & Hebard, 1938, Diapherodes Gray, 1835, Haplopus Burmeister, 1838 and Paracranidium Brock, 1998 were misplaced in Cranidiini and are transferred to Haplopodini. On the basis of numerous morphological characters of the insects and eggs Hesperophasmatini is removed from Pseudophasmatidae: Xerosomatinae and re-transferred to its previous position in the subfamily Cladomorphinae sensu lato. A detailed newdiagnosis of Hesperophasmatini is presented, but is only provisional since the true diversity is as yet only fractionally known. The lack of a gula distinguishes Hesperophasmatini from all other tribes. The genus Laciphorus Redtenbacher, 1908 is removed from Hesperophasmatini and transferred to Diapheromeridae: Diapheromerinae: Diapheromerini. The new tribe Pterinoxylini n. trib. is established to contain only the type-genus Pterinoxylus Audinet-Serville, 1838. It is closely related and perhaps the sister taxon of Hesperophasmatini, with which it shares the presence of rough sensory areas on the probasisternum and profurcasternum. It differs from Hesperophasmatini and Haplopodini by the presence of a tympanal region (= stridulatory organ) in the alae of females and the alveolar eggs, which possess peripheral opercular and polar structures. Haplopodini is likely to be the sister group of Pterinoxylini n. trib. + Hesperophasmatini. The tribe Haplopodini rev. stat. is revised at the species level and comprises eight almost exclusively Caribbean genera, four of which are newly described. All eight genera now contained in Haplopodini are described in detail, differentiated from their closest relatives and their relationships and systematic position within Haplopodini are discussed. Keys and maps showing their distributions are presented along with a discussion of the distributional patterns. Detailed descriptions, differential diagnoses, synonymic listings, illustrations, material listings and measurements are given of all 26 currently known species and subspecies of Haplopodini. Four new genera are described within Haplopodini. The monotypical Apteroplopus n. gen. (type-species: Dyme grosse-tuberculata Brunner v. Wattenwyl, 1907) from Honduras is the only taxon of the tribe represented in Central America. It is only known from the male which differs from all other genera by being entirely apterous. Cephaloplopus n. gen. (type-species: Cephaloplopus pulchellus n. sp.) and Parhaplopus n. gen. (type-species: Haplopus cubensis Saussure, 1868) occur only on Hispaniola and Cuba. Both are closely related to Haplopus Burmeister, 1838 but in addition to having noticeably different eggs, both genera differ from Haplopus in several morphological characters. The monotypical Venupherodes n. gen. (type-species: Platycrana venustula Audinet-Serville, 1838) is endemic to Cuba, and in females being apterous resembles the second exclusively Cuban genus Aploploides Rehn & Hebard, 1938. It however differs from all other members of Haplopodini by the laterally expanded mesonotum of females, which overlaps the mesopleurae, as well as the morphology of the eggs. Two species-groups are recognized within Diapherodes Gray, 1835. The gigantea species-group comprises the species from the Lesser Antilles, which are: D. angulata (Fabricius, 1793), Diapherodes dominicae (Rehn & Hebard, 1938), D. gigantea gigantea (Gmélin, 1789), D. gigantea saintluciae n. ssp. and Diapherodes martinicensis Lelong & Langlois, 2005. The three species of the jamaicensis species-group, which are D. achalus (Rehn, 1904), D. jamaicensis (Drury, 1773) and D. laevicollis Redtenbacher, 1906, are restricted to the two Greater Antillean islands Jamaica and Puerto Rico. Haplopus Burmeister, 1838 is the most widely distributed genus being represented on all islands of the Greater Antilles except Jamaica, and also in the Virgin Islands, Bahamas, Florida Keys, Dry Tortugas and as far southwest as the Cayman Islands and Swan Islands. Nine new species and one new subspecies are described: Cephaloplopus alope n. sp. and Haplopus sobrinus n. sp. from Cuba, Cephaloplopus euchlorus n. sp., Cephaloplopus laetus n. sp., Cephaloplopus pulchellus n. sp., Haplopus brachypterus n. sp., Haplopus intermedius n. sp. and Parhaplopus navarroi n. sp. from Hispaniola, Haplopus woodruffi n. sp. from Cayman Brac (Cayman Islands) and Diapherodes gigantea saintluciae n. ssp. from Saint Lucia. Seven of these are described from both sexes but Cephaloplopus alope n. sp. and Haplopus sobrinus n. sp. are only known from the females and Cephaloplopus laetus n. sp. only from the males. The previously unknown males of Diapherodes angulata (Fabricius, 1793), Diapherodes laevicollis Redtenbacher, 1908, Haplopus bicuspidatus de Haan, 1842 and Parhaplopus cubensis (Saussure, 1868) as well as the previously unknown female of Parhaplopus evadne (Westwood, 1859) n. comb. are described and illustrated for the first time. Descriptions and illustrations of the eggs of eleven species


HENNEMANN ET AL.

are presented: Cephaloplopus euchlorus n. sp., Cephaloplopus pulchellus n. sp., Diapherodes achalus (Rehn, 1904), Diapherodes dominicae (Rehn & Hebard, 1938), Diapherodes gigantea gigantea (Gmélin, 1789), Diapherodes martinicensis Lelong & Langlois, 2005, Diapherodes jamaicensis (Drury, 1773), Haplopus bicuspidatus de Haan, 1842, Haplopus micropterus St. Fargeau & Audinet-Serville, 1825, Parhaplopus navarroi n. sp. and Venupherodes venustula (Audinet-Seville, 1838) n. comb.. Type specimens of the newly described taxa are deposited in the collections of ANSP, NHMUK, IIBZ, FSCA, MCZC, MNHN and USNM. Six species are transferred to other genera (n. comb.): Bacteria grossetuberculata (Brunner v. Wattenwyl, 1907) to Apteroplopus n. gen.; Haplopus cubensis Saussure, 1868 and Haplopus evadne Westwood, 1859 to Parhaplopus n. gen.; Diapherodes venustula (Audinet-Serville, 1838) to Venupherodes n. gen.; Haplopus jamaicensis (Drury, 1773) and Haplopus achalus Rehn, 1904 to Diapherodes Gray, 1835. Mantis angulata Fabricius, 1793 and Diapherodes gigantea dominicae Rehn & Hebard, 1938 are removed from synonymy with D. gigantea (Gmélin, 1789) and shown to be valid species (n. stat.). Fifteen new synonymies are revealed amongst the species studied: Diapherodes longiscapha Redtenbacher, 1908 = Diapherodes achalus (Rehn, 1904) n. syn.; Haplopus grayi Kaup, 1871 = Diapherodes angulata (Fabricius, 1793) n. syn.; Diapherodes glabricollis Gray, 1835 = Diapherodes jamaicensis (Drury, 1773) n. syn.; Diapherodes pulverulentus Gray, 1835 = Diapherodes jamaicensis (Drury, 1773) n. syn.; Diapherodes christopheri Westwood, 1859 = Diapherodes jamaicensis (Drury, 1773) n. syn.; Haplopus murinus Redtenbacher, 1908 = Diapherodes jamaicensis (Drury, 1773) n. syn.; Haplopus bituberculatum de Haan, 1842 = Haplopus micropterus (St. Fargeau & Audinet-Serville, 1825) n. syn.; Haplopus cythereus Westwood, 1859 = Haplopus micropterus (St. Fargeau & Audinet-Serville, 1825) n. syn.; Haplopus ligiolus Redtenbacher, 1908 = Haplopus micropterus (St. Fargeau & Audinet-Serville, 1825) n. syn.; Haplopus ligia Westwood, 1859 = Haplopus micropterus (St. Fargeau & Audinet-Serville, 1825) n. syn.; Haplopus mayeri Caudell, 1905 = Haplopus scabricollis (Gray, 1835) n. syn.; Aplopus similis Rehn, 1904 = Haplopus scabricollis Gray, 1835 n. syn.; Diapherodes spinipes Gray, 1835 = Haplopus micropterus (St. Fargeau & Audinet-Serville, 1825) n. syn.; Haplopus obtusus Redtenbacher, 1908 = Haplopus micropterus (St. Fargeau & Audinet-Serville, 1825) n. syn. and Haplopus juvenis Redtenbacher, 1908 = Venupherodes venustula (Audinet-Serville, 1838) n. syn.. The previously presumed lost holotype of Cyprocrana microptera St. Fargeau & Audinet-Serville, 1825 (= Phasma angulata Stoll, 1813), was traced in the collection of RMNH. The designation of a neotype has become necessary for Mantis angulata Fabricius, 1793 (in MNCN) and Platycrana venustula Audinet-Serville, 1838 (in MNHU). Lectotypes are designated for eight species: Diapherodes longiscapha Redtenbacher, 1908; Diapherodes scabricollis Gray, 1835; Haplopus christopheri Westwood, 1859; Haplopus cytherea Westwood, 1859; Haplopus juvenis Redtenbacher, 1908; Haplopus ligiolus Redtenbacher, 1908; Haplopus ligia Westwood, 1859 and Haplopus murinus Redtenbacher, 1908. Key words: Phasmatodea, Phasmatidae, Cladomorphinae, Cladomorphini, Cladoxerini, Hesperophasmatini, Pterinoxylini n. trib., Haplopodini, West Indies, Honduras, Aploploides, Apteroplopus n. gen., Cephaloplopus n. gen., Diapherodes, Haplopus, Paracranidium, Parhaplopus n. gen., Venupherodes n. gen., revision, keys, new genera, new species, new subspecies, new synonyms, new combinations, lectotypes, neotypes, descriptions, differentiations, illustrations, eggs, phylogeny, biogeography, biology, habitats, ecology.

2. INTRODUCTION The phasmid fauna of the Greater Antilles is by far richer and more diverse than supposed in the past. This is particularly obvious through the availability and examination of extensive new material from the island of Hispaniola. Collections of orthopteroid insects by the third author and collaborators in the Dominican Republic during the past 20 years have revealed many interesting Phasmatodea, which multiply the number of species previously known from Hispaniola. The higher elevations of the island in particular still seem to harbor many so far unrecognized species. Three previous papers by the authors have already described nine new species and a new genus (Conle, Hennemann & Perez-Gelabert, 2006; Conle, Hennemann & Perez-Gelabert, 2008; Conle, Hennemann & Perez-Gelabert, 2014). The subfamily Cladomorphinae Bradley & Galil, 1977 is represented in the West Indies by the three tribes Hesperophasmatini Bradley & Galil, 1977 (here re-transferred from Pseudophasmatidae: Xerosomatinae), Pterinoxylini (here established) and Haplopodini Günther, 1935 (here re-established). The latter tribe in particular has colonized almost the complete West Indies and contains the largest, most striking and colourful phasmatodeans of the Caribbean Subregion, with females of some species achieving body lengths up to almost 20 cm. The predominantly West Indian Haplopodini include such well known genera as Haplopus Burmeister, 1938 and Diapherodes Gray, 1835, but the systematics is still poorly understood. Most taxa have so far remained poorly



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studied and traditionally species have been assigned to either Haplopus or Diapherodes on the basis of ♀♀ being more slender (Haplopus) or rather broader (Diapherodes). The latest studies on members of Haplopodini represented in the West Indies are the unpublished PhD-thesis of Moxey (1972) as well as the papers of Langlois & Lelong (1997), Langlois, Lelong & Dorel (2006), Langlois, Lelong, Rastel & Dorel (2000) and Lelong & Langlois (2001 & 2005) covering the taxa represented in the Lesser Antillean islands Dominica, Guadeloupe, Martinique and Saint Lucia. Most of the last major changes introduced within the classification of Cladomorphinae by Zompro (2004) contradict previous opinions (e.g. Günther, 1953; Bradley & Galil, 1977), extensive phylogenetic studies based on morphological features (Bradler, 2009) and molecular data (Whiting et al., 2003), and can also not be supported herein. Consequently, several of the taxonomic changes proposed by Zompro (2004) are here revised and clarified, e.g. Hesperophasmatini is transferred back to Cladomorphinae, the tribe Cranidiini shown to be monotypical, Haplopodini re-established, and Baculini Günther, 1953 shown to be a synonym of Cladoxerini Karny, 1923. The collected material contains several species of Haplopodini, several of which are new to science and some of which could not be attributed to any existing genus. As a result, nine new species, a new subspecies and four new genera are described herein. In addition to the Hispaniolan material, specimens from various institutions and collections throughout Europe and the United States were examined, including all the necessary type-specimens. The present paper is the 16th part of an on-going study of the New World Phasmatodea and provides a detailed revision of the entire tribe Haplopodini Günther, 1953 at the species-level. For clarification of the systematic position and distinction of Haplopodini a review of the subfamily Cladomorphinae Bradley & Galil, 1977 is presented, along with new diagnoses and keys to the tribes.

3. MATERIAL & METHODS This study is based on the examination of all necessary type-material, which has received much support from the curators of all corresponding museums and institutions. In a very few cases the examination of the type specimens was not possible, but detailed photographs, data and measurements were available. The material at hand for this study is exclusively dried and pinned. Insects and eggs were examined using an entomological lens with 4x magnification and a stereoscope (Zeiss Stemi SV 6). Eggs were examined at 10x magnification. Measurements were taken using a long ruler or a digital caliper and are given to 0.1 mm. If more than one egg has been examined, average measurements are given. All eggs examined were already laid or removed from the female's ovipositor, hence are fully developed. The terminology used for the descriptions of external and internal egg structures follows that of Clark-Sellick (1997a). If not differently cited and no live material was available, the colouration is described from dried specimens.

3.1. Abbreviations AMNH: ANSP: BMAG: NHMUK: CASC: CMNH: CUIC: ESUW: ETHZ: FSCA: HLDH: HMUG: IIBZ: LSUK: MCZC:

American Museum of Natural History, New York / USA. Academy of Natural Sciences, Philadelphia, Pennsylvania / USA. Bristol Museum and Art Gallery, Bristol / Great Britain. Natural History Museum, London / Great Britain. California Academy of Sciences, San Francisco, California / USA. Carnegie Museum of Natural History, Pittsburgh, Pennsylvania / USA. Cornell University Insect Collection, Ithaca, New York / USA. University of Wyoming, Wyoming / USA. Eidgenössische Technische Hochschule, Entomologisches Institut, Zürich / Switzerland. Florida State Collection of Arthropods, Gainesville / USA. Hessisches Landesmuseum, Darmstadt / Germany. Hunterian Museum, Glasgow / Scotland. Instituto de Investigaciones Botánicas y Zoológicas, Santo Domingo / Dominican Republic. Linnean Society, Burlington House, London / Great Britain. Museum of Comparative Zoology, Harvard University, Cambridge / USA.


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MHNG: MNCN: MNHN: MNHNSD: MNHU: NHMB: NHMJ: NHMW: OXUM: RMNH: RNV: SMNS: SMTD: UMMZ: USNM: ZMUH: ZSMC: FH: OC: HT: AT: PT: ST: LT: PLT: NT:

Museum d’Histoire Naturelle, Geneva / Switzerland. Museo Nacional de Ciencias Naturales, Madrid / Spain. Museum d’Histoire Naturelle, Paris / France. Museo Nacional de Historia Natural, Santo Domingo / Dominican Republic. Museum für Naturkunde der Humboldt-Universität, Berlin / Germany. Naturhistorisches Museum, Basel / Switzerland. Natural History Museum of Jamaica, Kingston / Jamaica. Naturhistorisches Museum, Wien / Austria. Oxford University Museum of Natural History, Oxford / Great Britain. Nationaal Natuurhistorisch Museum, Leiden / Netherlands. Reserva Natural Vale, Espírito Santo, Linhares / Brazil. Staatliches Museum für Naturkunde, Stuttgart / Germany. Staatliches Museum für Tierkunde, Dresden / Germany. University Museum of Zoology, Ann Arbour, Michigan / USA. United States National Museum of Natural History, Washington D.C. / USA. Zoologisches Museum und Universität Hamburg / Germany. Zoologische Staatssammlung, München / Germany. Private collection of Frank H. Hennemann, Kaiserslautern / Germany. Private collection of Oskar V. Conle, Bolsterlang / Germany. Holotype Allotype Paratype Syntype Lectotype Paralectoptype Neotype

4. SUBFAMILY CLADOMORPHINAE BRADLEY & GALIL, 1977 Genus eponymum: Cladomorphus Gray, 1835: 15. Cladomorphinae Bradley & Galil, 1977: 186. Zompro, 2004: 134. Otte & Brock, 2005: 32. Cladomorphidae Brunner v. Wattenwyl, 1893: 90, 98. Bacteriinae Kirby, 1904a: 348 (in part). Kevan, 1982: 282 (in part). Cladoxerinae Karny, 1923: 237 (in part). Diapheromerinae, Zompro, 2001: 228 (in part). Zompro, 2004: 139 (in part). Phibalosomini (SectioV: Phibalosomata) Redtenbacher, 1908: 399 (in part). Phibalosominae Günther, 1953: 557. Phibalosomatinae, Moxey, 1971: 67. Xerosomatinae, Zompro, 2004: 139 (in part).

Günther (1953: 557) established the subfamily Phibalosominae (= Cladomorphinae Bradley & Galil, 1977) for sections of Phibalosomini Redtenbacher, 1908 and originally subdivided it into the four tribes Cladoxerini, Phibalosomini, Haplopodini and Cranidiini. Bradley & Galil (1977: 186) took over Günther's arrangement of the Phibalosominae and generally restricted themselves to translating his keys to the tribes into English. Taxonomic changes by these authors were limited to re-naming Günther's Phibalosominae to Cladomorphinae and Günther's tribes Phibalosomini, Haplopodini and Cranidiini to Cladomorphini, Hesperophasmatini and Craspedoniini. The new name Craspedoniini for Cranidiini Günther, 1953 was not justified and introduced because Bradley & Galil confused the type-species of Craspedonia Westwood, 1841 (see Brock, 1998b: 26), hence Cranidiini remains valid. The monophyly of Cladomorphinae sensu Bradley & Galil, 1977 seems poorly supported and the subfamily



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divides into at least two well recognized sub-divisions or clades, which do not appear particularly closely related, and indicate Cladomorphinae as presently recognized may not be a natural grouping (→ see comments below). Zompro (2004) introduced several major taxonomic changes within the classification of Cladomorphinae, which however contradict all previous opinions (e.g. Günther, 1953; Bradley & Galil, 1977), extensive phylogenetic studies based on morphological features (Bradler, 2009) and molecular data (Whiting et al., 2003). The paper by Zompro (2004) postulates numerous questionable relationships with resulting erroneous taxonomic changes and several genera are omitted. Zompro (2004: 135) transferred the three genera Aploploides Rehn & Hebard, 1938, Diapherodes Gray, 1835 and Haplopus Burmeister, 1838) from Hesperophasmatini to Cranidiini. But in fact neither genus is closely related to Cranidium Westwood, 1843, the type-genus of Cranidiini, which is proven by a wide range of morphological characters that are discussed in detail below (→ 4.2.3 and table 1). Surprisingly, the only common character for these four genera that Zompro (2004: 135) mentioned to justify his action is “the smooth and shining body”. The striking and numerous fundamental morphological differences between Cranidium and the other three genera were completely overlooked by Zompro (2004: 135). Consequently, the exclusively Antillean Aploploides, Haplopus, Diapherodes and Paracranidium Brock, 1998 are here removed from the South American Cranidiini and shown to form a well-defined clade that can be easily separated not only from Cranidiini but also from Hesperophasmatini. Therefore, Haplopodini Günther, 1953 (type-genus Haplopus Burmeister, 1838) is here re-established (rev. stat. → 5) to comprise these as well as four newly described Antillean genera. Also Aplopocranidium Zompro, 2004 (Type-species: Bacteria waehneri Günther, 1940) from northwestern South America is not a member of Cranidiini as postulated by Zompro (2004), but in fact shows very close relation to Jeremia Redtenbacher, 1908 and Jeremiodes Hennemann & Conle, 2007, hence was transferred to Cladomorphini by Hennemann & Conle (2010: 104). As a result, Cranidiini remains a monotypical tribe which only contains its type-genus, the very distinctive northeast South American Cranidium. Close relationship between Cranidiini to the South American Cladomorphini is indicated by features such as the presence of a gula in both sexes, the long and filiform gonapophyses VIII, presence of gonoplacs and a distinct praeopercular organ of ♀♀, specialized poculum of ♂♂ and the egg-morphology, which includes the presence of a median line. A new diagnosis and differentiation of Cranidiini is provided below (→ 4.2.3). Although all genera retained in Hesperophasmatini by Zompro (2004), namely Hesperophasma Rehn, 1901, Hypocyrtus Redtenbacher, 1908, Lamponius Stål, 1875, Rhynchacris Redtenbacher, 1908 and Taraxippus Moxey, 1971, lack an area apicalis on the tibiae, Zompro (2004: 139) interpreted Hesperophasmatini as a subordinate taxon of the areolate subfamily Xerosomatinae (family Pseudophasmatidae). In addition to a “similar general resemblance” of the insects Zompro (2004: 139) mentioned common features of the extremities (“interodorsal carina of the protibiae lamelliform” and “mid- and hindlegs agree completely”) and eggs (“The eggs are bulletshaped”), stating Hesperophasmatini were “…simply representing derived Xerosomatinae, in which the area apicalis is reduced“ (Zompro, 2004: 139). As shown by Bradler (2009: 99) and confirmed herein these features do however not indicate any close relation to Pseudophasmatidae: Xerosomatinae and Zompro's placement of the tribe is not supported at all. This is also confirmed by molecular data presented by Whiting et al. (2003). In fact, close relation between Hesperophasmatini and the here newly recognized and re-established Haplopodini, which comprises genera formerly placed in Haplopodini (Günther, 1953) or Hesperophasmatini respectively (Bradley & Galil, 1977), is very obvious and proven by a wide range of taxonomically and phylogenetically relevant characters of the insect and egg-morphology (→ 4.2.4). Both tribes share a distribution mainly in the West Indies with only a few taxa represented in northern Central America, whereas members of the Xerosomatinae are restricted to South and Central America. Consequently, Hesperophasmatini is here removed from Pseudophasmatidae: Xerosomatinae and transferred back to Cladomorphinae. A detailed discussion is presented below (→ 4.2.4). Zompro (2004: 135) transferred Baculini Günther, 1953 from Phasmatidae: Phasmatinae to Phasmatidae: Cladomorphinae, however without providing a diagnosis or a list of genera contained. Günther (1953: 555) treated Baculini as an Oriental tribe, but because the type-genus Baculum Saussure, 1861 was described from Brazil, the tribe is an exclusively New World taxon. Baculini is here shown to be a junior synonym of Cladoxerini Karny, 1923 (n. syn.), since Baculum is a synonym of Cladoxerus St. Fargeau & Audinet-Serville, 1827, the type-genus of Cladoxerini (→ 4.2.1). The striking genus Pterinoxylus Audinet-Serville, 1838 is distributed throughout southern Central America, the northern portion of South America and also represented by one species on the southern Lesser Antilles. The genus was originally placed in Haplopodini (Günther, 1953: 557) and subsequently moved to Hesperophasmatini


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(Bradley & Galil, 1977: 188). Zompro (2004: 139) omitted Pterinoxylus in his treatment of Cranidiini and Hesperophasmatini. Detailed examination of the insect and egg-morphology of Pterinoxylus and careful comparison with taxa of Hesperophasmatini and Haplopodini have revealed several features that separate it from either tribe, e.g. the tympanal region or stridulatory organ in the basal portion of the alae of ♀♀, conspicuously displaced medioventral carina of the meso- and metatibiae, longitudinal ventral keel in the basal portion of the antennae and elongate alveolar eggs, which exhibit distinct hollow, peripheral extensions on the polar-area and capitulum. Consequently, Pterinoxylini n. trib. is here established to contain solely the striking Pterinoxylus and a detailed discussion and justification is provided below (→ 4.2.5 and table 2). Relationships within Cladomorphinae (Fig. 389): Detailed investigation and comparison of the six tribes currently contained in Cladomorphinae indicate the subfamily as presently treated may not form a natural group and does not support its monophyly. Based on several morphological characters of both the insects and eggs two distinct groups can be recognized within the present Cladomorphinae. The first group, here also regarded as the Cladomorphinae sensu stricto, is formed by the principally South American Cladomorphini + Cranidini + Cladoxerini, while the predominantly Antillean Hesperophasmatini + Pterinoxylini n. trib. + Haplopodini form a second fairly well distinguished and presumably monophyletic clade (Fig. 409). Examination of certain taxa currently attributed to other subfamilies (e.g. Diapheromerinae and Heteronemiinae) furthermore suggest Cladomorphinae sensu stricto to be paraphyletic (Hennemann & Conle, in preparation). In Cladomorphini and Cranidiini the gonapophyses VIII of ♀♀ are strongly elongated, filiform and project considerably over the apex of the anal segment (Figs. 8, 9, 16). Gonoplacs are present (Figs. 8, 16). The praeopercular organ of ♀♀ on abdominal sternum VII is very distinct and usually formed by one (Cranidiini) or two to three (Cladomorphini) spiniform appendages. Males have the terminal abdominal segments specialized in various ways, e.g. an elongated sometimes tube-like or remarkably enlarged and convex poculum, strongly enlarged hook-shaped or in-curving cerci or a spinose phallus. These specializations of the ♂♂ terminalia are likely to be apomorphic character states, but this deserves evaluation by a comprehensive phylogenetic study. The medioventral carina of the profemora of Cladomorphini is very prominent, lamellate and distinctly displaced towards the anteroventral carina and the profemora are triangular in cross-section with the anterodorsal carina conspicuously raised (♀♀ in particular). There are never sensory areas on the probasisternum or profurcasternum and both tribes have a well-developed gula. The antennae are very long and filiform with strongly elongated antennomeres. The eggs possess a median line below the micropylar plate although this is fairly different in Cladomorphini and Cranidiini. The eggs of Cladomorphini possess an operculum which is similar in structure to the opercula seen in genera of Diapheromerinae: Diapheromerini: “Phanocles-group” (→ 4.2.2), being an open and hollow net-work, and all have a ± parallel-sided micropylar plate which is open internally and exhibits a distinct median line (Fig. 50). Eggs of Cranidiini lack the aforementioned opercular structures and have a broad ovoid micropylar plate, which has a narrowing of the posterior opening before it widens into the notch. The short median line is strongly displaced towards the polar-area (Fig. 49). The striking differential characters of Cranidiini are summarized below and several of these are believed to be autapomorphies of that tribe (→ 4.2.3 and Table 1). The position of Cladoxerini within the Cladomorphinae remains somewhat questionable, but several characters support it as the sister-taxon of Cladomorphinae, which consequently would place Cranidiini as the sister-group of Cladomorphinae + Cladoxerini (Fig. 409). Close relation to Cladomorphini in particular is indicated by the distinctly triangular cross-section of the profemora, which have the anterodorsal carina strongly raised and the lamellate medioventral carina considerably displaced towards the anteroventral carina. The elongated gonapophyses VIII and presence of gonoplacs in ♀♀ are shared with both Cladomorphini and Cranidiini. However, the gonapophyses VIII are increasingly less elongated and hardly project over the apex of the anal segment, which is likely to be an autapomorphy of Cladoxerini. The genitalia of ♂♂ do not show any significant specializations as in these two tribes, but as in Cladomorphini and Cranidiini there is a well developed gula. Also the egg-morphology supports close relations between these three tribes, the operculum bearing a capitular structure (Fig. 5) and the open internal micropylar plate having a median line. Females are always apterous and the anal fan of ♂♂ is transparent as in Cladomorphini and Cranidiini. The remarkably shortened antennae of ♀♀ (Fig. 3), which consist of no more than 30 antennomeres, and serrate posterodorsal carina of the profemora of both sexes readily distinguish the tribe from Cladomorphini and Cranidiini and might also be autapomorphies of Cladoxerini (→ 4.2.1).



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Hesperophasmatini + Pterinoxylini n. trib. + Haplopodini are likely to form a monophyletic clade that differs from Cladomorphinae sensu stricto by a variety of characters. All three tribes lack gonoplacs and do not have conspicuously elongated or filiform gonapophyses VIII, these being short, not or hardly longer than gonapophyses IX and fully hidden under the anal segment. A praeopercular organ on abdominal sternum VII of ♀♀ is present, but it is much less developed and usually represented merely by a small wart-like swelling. The terminal abdominal segments of ♂♂ do not show any significant specializations and are fairly typical for Phasmatodea with a developed vomer. The medioventral carina of the profemora is not lamellate and more or less central on the ventral surface of the profemora. While Hesperophasmatini and Pterinoxylini n. trib. possess rough sensory-areas on the probasisternum and profurcasternum (Figs. 28–30, 41–43), these are lacking in Haplopodini. The presence of sensory-areas is likely to be a synapomorphy of Pterinoxylini n. trib. + Hesperophasmatini and might support a sister-group relationship between these two tribes, which would consequently place Haplopodini as the sistergroup of these two tribes (Fig. 409). Pterinoxylini n. trib. is characteristic for having a tympanal region (= stridulatory organ) in the basal portion of the alae of ♀♀, which quite certainly is an autapomorphy (Fig. 409) since it is not present in any other representative of the entire subfamily. The presence of a gula in Pterinoxylini n. trib. appears to be plesiomorphic because it is present in all three tribes of Cladomorphini sensu stricto and also distinguishes this tribe from Hesperophasmatini and Haplopodini. Females of all three tribes often possess shortened or vestigial wings and the anal fan of both sexes is mostly coloured, reticulate, tessellate or bears dark markings. The antennae are considerably more robust, on average shorter and less filiform than in members of Cladomorphinae sensu stricto (exception Cladoxerini), either more or less oval in cross-section (Pterinoxylini n. trib. and Hesperophasmatini) or with the antennomeres increasingly shortened and perlamorph (Haplopodini). The eggs of all three tribes agree in aspect of the morphology of the internal micropylar plate, which is open with a large and wide posterior gap but lacks a median line as in Cladomorphinae sensu stricto (Figs. 45–48). The lack of a median line is likely to represent a synapomorphy of Hesperophasmatini + Pterinoxylini n. trib. + Haplopodini. Furthermore, eggs lack the hollow net-like structures on the operculum seen in Cladomorphini and have rather short, shield-shaped or anteriorly pointed micropylar plates. The internal surface of the capsule is usually shiny sepia with the micropylar plate a contrasting white. Consequently, Hesperophasmatini + Pterinoxylini n. trib. + Haplopodini form a well separated group within Cladomorphinae as currently defined and are likely to be a monophyletic clade. This hypothesis however deserves evaluation by a comprehensive phylogenetic analysis of the Cladomorphinae and potentially closely related taxa. Previous phylogenetic analyses (e.g. Whiting et al., 2003; Buckley et al., 2009; Buckley et al., 2010) have provided support for a sister-group relationship between the Cladomorphinae and the tribe Stephanacridini Günther, 1953, an Old World clade that is today mostly restricted to some of the Pacific Islands, New Guinea and parts of Wallacea. This tribe is characterised by the strongly elongated gonapophyses VIII and well developed gonoplacs in ♀♀, the presence of a gula and a distinctly triangular cross-section of the profemora which has the anterodorsal carina conspicuously raised. As in most Cladomorphinae ♀♀ of Stephanacridini have a strongly elongated either lanceolate or spatulate subgenital plate and ♂♂ have rather basal terminalia with a well developed vomer and a pair of thorn-pads on the posterior margin of the anal segment. Most genera of Stephanacridini have shortened antennae (♀♀ in particular), that often are hardly longer than the profemora. If this sister-group relationship between Stephanacridini and Cladomorphinae (sensu lato) supposed by e.g. Whiting et al. (2003), Buckley et al. (2009) and Buckley et al. (2010) proves true, the elongated gonapophyses VIII and presence of gonoplacs in Cladomorphinae sensu stricto as well as the presence of a gula in Cladomorphinae sensu stricto and Pterinoxylini n. trib. must be regarded as plesiomorphic, while the increasingly elongated and filiform antennae in Cladomorphini and Cranidiini in particular will have to be interpreted as apomorphic. Furthermore, this hypothesis postulates a Gondwanan origin for the Cladomorphinae and suggests invasion of its ancestors via Gondwanan land connections from the Australasian region through Antarctica to South America perhaps during the late Cretaceous or late Tertiary. In addition to a complete taxonomic revision of Haplopodini rev. stat. at the species level, keys to the six tribes currently in Cladomorphinae, brief discussions of Cladoxerini and Cladomorphini as well as detailed new descriptions and discussions of Cranidiini (→ 4.2.3) and Pterinoxylini n. trib. (→ 4.2.5) and a more detailed discussion of Hesperophasmatini (→ 4.2.4) appear necessary and are presented below along with lists of the genera included in each of these tribes.


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Tribes of Cladomorphinae Bradley & Galil, 1977: 1. Cladoxerini Karny, 1923: 237. Type-genus: Cladoxerus St. Fargeau & Audinet-Serville, 1828. = Baculini Günther, 1953: 555. Type-genus: Baculum Saussure, 1862. n. syn. 2. Cladomorphini Bradley & Galil, 1977: 189. Type-genus: Cladomorphus Gray, 1835. = Phibalosomini Günther, 1953: 557. Type-genus: Phibalosoma Gray, 1835. 3. Cranidiini Günther, 1953: 557. Type-genus: Cranidium Westwood, 1843. 4. Haplopodini Günther, 1953: 557 rev. stat. Type-genus: Haplopus Burmeister, 1838. 5. Hesperophasmatini Bradley & Galil, 1977: 188. Type-genus: Hesperophasma Rehn, 1901. [Here re-transferred from Pseudophasmatidae: Xerosomatinae] 6. Pterinoxylini n. trib. Type-genus: Pterinoxylus Audinet-Serville, 1838.

4.1. Keys to the tribes of Cladomorphinae ♀♀ 1. 2. 3. 4. 5. -

Body cylindrical or oval in cross-section; abdomen not considerably dilated laterally; mesosternum not tectiform . . . . . . . . . 2 Body strongly flattened; abdomen prominently dilated laterally (Fig. 13); mesosternum tectiform and with a granulose longitudinal median keel (Fig. 15); NE South America . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cranidiini Antennae much longer than head and pronotum combined, consisting of >50 segments; profemora without serrations dorsally ................................................................................................... 3 Antennae distinctly shortened and hardly longer than head and pronotum combined, with 1/2 of capsule length; posteromedian notch of internal micropylar plate narrowed before it widens into the gap (Fig. 49) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Cranidiini



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4. 5. -

Capsule ovoid to bullet-shaped, < 2.5x longer than wide; polar-area and operculum without raised hollow extensions . . . . . 5 Capsule > 3x longer than wide, alveolar; polar area and operculum with hollow, peripheral or crest-like extensions (Fig. 44). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pterinoxylini n. trib. Capsule surface and operculum covered with hairy structures (Figs. 22–25)* . . . . . . . . . . . . . . . . . . . . . . . Hesperophasmatini Capsule surface without hairy structures . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Haplopodini

* This character only holds true for the currently known members of Hesperophasmatini (see → 4.2.4)

4.2. Discussion of the tribes of Cladomorphinae Bradley & Galil, 1977 Below, the six tribes currently assigned to Cladomorphinae are discussed. New diagnoses and differentiations are presented for Cranidiini and Pterinoxylini n. trib.. Only brief discussions of Cladoxerini and Cladomorphini are provided, since both tribes appear paraphyletic and deserve more comprehensive study. Hesperophasmatini is discussed in more detail, but no diagnosis is provided since this would be premature and only of provisional use. The authors are aware of a large number of still undescribed species and genera of Hesperophasmatini, which would need to be incorporated for a meaningful description.

4.2.1. Cladoxerini Karny, 1923 (Figs. 1–5) Type-genus: Cladoxerus St. Fargeau & Audinet-Serville, 1828: 445 Cladoxerinae Karny, 1923: 237 (in part). Kevan, 1982: 283. Cladoxerini Günther, 1953: 557 (in part). Bradley & Galil, 1977: 189. Otte & Brock, 2005: 32 (in part). Baculini Günther, 1953: 555. (Type-genus: Baculum Saussure, 1861) n. syn. Zompro, 2004: 134, 135. Phibalosomatini (Sectio V: Phibalosomata) Redtenbacher, 1908: 399 (in part).

Comments: Karny (1923: 237) originally established the subfamily Cladoxerinae to replace Redtenbacher's Phibalosomini. Günther (1953: 557) first used Cladoxerini for a tribe of his Phibalosominae and Bradley & Galil (1977: 189) listed it as a tribe of Cladomorphinae, while Kevan (1982: 283) treated it as a subfamily of his Bacteriidae. Baculini was originally established by Günther (1953: 555) as a tribe of Phasmatidae: Phasminae (= Phasmatinae Bradley & Galil, 1977) and included exclusively Oriental taxa. Günther's use of Baculini was based on the erroneous interpretation of Baculum Saussure, 1861 by Kirby (1904a: 327), who erroneously synonymised it with the Oriental Clitumnus Stål, 1875. In fact, Saussure (1861: 127) originally described Baculum as a subgenus of Bacillus Latreille, 1825, the type-species being the Brazilian Baculum ramosum Saussure, 1861 by monotypy. Kirby (1904a: 327) erroneously selected the Sri Lankan Bacillus cuniculus Westwood, 1859 as the type-species of Baculum, although Saussure had not included this species in the genus. Thus it was not available from the viewpoint of zoological nomenclature and cannot serve as the type-species for Baculum. Hennemann (2002) has partly clarified the confusion around the Oriental taxa erroneously assigned to Baculum and selected the Sri Lankan Bacillus cuniculus Westwood as the type-species of Cuniculina Brunner v. Wattenwyl, 1907, an exclusively Oriental genus. Hence, Hennemann & Conle (2008: 67) re-established Clitumnini Brunner v. Wattenwyl, 1893 for the Oriental genera that Günther (1953: 555) contained in Baculini. Zompro (2004: 135) discussed the genus Baculum, provided a re-description of the type-species B. ramosum, whose type is lost, and correctly assigned Baculini to the Cladomorphinae. However, Zompro did not identify the close relation between the newly recognized Baculini and Cladoxerini. The species which Zompro (2004: 136, fig. 2) described as Baculum ramosum is without any doubt congeneric with the type-species of Cladoxerus St. Fargeau & Audinet-Serville, 1828, i.e. C. gracilis St. Fargeau & Audinet-Serville, 1828. Hence, Baculum Saussure, 1861 falls as a junior synonym of Cladoxerus (n. syn.) and the tribe Baculini consequently becomes a junior synonym of Cladoxerini Karny, 1923 (n. syn.).


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FIGURES 1–5. Tribe Cladoxerini Karny, 1923. 1. Cladoxerus gracilis (St. Fargeau & Audinet-Serville, 1828) ♀: Brazil, Bahia, Santa Catharina [NHMW, No. 781]; 2. Cladoxerus gracilis (St. Fargeau & Audinet-Serville, 1828) ♂: Brazil, Bahia, Santa Catharina [NHMW, No. 781]; 3. Cladoxerus gracilis (St. Fargeau & Audinet-Serville, 1828) ♀ closeup of head showing the shortened antennae: Brazil, Espírito Santo, Linhares [RNV]; 4. Cladoxerus gracilis (St. Fargeau & Audinet-Serville, 1825) ♂ apex of abdomen: Brazil, Bahia, Santa Catharina [NHMW, No. 781]; 5. Cladoxerus gracilis (St. Fargeau & Audinet-Serville, 1828) egg in dorsolateral view: Brazil, Sao Paulo [© B. Kneubühler]. Po = Poculum, Vo = Vomer, Cer = Cerci, ML = Micropylar line.

The genus Parabactridium Redtenbacher, 1908 (Type-species: Parabactridium mirum Redtenbacher, 1908: 403) was described on the basis of ♀♀ from Madagascar in MNHN and NHMW and originally placed in close relation to Cladoxerus St. Fargeau & Audinet-Serville, 1828. Hence, subsequent authors (e.g. Bradley & Galil, 1977 or Otte & Brock, 2005) have placed the genus in the tribe Cladoxerini. However, the type-specimens appear to be lost and the systematic position within Cladoxerini has already been doubted by Cliquennois (2005: 119), who hypothesized it might be related to the Madagscan tribe Achriopterini Günther, 1953 if the type-locality given as “Madagascar” was correct. Indeed, Parabactridium is most unlikely to belong in Cladoxerini, which is supported by certain characters mentioned in the original description by Redtenbacher (1908: 403) i.e. the small size (body-length including subgenital plate 90.0 mm), uni-spinose head, fairly short median segment, proportionally very short legs and abbreviated tarsi. Some of these characters rather suppose relation to the Madagascan subfamily Antongiliinae. Consequently, the genus is here only retained in Cladoxerini until its actual position is clarified. A confirmed placement of Parabactridium however remains as yet impossible.



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A detailed diagnosis of the tribe would be premature, since investigation of supposedly closely related genera currently attributed to the Heteronemiinae: Paraleptyniini suggest Cladoxerini in its present recognition to be paraphyletic (→ see comments below). Hence, only a brief diagnosis mainly based on the type-genus Cladoxerus is provided here: Members of Cladoxerini are well characterized and distinguished from all other five tribes currently attributed to Cladomorphinae by the dorsally serrate profemora of both sexes and shortened antennae, which consist of no more than 30 antennomeres. These are somewhat shorter than the profemora and have the antennomeres fairly elongate in ♂♂, while they are never considerably longer than the head and pronotum combined with the basal antennomeres strongly shortened and perlamorph in ♀♀ (Fig. 3). There are no ocelli. Both sexes are very slender and elongate stick-like insects, the ♀♀ being apterous, the ♂♂ (at least those of the type-genus Cladoxerus) having scale-like tegmina and well developed alae (Fig. 2). The anal fan of the alae of ♂♂ is transparent. Most representatives are fairly large, ♀♀ of some species achieving body lengths of up to 240.0 mm including the long subgenital plate. The body surface is usually ± smooth and sub-glabrous. Occasionally there is a pair of spines on the head of ♀♀. A well developed and fairly large gula is present. Both sexes have a long median segment that is considerably longer than the metanotum. Females have a moderately developed praeopercular organ, which is formed by a median gap or slit near the posterior margin of abdominal sternum VII, and usually is marked by two bulge-like ridges or elevations. The subgenital plate of ♀♀ is strongly elongated, lanceolate, naviculate or spatulate and projects considerably over the apex of the abdomen (Fig. 1). The gonapophyses VIII are elongated, filiform but at best slightly exceed the posterior margin of the anal segment. Gonoplacs are present and moderately elongated. Males have the vomer well developed, sclerotized and elongately triangular with a pointed terminal hook. The poculum is small and angularly cup-shaped (Fig. 4). The profemora are distinctly triangular in cross-section, with the anterodorsal carina raised and ± distinctly serrate (Fig. 1); occasionally there may also be some teeth on the posteroventral carina. The medioventral carina is lamellate and strongly displaced towards the anteroventral carina. The mid and hind legs are trapezoidal in cross-section and ± prominently armed with a variable number of teeth or lobes. The tarsi are very elongate with the basitarsus carinate dorsally with the two dorsal carinae melted with another and considerably longer than the following tarsomeres; in ♀♀ the basitarsi often have a raised dorsal carina. Eggs are fairly small, longer than wide, laterally compressed, have the polar-area flattened and the anterodorsal portion of the capsule conspicuously convex. The capsule-surface is strongly sculptured. The micropylar plate is oval and fairly small, covering less than half the length of capsule. A median line is present. The operculum bears a raised but flattened structure, which is encircled by several conspicuous carinae on the outer surface (Fig. 5). The shape of the profemora, having the lamellate medioventral carina strongly displaced towards the anteroventral carina, dorsally carinate basitarsi, and the well-developed gula of both sexes, the presence of gonoplacs and the moderately elongated, filiform gonapophyses VIII of ♀♀ as well as the presence of a median line in the eggs are shared with Cladomorphini, which supports close relation between these two tribes and suggests a sister-group relationship between Cladoxerini and Cladomorphini (Fig. 409). However, since close relation to the tribe Paraleptyniini Zompro, 2004 (currently in Heteronemiidae: Heteronemiinae) is indicated by a good number of characters, Cladoxerini might by paraphyletic and any broader discussion on the true relationships of Cladoxerini would be premature at this point. The serrate anterodorsal carina of the profemora and increasingly shortened gonapophyses VIII of ♀♀ are likely to be autapomorphies of Cladoxerini. Whether the conspicuously shortened antennae of ♀♀ are plesiomorphic or apomorphic deserves further evaluation but is a priori interpreted as an autapomorphy of Cladoxerini in Fig. 409, since this character is unique within Cladomorphinae. Distribution: Southeastern South America (SE-Brazil, Paraguay, Uruguay, Argentina & Bolivia). Genera included: 1. Cladoxerus St. Fargeau & Audinet-Serville, 1828: 445. Type-species: Cladoxerus gracilis St. Fargeau & Audinet-Serville, 1828: 445, by monotypy. (Figs. 1–2) = Baculum Saussure, 1861: 127. Type-species: Bacillus (Baculum) ramosum Saussure, 1861: 127, by monotypy. [Originally described as a subgenus of Bacillus Latreille, 1825] n. syn. = Abrachia Kirby, 1889 : 503. Type-species: Abrachia brevicornis Kirby, 1889, by monotypy. [Synonymised with Baculum Saussure, 1861 by Zompro, 2004: 135] = Ceratiscus Caudell, 1904: 188. Type-species: Ceratiscus laticeps Caudell, 1904, by original designation. [Synonymised with Baculum Saussure, 1861 by Zompro, 2004: 135]


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2. Parabactridium Redtenbacher, 1908: 403. Type-species: Parabactridium mirum Redtenbacher, 1908: 403, by monotypy. [Presumably related to the Madagascan Antongiliini → see above] 3. Wattenwylia Toledo Piza, 1938: 4. Type-species: Wattenwylia foliata Toledo-Piza, 1938: 6, by original designation. [Here transferred from Pachymorphinae: Gratidiini]

FIGURES 6–12. Tribe Cladomorphini Brunner v. Wattenwyl, 1893. 6. Cladomorphus phyllinus Gray, 1835 ♀: Brazil, Espírito Santo [coll. FH, No. 0184-1]; 7. Cladomorphus phyllinus ♂: Brazil, Espírito Santo [coll. FH, No. 0184-2]; 8. Cladomorphus phyllinus: apex of ♀ abdomen (lateral view) [coll. FH, No. 0184-1]; 9. Cladomorphus phyllinus: apex of ♀ abdomen (dorsolateral view) [coll. FH, No. 0184-1]; 10. Cladomorphus phyllinus: apex of ♂ abdomen: Brazil [coll. FH, No. 0184-2]; 11. Cladomorphus phyllinus: egg from Brazil, Sao Paulo (dorsal view) [coll. FH, No. 0184-E2]; 12. Cladomorphus phyllinus: egg from Brazil, Sao Paulo (lateral view) [coll. FH, No. 0184-E2]. Ep = Epiproct, Cer = Cerci, Pa = Paraproct, Gpl = Gonoplacs, Gap8 = Gonapophyses VIII, PrOrg = Praeopercular organ, Vo = Vomer, Po = Poculum, ML = Micropylar line.

4.2.2. Cladomorphini Bradley & Galil, 1977 (Figs. 6–12, 50) Type-genus: Cladomorphus Gray, 1835: 15. Cladomorphini Bradley & Galil, 1977: 189. Otte & Brock, 2005: 32.



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Hennemann & Conle, 2010: 101. Cladoxerinae Karny, 1923: 237 (in part). Cranidiini, Zompro, 2004: 134 (in part). Otte & Brock, 2005: 32 (in part). Phibalosomatini (Sectio V: Phibalosomata) Redtenbacher, 1908: 399 (in part). Phibalosomini Günther, 1953: 557.

Comments: Günther (1953: 557) established Phibalosomini as a tribe of Phibalosominae (= Cladomorphinae Bradley & Galil, 1977) and distinguished it from the other three tribes in that subfamily i.e. Cladoxerini, Haplopodini and Cranidiini, either by the long antennae, ♀♀ lacking tegmina and alae or distinct medioventral carina of the profemora, which is conspicuously displaced towards the anteroventral carina. As the type-genus Phibalosoma Gray, 1835 is a synonym of Cladomorphus Gray, 1835, Bradley & Galil (1977: 188, 189) renamed Günther's subfamily and tribe Cladomorphinae and Cladomorphini. Günther contained exclusively Neotropical genera except for the Madagascan Parabactridium Redtenbacher, 1908, but since the type specimen(s) of the typespecies Parabactridium mirum Redtenbacher, 1908 are lost a confirmed assignment to an existing subfamily or tribe is currently impossible. Certainly however, Parabactridium is rather unlikely to be a member of Cladomorphini and presently attributed to Cladoxerini (→ 4.2.1). The genus Aplopocranidium Zompro, 2004 was misplaced in Cranidiini and transferred to Cladomorphini by Hennemann & Conle (2010: 104). The strongly elongated and filiform gonapophyses VIII, which considerably project over the apex of the anal segment (Figs. 8–9), and distinct praeopercular organ (Fig. 8) are shared with ♀♀ of Cranidiini. The developed gonoplacs and presence of a gula are in common with Cranidiini and Cladoxerini. A sister-group relationship with Cladoxerini (Fig. 409) is supported by the morphology of the profemora, fairly slender, stick-like body of ♀♀ and morphology of the eggs. As currently recognized Cladomorphini appears to be paraphyletic. In particular, genera that were placed in the “Phanocles-group” of Diapheromeridae: Diapheromerinae: Diapheromerini by Zompro (2001: 195) share numerous morphological characters that support close relation. These include the elongated and filiform gonapophyses VIII, well developed gonoplacs and distinct praeopercular organ of ♀♀, various specializations of the genitalia of ♂♂ as well as the prominent, lamellate and considerably displaced medioventral carina of the profemora and presence of a gula in both sexes. Also the egg-morphology of members of the “Phanocles-group” supports close relationship with Cladomorphini, the capitulum bearing a raised hollow structure and the internal micropylar plate being open with a distinct median line. A more detailed discussion of the tribe Cladomorphini and revision that also covers the genera attributed to the “Phanocles-group” by Zompro (2001: 195) is subject of a forthcoming study by the two first authors (Hennemann & Conle, in preparation). Since this will clarify the here suggested relationships, no broader discussion shall be presented at this point. The following brief and certainly preliminary diagnosis of the tribe is based on the eight genera that are currently attributed to Cladomorphini: Large to very large (♀♀ up to 245.0 mm), moderately slender to fairly stocky and robust Cladomorphinae with strong sexual dimorphism. Females are apterous and considerably larger than ♂♂, which are slender, stick-like insects mostly with scale-like tegmina and well developed alae (exception are the apterous ♂♂ of Otocraniella Zompro, 2004). The anal region of the alae is transparent grey or brown. Ocelli are lacking in both sexes. The body surface is smooth (spines of mesonotum disregarded) and slightly shiny (Jeremia Redtenbacher, 1908 or Aplopocranidium Zompro, 2004) to densely granulose (Jeremiodes Hennemann & Conle, 2006 or Cladomorphus Gray, 1835) or rugose (Xylodus Saussure, 1859). The head is either elongate or oval with the vertex almost flat and unarmed (Jeremiodes), has the vertex ± convex and either unarmed (e.g. Jeremia & Hirtuleius Stål, 1875), tuberculate (Cladomorphus), or bears two prominent horns or foliaceous appendages (Otocrania Redtenbacher, 1908 and Xylodus). A gula is present. The antennae are filiform, longer than the mesothorax and consist of >50 antennomeres. The scapus is dorsoventrally flattened, the remaining antennomeres cylindrical. The mesothorax is elongate, at least 2x longer than the head and pronotum combined and often armed with enlarged tubercles, spines or irregular swellings dorsally. The median segment is longer than the metanotum. Abdominal tergum V often bears crest-like posterior lobes and on sternum VII of ♀♀ there is a ± distinct praeopercular organ (Fig. 8). Occasionally, also tergites VI and VII of ♀♀ may bear posterolateral lobes. Females have the gonapophyes VIII strongly elongated, filiform and projecting considerably over the apex of the abdomen (Figs. 8–9). Gonoplacs are well developed. The subgenital plate is elongated, either lanceolate or irregularly spatulate, and extends


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considerably over the apex of the abdomen (Figs. 8–9). Males show conspicuous specializations of the genitalia, either having the cerci strongly enlarged and hook-like (Aplopocranidium, Jeremia and Jeremiodes), the poculum apically elongated into a tube-like or spatulate appendage (Cladomorphus, Otocraniella, Otocrania and Xylodus, Fig. 10) or the posterior margin of the poculum with two terminal teeth (Jeremiodes and Jeremia). The vomer is well developed and sclerotized, being a rather elongate hook-like organ with one (e.g. Jeremia and Jeremiodes) or two terminal points (Cladomorphus, Otocraniella and Xylodus). The profemora are triangular in cross-section, having the anterodorsal carina raised and ± lamellate and the posterodorsal carina considerably reduced. The medioventral carina is distinct, mostly lamellate and ± conspicuously displaced towards the anteroventral carina. The meso- and metafemora and tibiae are trapezoidal in cross-section and often bear single enlarged teeth, spines or lobes. The basitarsi are carinate dorsally with the the two dorsal carinae melted with another (exceptions are Aplopocranidium and Jeremiodes). The eggs are ± laterally compressed, have a hollow, net-like capitulum and a ± elongate, roughly parallel-sided micropylar plate, which covers at least half of the dorsal capsule surface (Figs. 11–12). Internally the plate is open with a narrow posteromedian gap and a distinct but clearly separated median line (Fig. 50). A well supported autapomorphy of Cladomorphini is represented by the conspicuous specializations of the ♂♂ genitalia, some of which are unique within the entire Phasmatodea (i.e. the strongly elongated and tube-like or spatulate poculum seen in Cladomorphus, Xylodus and Otocraniella, which projects considerably over the apex of the abdomen and resembles an elongated ♀ subgenital plate, Fig. 10). Distribution: Northern half of South America. Genera included: 1. Aplopocranidium Zompro, 2004: 134. Type-species: Bacteria waehneri Günther, 1940: 456, 495, by original designation of Zompro, 2004: 134. 2. Cladomorphus Gray, 1835: 15. Type-species: Cladomorphus phyllinus Gray, 1835: 15, by subsequent designation of Rehn, 1904: 61. (Figs. 3–8, 45) = Phibalosoma Gray, 1835: 42. Type-species: Phibalosoma lepelletieri Gray, 1835: 42, by monotypy. [Synonymised by Kirby, 1904a: 356] 3. Hirtuleius Stål, 1875: 29. Type-species: Hirtuleius laeviceps Stål, 1875: 81, by monotypy. 4. Jeremia Redtenbacher, 1908: 425. Type-species: Jeremia grossedentata Redtenbacher, 1908: 425, by monotypy. 5. Jeremiodes Hennemann & Conle, 2007: 2. Type-species: Jeremiodes guianensis Hennemann & Conle, 2007: 6, by original designation of Hennemann & Conle, 2007: 2. 6. Otocrania Redtenbacher, 1908: 423. Type-species: Bacteria aurita Burmeister, 1838: 565, by subsequent designation of Brock, 1998: 26. 7. Otocraniella Zompro, 2004: 137. Type-species: Otocraniella flagelloantennata Zompro, 2004: 137, by original designation of Zompro, 2004: 137. 8. Xylodus Saussure, 1859: 62. Type-species: Xylodus adumbratus Saussure, 1859: 62, by monotypy.

4.2.3. Tribe Cranidiini Günther, 1953 (Figs. 13–19, 49) Type-genus: Cranidium Westwood, 1843: 49. Cranidiini Günther, 1953: 557. Brock, 1998b: 26. Clark-Sellick, 1998: 221. Zompro, 2004: 135 (in part). Otte & Brock, 2005: 32 (in part). Hennemann, Conle & Delfosse, 2007: 358 (in part). Bacteriinae Kirby, 1904a: 348 (in part). Craspedoniini Bradley & Galil, 1977: 187. [Unnecessary replacement name] Phibalosomini (Sectio V: Phibalosomata) Redtenbacher, 1908: 399 (in part).

Description: ♀♀/♂♂ (Figs. 9–10). Medium-sized to large (body lengths: ♀♀ incl. subgenital plate 107.0–161.0 mm; ♂♂ 78.0–115.0 mm), Cladomorphinae with remarkable sexual dimorphism. Body surface shiny, ♀♀ plain bright green. ♂♂ with short tegmina and well developed alae (48.2–53.2 mm), ♀♀ apterous. ♂♂ elongate and moderately slender with body cylindrical. ♀♀ considerably larger and much broader than ♂♂, body strongly REVISION OF HAPLOPODINI


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dorsoventrally flattened with meso-, metathorax and abdomen strongly laterally dilated (maximum body width 28.0–34.0 mm); body rhombic in cross-section. Head longer than wide, strongly globose and convex; vertex smooth. No ocelli. A gula is present. Antennae very long, slender and filiform, longer than head, thorax and median segment combined and consisting of >80 antennomeres. Pronotum distinctly narrower than head and mesonotum. Mesothorax of ♂♂ elongate and slender, 2.5x longer than head and pronotum combined, minutely granulose; mesosternum with a fine longitudinal median carina. Mesothorax of ♀♀ broadly and semi-circularly expanded, 1.5x longer than wide. Mesonotum strongly convex dorsally and armed with a cluster of blunt tubercles in the raised anterior portion; lateral margins expanded, lamellate and bluntly dentate. Mesopleurae strongly dilated, forming a longitudinal lamella with the outer margin dentate. Mesosternum strongly tectiform and with a prominent and broad, tuberculose longitudinal median keel; lateral margins slightly expanded and with a longitudinal row of tubercles (Fig. 15). Median segment distinctly longer (♂♂) or roughly equal in length to metanotum (♀♀). Abdominal segments II–VI of ♂♂ parallel-sided and distinctly longer than wide. In ♀♀ tergites and sternites II–VI with lateral margins strongly expanded and lamellate; all transverse. Praeopercular organ of ♀♀ represented by a small spiniform projection on sternum VII (Fig. 16). Epiproct and cerci of both sexes very small and considerably shorter than anal segment. Gonapophyses VIII of ♀♀ elongated, up-curving and projecting well beyond posterior margin of anal segment (Fig. 16). Gonoplacs developed (Fig. 16). Vomer of ♂♂ produced, well sclerotized and elongate-triangular. Phallus totally covered with minute black denticles. Poculum very bulgy, strongly convex and extending ventrally by more than 2x the body width (Fig. 17). Subgenital plate of ♀♀ distinctly keeled longitudinally, boat-shaped, tapered towards a narrow ± acute apex and projecting considerably over anal segment (Fig. 16). All legs elongate and slender (♂♂) to moderately robust (♀♀), all distinctly carinate but entirely unarmed. Femora trapezoidal, tibiae almost triangular in cross-section with dorsal carinae strongly approaching each other. Profemora compressed and curved basally with posterodorsal carina reduced and considerably lower than anterior carina; medioventral carina distinct and roughly midways on ventral surface of femur. Medioventral carina of meso- and metafemora very faint. Tarsi simple. Basitarsi at least as long as following three tarsomeres combined and carinate dorsally with the two dorsal carinae melted with another. Eggs (Figs. 18–19, 49): Medium-sized, strongly globose and almost spherical in lateral aspect; slightly oval in cross-section. Polar-area with a wide depression. Operculum elliptical, flat and with a convex, net-like hollow expansion which covers the complete disc. Micropylar plate covering about half of capsule-length, broadly ovoid with a raised outer margin and convex central region. Internal micropylar plate open and with a narrowing of the posteromedial gap before it widens into the notch. Median line indistinct, short, very well separated from the plate and distinctly displaced towards the polar-area (Fig. 49). Differentiation (Table 1): Several features such as the presence of a gula, filiform antennae, the elongate, filiform gonapophyses VIII and presence of gonoplacs in ♀♀, specialized poculum of ♂♂ and presence of a median line in the eggs prove close relation to the South American Cladomorphini (→ 4.2.2). Cranidiini however clearly differs from Cladomorphini by: the shiny body surface; green general colouration; entirely unarmed legs; indistinct medioventral carina of the profemora which is roughly central on the ventral surface of the profemur; roughly trapezoidal cross-section of the profemora and very faint medioventral carina of the meso- and metafemora of both sexes. Furthermore, ♀♀ readily differ by: the strongly flattened and laterally dilated, leaf-like body; convex mesonotum and strongly longitudinally tectiform mesosternum (Fig. 15). Males are also well distinguished by the strongly enlarged and very bulgy poculum which ventrally extends by more than 2x the body diametre, as well as the specialized phallus which is totally covered with minute black denticles. The eggs differ from those of Cladomorphini by having the micropylar plate oval and relatively shorter (< ½ of the dorsal capsule surface) and the posteromedian notch of the plate with a narrowing before it widens into the gap (Fig. 49). The completely reduced leg armature and tectiform mesosternum of both sexes, the prominently dilated mesothorax and abdomen of ♀♀ as well as the prominently enlarged poculum and spinulose phallus of ♂♂ appear to be autapomorphies of Cranidiini, which suggest the tribe might be the sister-group of Cladomorphini + Cladoxerini (Fig. 409) and at once distinguish it from these two tribes. The elongated gonapophyses VIII and presence of gonoplacs in ♀♀ as well as the developed gula are shared with Cladomorphini and Cladoxerini. Because of the erroneous and confusing treatment of Cranidiini by Zompro (2004: 135) a detailed comparison and differentiation of Cranidiini and Haplopodini is presented in Table 1, which emphasizes the striking differences between these two tribes and confirms the separate status of Haplopodini as well as the monotypy of Cranidiini.


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FIGURES 13–19. Tribe Cranidiini Günther, 1953. 13. Cranidium gibbosum (Burmeister, 1838) ♀: captive reared from French Guiana [coll. FH, No. 0470-11]; 14. Cranidium gibbosum ♂: captive reared from French Guiana [coll. FH, No. 0470-9]; 15. Cranidium gibbosum: mesosternum of ♀ [coll. FH, No. 0470-1]; 16. Cranidium gibbosum: apex of ♀ abdomen (lateral view) [coll. FH, No. 0470-17]; 17. Cranidium gibbosum: apex of ♂ abdomen (lateral view) [coll. FH, No. 0470-19]; 18. Cranidium gibbosum: egg (dorsal view) [coll. FH, No. 0470-E]; 19. Cranidium gibbosum: egg (lateral view) [coll. FH, No. 0470-E]. Gpl = Gonoplacs, Gap8 = Gonapophyses VIII, Gap9 = Gonapophyses IX, PrOrg = Praeopercular organ, Po = Poculum, Cer = Cerci, Vo = Vomer, ML = Micropylar line.



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Comments: When Günther (1953: 557) established the tribe Cranidiini, Cranidium Westwood, 1843 was the only genus contained. A comprehensive treatment and re-description of the type-genus was presented by Hennemann, Conle & Delfosse (2007). Brock (1998b) described a second Jamaican genus in the tribe, Paracranidium. This however, is here shown to be not closely related and to take on a rather isolated position within the re-established tribe Haplopodini Günther, 1953 (→ 5.6), hence is removed from Cranidiini. Another South American genus, Aplopocranidium, was added by Zompro (2004: 135) but is also not closely related to Cranidium and was transferred to Cladomorphini by Hennemann & Conle (2010: 104) who provided a comprehensive treatment of Aplopocranidium including a description of the previously unknown ♂♂. Zompro (2004: 135) removed the genera Aploploides Rehn & Hebard, 1938, Haplopus Burmeister, 1838 and Diapherodes Gray, 1835 from Hesperophasmatini Bradley & Galil, 1977 and transferred them to Cranidiini. It is surprising that Zompro (2004: 135) argued the supposed close affinity of these three Antillean genera to Cranidiini exclusively on the fact that they have the same “smooth and shining body” and mentioned this as the only feature that in his opinion characterizes Cranidiini within Cladomorphinae. This act however is very arbitrary, since a wide range of very striking and fundamental morphological differences are present between Cranidiini and the mentioned three genera (→ Table 1). Nor does this “feature” hold true for all taxa that Zompro transferred to Cranidiini, e.g. ♀♀ of Haplopus-species and ♂♂ of certain Diapherodes-species have a rather dull and ± sculptured body surface. As can be seen in Table 1 Aploploides, Haplopus and Diapherodes are not at all closely related to Cranidium and here shown to form a very well separated clade, defined as the supposedly monophyletic tribe Haplopodini rev. stat. (→ 5.). Consequently, these three genera are here removed and Cranidiini is a monotypical and very well separated tribe within the subfamily Cladomorphinae. Clark-Sellick (1998: 221, fig. 32a, b) and Hennemann, Conle & Delfosse (2007: 360) stated the egg of Cranidiini to lack a median line, although the latter authors showed there to be a median line in the illustration of the internal plate (2007: 365, fig. 12). Subsequent examination of more examples has shown a short and rather indistinct median line to be present. However, it is rather short and very well separated from the micropylar plate, being displaced towards the polar-area by more than its own length. A captive reared ♀ of C. gibbosum at hand from the first author's collection (coll. FH, No. 0470-14) represents the new length record for this species, measuring 161.0 mm including the subgenital plate. Distribution: Restricted to Northeastern South America and so far recorded from Northern Brazil, French Guiana and Suriname (Hennemann, Conle & Delfosse, 2007: 366). Genus included: 1. Cranidium Westwood, 1843: 49. Type-species: Diapherodes (Cranidium) serricollis Westwood, 1843: 49, pl. 6: 1 (= Diapherodes gibbosa Burmeister, 1838), by subsequent designation of Bradley & Galil, 1977: 187. = Phasmilliger Carrera, 1960: 100. Type-species: Diapherodes (Cranidium) serricollis Westwood, 1843: 49, by indication. [Unnecessary replacement name, hence a junior objective synonym]

TABLE 1. Comparison and differentiation of Cranidiini and Haplopodini Cranidiini

Haplopodini

Strongly dorsoventrally flattened and laterally dilated; leaf-like

Moderately slender and stick-like to very robust

Rhombic

Cylindrical to oval (or triangular*)

Yes

No

Filiform; > 80 antennomeres

Robust; < 80 antennomeres

Mesonotum (♀♀)

Strongly convex and rounded with lateral margins expanded, lamellate and dentate

Simple (or with a prominent semicircularly raised longitudinal keel*); lateral margins not considerably expanded and unarmed

Mesopleurae (♀♀)

Strongly laterally expanded, lamellate and with a longitudinal marginal row of teeth (Fig. 15)

Simple and tuberculose or with a dentate longitudinal keel; rarely unarmed

Body shape (♀♀) Body cross-section (♀♀) Gula Antennae

......continued on the next page


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TABLE 1. (Continued) Cranidiini

Haplopodini

Mesosternum (♀♀)

Tectiform; with a prominent, tuberculose longitudinal median keel (Fig. 15)

Simple; may be granulose / tuberculose

Mesosternum (♂♂)

With a fine longitudinal median carina

Simple; may be granulose / tuberculose

Abdominal segments II–VII (♀♀)

Lateral margins strongly laterally dilated and lamellate (Fig. 13)

Not considerably dilated laterally or lamellate; VII often with a ± distinct lateral lobe or expansion

Gonapophyses (♀♀)

Elongated and filiform; projecting considerably beyond apex of anal segment (Fig. 16)

Short; hidden under anal segment

Phallus (♂♂)

Covered all over with minute black denticles

Unarmed

Poculum (♂♂)

Very bulgy; strongly enlarged, boatlike and convex; extending ventrally by more than 2x of body diametre (Fig. 17)

Cup-like; not considerably enlarged or specialised

Leg armature

None

At least with sub-apical spines on the two outer ventral carinae of meso- and metafemora; often with further spines, teeth or lobes

Very faint

Distinct; often spinose

Carinate dorsally; the two dorsal carinae melted with another

Sulcate dorsally; the two dorsal carinae clearly separated

Posteromedian notch with a narrowing before it widens towards the micropylar cup (Fig. 44)

Posteromedian notch large and widely triangular (Fig. 40)

Yes; very short and considerably displaced towards polar-area (Fig. 49)

No (Fig. 45)

NE South America

West Indies, Florida Keys & Honduras (Fig. 52)

Medioventral carina of mesoand metafemora Basitarsus Egg (micropylar plate— internally) Egg (Median line)

Distribution * Paracranidium Brock, 1998

4.2.4. Hesperophasmatini Bradley & Galil, 1977 (Figs. 20–35, 47–48) Type-genus: Hesperophasma Rehn, 1901: 271. Hesperophasmatini Bradley & Galil, 1977: 188 (in part). Cladoxerinae Karny, 1923: 237 (in part). Diapheromerini “Clonistria-group” Zompro, 2001: 228 (in part). Haplopodini Günther, 1953: 557 (in part). Phibalosomini (Sectio V: Phibalosomata), Redtenbacher, 1908: 399 (in part). Phibalosominae, Shelford, 1908: 343, 355 (in part). Phibalosomatinae, Moxey, 1971: 44 (in part).



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FIGURES 20–23. Tribe Hesperophasmatini Bradley & Galil, 1977. 20. Lamponius guerini (Saussure, 1868) ♀: captive reared from Guadeloupe, Bassé Terre [coll. FH, No. 0084-39]; 21. Lamponius portoricensis Rehn, 1903 ♂: captive reared from Puerto Rico, Luquillo Experimental Forest [coll. FH, No. 0467-6]; 22. Hesperophasma planulum (Westwood, 1859) ♂: Dominican Republic, Province Barahona [USNM]; 23. Hesperophasma planulum (Westwood, 1859) ♀: Dominican Republic [MNCN].


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FIGURES 24–30. Tribe Hesperophasmatini Bradley & Galil, 1977. 24. Lamponius guerini (Saussure, 1868) egg (dorsolateral view): captive reared from Guadeloupe, Bassé Terre [coll. FH, No. 0084-E]; 25. Agamemnon cornutus (Burmeister, 1838) egg (dorsal view): captive reared from Virgin Islands, Tortola [coll. FH, No. 0589-E]; 26. Rhynchacris ornata Redtenbacher, 1908 egg (dorsal view): captive reared from S-Costa Rica, Prov. Limón [coll. FH, No. 0134-E]; 27. Hypocyrtus scythrus (Westwood, 1859) egg (dorsolateral view): captive reared from Mexico, Veracruz, Tuxtlas [coll. FH, No. 0638-E]; 28. Rhynchacris ornata Redtenbacher, 1908 ♀: sensory-area of profurcasternum [coll. FH, No. 0134-12]; 29. R. ornata ♀: left sensory area of prosternum [coll. FH, No. 0134-12]; 30. Hypocyrtus scythrus (Westwood, 1859) ♀: sensory-area of profurcasternum [coll. FH, No. 0638-1].

Differentiation (Table 2): Close relation to Pterinoxylini n. trib. and Haplopodini is obvious and a detailed but as yet provisional differentiation from these two tribes is presented in Table 2. These three tribes are likely to represent a monophyletic clade that differs from Cladomorphinae sensu stricto by a number of characters (→ see above). The differentiation can at this point only be provisional and deserves considerably more research, since the authors have at hand several as yet undescribed genera and species that would need to be incorporated for providing a sufficient and satisfying differential diagnosis (→ see comments below). Eggs of the currently known genera readily differ from both these tribes by the distinct hairy structures of the capsule and operculum (Figs. 24–27), but eggs of some of the as yet undescribed taxa have almost entirely smooth capsules. A character that is unique within this clade and the entire Cladomorphinae is the secondary beak-like ovipositor in ♀♀ of Agamemnon Moxey, 1971 and Rhynchacris Redtenbacher, 1906, which is formed by an elongated subgenital plate and epiproct. The most obvious characters that distinguish Hesperophasmatini from Pterinoxylini n. trib. are: the much smaller size (body lengths: ♀♀ 33.0–94.0 mm, ♂♂ 24.5–90.0 mm); more robust and stocky body (♀♀ in particular); rather rectangular cross-section of the profemora (distinctly triangular with the anterodorsal strongly raised in Pterinoxylini n. trib.); not broadly expanded and lamellate dorsal carinae of the protibiae and not displaced medioventral carina of the meso- and metafemora of both sexes. Furthermore, Hesperophasmatini lack developed tegmina or alae in ♀♀ and at best have very small and scale-like rudiments of alae in Hypocyrtus



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Redtenbacher, 1908. Males are either apterous, brachypterous or have fully developed alae. The presence of sensory areas on the probasisternum and/or profurcasternum (Figs. 28–30) is shared with Pterinoxylini n. trib., which is likely to be a synapomorphy of Pterinoxylini n. trib. + Hesperophasmatini. This would support a sistergroup relationship between these two tribes and distinguishes both from their possible sister-group, the Haplopodini (Fig. 409). The lack of a gula In Hesperophasmatini might be an autapomorphy but this deserves evaluation since this character is shared with Haplopodini. From Haplopodini members of this tribe at once differ by: the generally more robust body and shorter legs (♀♀ in particular); presence of rough sensory areas on the probasisternum and/or profurcasternum (exception Lamponius Stål, 1875); generally more elongate antennomeres, and more rectangular cross-section of the profemora (± triangular in Haplopodini). Males of the currently known genera furthermore differ by being mostly brown or grey, while those of Haplopodini are often very colourful insects. Comments: No detailed and definite diagnosis of Hesperophasmatini is presented here, since it appears as yet premature. The authors have at hand a large number of new species and even new genera mostly from Hispaniola, whose description will be the subject of forthcoming publications. Several of these new taxa exhibit characters, which deserve comprehensive examination and evaluation and will change our present view of current Hesperophasmatini in various aspects. Hence, any diagnosis would now only be preliminary and cause more confusion instead of supporting the actual systematic position and clarifying the relationships of Hesperophasmatini within the subfamily Cladomorphinae. Some morphological features of the seven currently recognized genera however warrant a more detailed discussion, which is presented below. Bradley & Galil (1977: 188) established Hesperophasmatini to replace Haplopodini Günther, 1953. Zompro (2004: 135) removed the genera Aploploides Rehn & Hebard, 1938, Haplopus Burmeister, 1838 and Diapherodes Gray, 1835 and erroneously placed them in the tribe Cranidiini (→ 4.2.3). The genera Hesperophasma Rehn, 1901, Lamponius Stål, 1875, Rhynchacris Redtenbacher, 1908 and Taraxippus Moxey, 1971 were retained in Hesperophasmatini by Zompro (2004: 139), who however omitted Agamemnon Moxey, 1971 and erroneously synonymised Hypocyrtus Redtenabcher, 1908 with Lamponius (see below). Examination has shown the monotypical genus Laciphorus Redtenbacher, 1908 (Type-species: Laciphorus lobulatus Redtenbacher, 1908: 351) from Peru is not a member of Hesperophasmatini but belongs in Diapheromerinae: Diapheromerini, hence is here removed from Hesperophasmatini (→ 8.3). A redescription of Laciphorus and a discussion of its systematic position within Diapheromerini by the authors is in progress (Hennemann & Conle, in preparation). The monotypical Tersomia Kirby, 1904 (Type-species: Tersomia brasiliensis Kirby, 1904) from Brazil is obviously misplaced and here removed from Hesperophasmatini. Females, the only sex known, are large and very slender, typically stick-like insects with a long, spatulate subgenital plate, a very short median segment, very long and slender, unarmed legs and a pair of horns between the eyes. These features as well as the remarkably short antennae, which consist of no more than 23 segments, and morphology of the genitalia place Tersomia in Heteronemiidae: Heteronemiinae: Paraleptyniini (→ 8.3). Although lacking an area apicalis, Zompro (2004: 139) transferred Hesperophasmatini from the anareolate Phasmatidae: Cladomorphinae to the areolate Pseudophasmatidae: Xerosomatinae. The author stated they simply represented derived Xerosomatinae, in which the area apicalis is reduced, and postulated Hesperophasmatini were a subordinate taxon of Xerosomatinae. However, the very few features on which Zompro (2004: 139) based his action appear arbitrary (e.g. “eggs bullet-shaped”, “interodorsal carina of the protibiae lamelliform” and “mid- and hind legs agree completely”). Indeed, some Hesperophasmatini resemble certain representatives of Xerosomatinae: Xerosomatini at first glance, but presuming a close relation between these two tribes mostly based on convergences such as a similar general resemblance, basally curved and compressed profemora (a character widely distributed throughout the entire Phasmatodea and also typical for all members of the family Phasmatidae and subfamily Cladomorphinae respectively) and trapezoidal meso- and metafemora is the result of superficial examination. The fact that Xerosomatini (e.g. Creoxylus Audinet-Serville, 1838 or Xylospinodes Zompro, 2004) of similar habitus occur in geographically close regions or even in the same habitats as certain Hesperophasmatini (e.g. Rhynchacris in Central America) is not at all surprising. It merely shows the “similar resemblance” discussed by Zompro (2004: 139) are convergent developments caused by similar or identical evolutional pressure in the common habitats of these taxa. Detailed examination and comparison of several representatives from both groups undertaken in the course of this work have revealed numerous morphological characters of the insects and eggs that clearly separate Hesperophasmatini not only from Xerosomatinae but from the entire family Pseudophasmatidae. This is clearly


HENNEMANN ET AL.

supported by phylogenetic studies based on external morphology (Bradler, 2009: 99) and molecular data (Whiting et al., 2003; Buckley et al., 2009; Buckley et al., 2010). Bradler (2009: 99) furthermore stated that he could not find any synapomorphies that Lamponius Stål, 1875 (the only genus of Hesperophasmatini this author examined) had in common with genera of Xerosomatinae and that none of the apomorphies of the family Pseudophasmatidae were present in this Hesperophasmatini genus. Consequently, Zompro's placement of Hesperophasmatini as a subordinate taxon of Pseudophasmatidae: Xerosomatinae cannot be supported and the tribe is here re-transferred back to Cladomorphinae, where it was originally placed by Günther (1953: 557). In fact, close relation to Pterinoxylini n. trib. and Haplopodini is obvious and the most important characters that support the position within Cladomorphinae are discussed in more detail below. Hesperophasmatini is a principally Antillean taxon, with only two genera represented in Central America (Hypocyrtus Redtenbacher, 1908 and Rhynchacris Redtenbacher, 1908). The vast number of taxa is distributed througout the West Indies and as yet only a fraction of the tribe's true diversity is known. The authors have at hand numerous new species and genera mostly from Hispaniola, which multiply the number of currently known taxa. No representatives of Hesperophasmatini are so far known from the Bahamas. The northernmost distributed representative of this tribe, the Central American Hypocyrtus deserves special mention, since the genital morphology of ♂♂ differs considerably from that of all other members of Hesperophasmatini. While there is a well-developed and sclerotized vomer (Figs. 31–33) in ♂♂ of all other currently known genera, it is strongly reduced or missing and concealed by the paraprocts (= sternum XI) in Hypocyrtus (Hennemann & Conle, 2012: 66, see Fig. 34). Furthermore, the anal segment is much more tectiform than in the other genera and has the interior portions of the posterior margin facing each other and armed with prominent in-curving spines (Fig. 35). In all other genera, which do have a well-developed vomer, the posteromedian portion is merely incised and interiorly set with a variable number of small denticles. These striking genital differences are particularly remarkable, since the ♀♀ genitalia perfectly match with those of other Hesperophasmatini as does the insect and egg-morphology. Despite these striking differences of the external ♂♂ genital morphology and other obvious distinguishing features Zompro (2004: 140) erroneously synonymised Hypocyrtus with the exclusively Caribbean Lamponius. This misinterpretation was corrected by Eilmus (2009: 30), who re-established Hypocyrtus. This was confirmed by Hennemann & Conle (2012), who provided a detailed revision of the genus at the species-level. The Central American Rhynchacris and the Antillean Agamemnon are the only two genera whose ♀♀ exhibit a beak-like secondary ovipositor, that is formed by the elongated subgenital plate and an elongated epiproct. The eggs of both genera are more elongate and bullet-shaped than those of other genera and instead of simply being dropped to the ground are laid into a substrate (Figs. 25–26). Distribution: West Indies (excluding Bahamas) and Central America, ranging from Central Mexico as far south as the northern portions of Colombia. Below is a more detailed discussion of certain important characters, which distinguish and clearly exclude Hesperophasmatini from Pseudophasmatidae: Xerosomatinae: 1. Area apicalis: Zompro (2004: 139) stated that Hesperophasmatini strikingly “resemble” species of Xerosomatinae: Xerosomatini, and that the lack of an area apicalis on the tibiae was the only significant difference between these two tribes. The author postulated the area apicalis was merely reduced and hence interpreted the lack of this structure as an autapomorphy of Hesperophasmatini. However, as not only Xerosomatinae, but the entire family Pseudophasmatidae have a distinct area apicalis on all three tibiae and there are no transitions between these two extremes, Hesperophasmatini is, based solely on this feature, most unlikely to belong in Pseudophasmatidae. 2. Genitalia (♀♀): The genitalia of ♀♀ of Hesperophasmatini differ fundamentally from those of Xerosomatinae by having a distinctly keeled, scoop-like subgenital plate, which is tapered towards the apex and ± distinctly projects over the apex of the abdomen. Two genera (Agamemnon and Rhynchacris) even have the epiproct conspicuously elongated to form a beak-like secondary ovipositor together with the elongated subgenital plate. Very different specializations of the ♀ ovipositor are observed in Xerosomatinae. The subgenital plate is either strongly reduced, very small and scale-shaped (e.g. Acanthoclonia Stål, 1875, Metriophasma Uvarov, 1940), or slender, ± tube-like and down-curving with the lower gonapophyses enlarged and sclerotized (e.g. certain species of Creoxylus) forming a conspicuous derived appendicular ovipositor. The gonoplacs are lacking in Hesperophasmatini, while they are present and often conspicuously enlarged in representatives of Xerosomatinae (Bradler, 2009: 99). All mentioned features do not at all support the treatment of Hesperophasmatini by Zompro



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(2004: 139). Instead, the elongated subgenital plate and lack of gonoplacs are shared with Pterinoxylini n. trib. and Haplopodini and hence clearly prove the very close relation to these two tribes, which together form a supposedly monophyletic clade within Cladomorphinae (Fig. 409). 3. Genitalia (♂♂): Also the genital morphology of ♂♂ of Hesperophasmatini differs fundamentally from that of Xerosomatini. In Xerosomatini the anal segment is more or less distinctly tectiform and has the posteromedian portion sub-fissately divided with the bounding lips facing each other, labiately thickened and on their interior surfaces with several small in-curving teeth. The vomer is strongly reduced and missing (see Bradler, 2009: 99, figs. 22b–d). In contrast, Hesperophasmatini have the anal segment flattened and the thorn-pads at the posterior margin directed downward. The vomer is well developed, sclerotized and appearing as a roundly triangular plate usually with a single, prominent hook-like apex (Figs. 31–33, also see Bradler, 2009: 99, fig. 22a). An exception is represented by Hypocyrtus whose genital morphology resembles members of Xerosomatini by the tectiform anal segment and reduced vomer (see above, Figs. 34–35). However this genus perfectly matches with Hesperophasmatini in every other aspect. These striking morphological differences of the genitalia also do not support the systematic placement of Hesperophasmatini suggested by Zompro (2004: 139), but instead show remarkable affinity to Pterinoxylini n. trib. and Haplopodini and confirm the close relation to these three tribes here suggested (Fig. 409). 4. Sensory areas: Most of the currently known genera of Hesperophasmatini (exceptions are Lamponius and Taraxippus) possess a large and well developed sensory area on the profurcasternum, an important feature completely neglected and overlooked by Zompro (2004). Sometimes, two further much smaller sensory areas are present near the exavations of the procoxae on the probasisternum (e.g. Rhynchacris, Fig. 29). The sensory area of the profurcasternum is often very large, slightly convex and may cover a considerable part of the segments surface (Figs. 28, 30). It is formed by a cluster of small granules or wart-like humps, each of which bears 1–5 hollow, tubelike spines or papillate evaginations. The sensory areas on the probasisternum of e.g. Rhynchacris are much smaller, consist of a single hump-like swelling and usually have the papillate evaginations relatively longer. In Lamponius these swellings of the probasisternum are present but lack the evaginations seen in Rhynchacris. The function of these sensory areas is as yet unknown, but most certainly they serve as tactile or olfactory organs, since the live insects are frequently observed rubbing their prosternum on various surfaces, such as twigs or foliage of the host plant. In addition to several other common features the presence of sensory areas on the profurcasternum and probasisternum support close relation to Pterinoxylini n. trib., which corresponds to the results presented by Bradler (2009) and Buckley et al. (2009) (→ 4.2.5). In Pterinoxylini n. trib. however the sensory area of the profurcasternum is rather small, elliptical and strongly convex and the evaginations are considerably shorter than in members of Hesperophasmatini, being roughly cone-shaped with a deep central pit. Similar sensory areas may be present on the profurcasternum of certain members of Xerosomatini (e.g. some species of Acanthoclonia or Creoxylus), while there are never sensory areas on the probasisternum. The sensory areas on the profurcasternum of certain Xerosomatini-members are however rather different in structure and in contrast to Hesperophasmatini cover almost the complete surface of the segment, being formed by a cluster of densely arranged small, wart-like swellings, which lack any kind of evaginations or papillate structures. The different structure of the sensory areas do also not support the position of Hesperophasmatini supposed by Zompro (2004: 139). An ultrastructural study of these sensory areas is necessary for any broader discussion on their function or importance for phylogeny (Bradler, 2009: 32). 5. Egg-morphology: Zompro (2004: 139) stated the eggs of Hesperophasmatini and Pseudophasmatidae: Xerosomatinae were similar in the aspect that they are “bullet-shaped”, and the micropylar plate is small, placed roughly in the “centre of the capsule” [certainly meaning centre of the dorsal egg surface] and projects from the capsule surface. These observations are surprising since eggs of Xerosomatini are never “bullet-shaped” as stated by Zompro (2004: 139) but ± ovoid. Eggs of Hesperophasmatini are variable in shape, ranging from distinctly elongate and bullet-like with the polar-area more or less decidedly tapered over ovoid to almost spherical. In Xerosomatinae the capsule surface is merely covered by ± distinctly raised, net-like structures, while the capsule surface and operculum are to a variable degree covered with conspicuous hairy structures in all currently known genera of Hesperophasmatini (Figs. 24–27). Xerosomatinae have a small, roughly circular to elliptical micropylar plate, which distinctly projects from the capsule surface and is open internally with a rather narrow posteromedial gap and a distinct median line (Fig. 51). In contrast, the micropylar plate of Hesperophasmatini never projects considerably from the capsule surface, is relatively larger and variable in shape, being slightly transverse and


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shield-shaped to distinctly longer than wide with the anterior end narrowed and the posterior portion broadened. Internally the plate is open with a wide, triangular notch but without a median line (Figs. 47–48). Consequently, also the egg-morphology does not support the relation between Hesperophasmatini and Xerosomatinae postulated by Zompro (2004: 139). Instead, the lack of a median line is shared with Pterinoxylini n. trib. and Haplopodini and also confirms the close relation to these two tribes revealed in the present study and also suggested by the results of Bradler (2009) and Buckley et al. (2009).

FIGURES 31–36. Tribes Hesperophasmatini Bradley & Galil, 1977 and Pterinoxylini n. trib., genitalia of ♂♂. 31. Lamponius portoricensis Rehn, 1903: ventral view of anal segment [coll. FH, No. 0467-6]; 32. Agamemnon cornutus (Burmeister, 1838): ventral view of anal segment [coll. FH, No. 0589-6]; 33. Rhynchacris ornata Redtenbacher, 1908: ventral view of anal segment [coll. FH, No. 0134-32]; 34. Hypocyrtus scythrus (Westwood, 1859): ventral view of anal segment [coll. FH, No. 0638-7]; 35. Hypocyrtus scythrus (Westwood, 1859): caudal view of anal segment [coll. FH, No. 0638-7]; 36. Pterinoxylus crassus Kirby, 1889: ventral view of anal segment [coll. FH, 0588-7].

Genera included: 1. Agamemnon Moxey, 1971: 71. Type-species: Agamemnon iphimedeia Moxey, 1971: 75, by original designation of Moxey, 1971: 71. 2. Hesperophasma Rehn, 1901: 271. Type-species: Phantasis saussurei Bolivar, 1888: 137, by subsequent designation of Kirby, 1904a: 343. [Replacement name for Phantasis Saussure, 1870] = Phantasis Saussure, 1870: 188. Type-species: Phantasis saussurei Bolivar, 1888: 137, by subsequent designation of Kirby, 1904a: 343. 3. Hypocyrtus Redtenbacher, 1908: 355. Type-species: Hypocyrtus substrumosus Redtenbacher, 1908: 357 (= Hypocyrtus postpositus Redtenbacher, 1908), by subsequent designation of Zompro, 2000: 95. 4. Lamponius Stål, 1875: 19, 74. Type-species: Pygirhynchus guerini Saussure, 1868: 64, by monotypy. = Antillophilus Carl, 1913: 38. Type-species: Antillophilus brevitarsus Carl, 1913: 38 (= Lamponius guerini (Saussure, 1868), by monotypy. 5. Rhynchacris Redtenbacher, 1908: 354. Type-species: Rhynchacris ornata Redtenbacher, 1908: 354, by monotypy. = Pseudoceroys Hebard, 1922: 354. Type-species: Pseudoceroys harroweri Hebard, 1922: 355, pl. 15: 1–2 by original designation. [Synonymised by Hennemann & Conle, 2012: 81]



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6. Tainophasma Conle, Hennemann & Perez-Gelabert, 2014: 29. Type-species: Tainophasma monticola Conle, Hennemann & Perez, Gelabert, 2014: 31, by original designation. 7. Taraxippus Moxey, 1971: 67. Type-species: Taraxippus paliurus Moxey, 1971: 70, by original designation of Moxey, 1971: 67.

4.2.5. Pterinoxylini n. trib. (Figs. 36–44, 46) Type-genus: Pterinoxylus Audinet-Serville, 1838: 226. Haplopodini Günther, 1953: 557 (in part). Hesperophasmatini Bradley & Galil, 1977: 188 (in part). Phibalosomini (Sectio V: Phibalosomata) Redtenbacher, 1908: 399 (in part).

Description: ♀♀/♂♂ (Figs. 37–38). Medium sized to large (body lengths: ♀♀ incl. subgenital plate 135.0–189.0 mm, ♂♂ 94.0–118.0 mm), elongate and moderately slender, stick-like, brown Cladomorphinae with distinct sexual dimorphism. ♂♂ with scale-like tegmina and alae exceeding abdominal tergum IV. ♀♀ brachypterous and alae with a tympanal area (= stridulatory organ) in the basal portion of the costal region; alae projecting over posterior margin of median segment. Body of ♂♂ cylindrical, of ♀♀ subcylindrical. Entire body surface ± prominently rugulose and wrinkled, thorax partly granulose or tuberculose. Mesothorax of ♂♂ occasionally spinulose. Head longer than wide, dorsoventrally flattened and ± parallel-sided. Vertex with ± prominent tubercles or spines and occasionally slightly convex (Figs. 39–40). No ocelli. A slender and transverse gula is present. Antennae moderately robust and not reaching (♀♀) or slightly projecting over posterior margin of mesothorax; consisting of 30–40 segments. Scapus dorsoventrally flattened and roundly deflexed laterally, pedicellus subcylindrical. Following antennomeres with a longitudinal median keel or bulge ventrally (Figs. 39–40). Pronotum rectangular, distinctly longer than wide and longer than head. Probasisternum with two rough sensory areas near lateral excavations of procoxae (Figs. 41, 43); profurcasternum with a raised, oval central sensory area (Fig. 42). Mesothorax elongate, at least 2x longer than head and pronotum combined (longer in ♂♂), parallel-sided. Metanotum quadrate to slightly transverse. Meso- and metasternum as well as pleurae simple. Tegmina ovate and scale-like with a rounded central hump; slightly projecting over posterior margin of metanotum. Anal region of alae dark grey to black with ± distinct transparent spots, markings or radially arranged stripes; in ♀♀ of a conspicuous convex contour when opened. Abdomen longer than head and thorax combined. Median segment about 2x longer than metanotum. Segments II–VI parallel-sided and either more than 2x longer than wide (♂♂) or just indistinctly longer than wide (♀♀). Tergum VII ± laterally expanded or with a lateral lobe (♀♀ in particular). VIII–X slightly narrower (♀♀) or a little broader than previous segments (♂♂). In ♀♀ tergites III–IX often with a posteromedian tubercle. Praeopercular organ of ♀♀ indistinct and usually formed by a median tubercle close to posterior margin of sternum VII. Anal segment quadrate to slightly transverse, the posterior margin rounded with a small median indentation (♂♂) or broadly emarginated (♀♀). Epiproct small and with a longitudinal median carina; posterior margin truncate. Vomer of ♂♂ well developed, sclerotized and broadly triangular with a single short terminal hook (Fig. 36). Cerci very small, subcylindrical and slightly in-curving. Gonapophyses VIII of ♀♀ short, not extending beyond anal segment. No gonoplacs. Subgenital plate of ♀♀ elongate, irregularly naviculate or spatulate and distinctly projecting over apex of anal segment; longitudinally keeled basally but broadened and flattened towards a rounded to angular apex. Poculum of ♂♂ gently convex, cup-like and with a median carina; posterior margin notched medially. Legs short and robust (♀♀ in particular) with more or less prominently expanded or lobed carinae. Protibiae with broad foliaceous lobes and expansions (♀♀ in particular). Meso- and metafemora and corresponding tibiae with a ± distinct sub-apical dorsal lobe or tooth. Profemora shorter than mesothorax, metafemora at best reaching half way along abdominal tergum IV. Profemora compressed and curved basally and distinctly triangular in cross-section with the anterodorsal carina conspicuously raised and ± lamellate; medioventral carina indistinct and slightly displaced towards anteroventral carina. Meso- and metafemora and all tibiae trapezoidal in cross-section, the two anterior carinae strongly approaching each other. Medioventral carina of meso- and metafemora indistinct, and unarmed or at best with a few minute spinules. Tibiae without an area apicalis. Medioventral carina of meso- and metatibiae conspicuously displaced towards the anteroventral carina in the median portion. Tarsi short and stout, probasitarsus with a distinct rounded dorsal lobe, that is formed by the melted dorsal carinae. Meso- and metabasitarsi simple, furcate dorsally with the two dorsal carinae strongly approaching each other but well separated, and no longer than following two tarsomeres combined.


HENNEMANN ET AL.

FIGURES 37–44. Tribe Pterinoxylini n. trib.. 32. Pterinoxylus crassus Kirby, 1889 ♀: captive reared from Martinique [coll. FH, No. 0588-1]; 33. Pterinoxylus crassus Kirby, 1889 ♂: captive reared from Martinique [coll. FH, No. 0588-12]; 34. Pterinoxylus crassus Kirby, 1889: head and pronotum of ♀ [coll. FH, No. 0588-1]; 35. Pterinoxylus crassus Kirby, 1889: head and pronotum of ♂ [coll. FH, No. 0588-7]; 36. Pterinoxylus crassus Kirby, 1889 ♂: Prosternum and profurcasternum showing sensory-areas (SA); 37. Pterinoxylus crassus Kirby, 1889 ♂: sensory-area of profurcasternum; 38. P. crassus Kirby, 1889 ♂: left sensory area of prosternum; 39. Pterinoxylus crassus Kirby, 1889 egg (dorsolateral view): captive reared from Martinique [coll. FH, No. 0588-E].



29

FIGURES 45–51. Internal micropylar plates of Cladomorphinae and Xerosomatinae. 45. Tribe Haplopodini: Haplopus micropterus (St. Fargeau & Audinet-Serville, 1825); 46. Tribe Pterinoxylini n. trib.: Pterinoxylus crassus Kirby, 1889; 47. Tribe Hesperophasmatini: Lamponius guerini (Saussure, 1868); 48. Tribe Hesperophasmatini: Rhynchacris ornata Redtenbacher, 1908; 49. Tribe Cranidiini: Cranidium gibbosum (Burmeister, 1838); 50. Tribe Cladomorphini: Cladomorpus rubus (Saussure, 1861); 51. Pseudophasmatidae: Xerosomatinae: Xerosomatini: Creoxylus spinosus (Fabricius, 1793).

Eggs (Figs. 44 & 46): Very large (overall length > 8.0 mm), alveolar, at least 3x longer than wide and cylindrical to oval in cross-section. Entire capsule surface minutely punctured and strongly sculptured, with prominent raised ridges and wrinkles. Polar-area with a more or less prominent, hollow peripheral extension. Operculum circular and with a peripheral extension on outer margin similar to that of the polar-area. Micropylar plate elongate, oval and pointed anteriorly; covering about ¼ of capsule length. Micropylar cup distinct. Internal micropylar plate open with a wide, triangular posteromedian gap. No median line (Fig. 46). Differentiation (Table 2): Pterinoxylini n. trib. is closely related to Hesperophasmatini and Haplopodini, with which the increasingly shortened gonapophyses VIII and lack of gonoplacs of ♀♀ and lack of a median line in the eggs are shared. The presence of sensory areas on the probasisternum and profurcasternum are shared with Hesperophasmatini and support a sister-group relationship with this tribe.


HENNEMANN ET AL.

The new tribe is characterised by: the distinctly triangular cross-section of the profemora; broadly expanded and lamellate carinae of the protibiae; dorsally lobed probasitarsus; longitudinal carina along the ventral surface of the basal antennomeres; distinctly displaced medioventral carina of the meso- and metatibiae (towards anteroventral carina); strongly laterally deflexed scapus; presence of a gula and possessing rough, oval sensory areas on both the probasisternum and profurcasternum of both sexes (Figs. 41–43). The presence of a gula is most certainly a plesiomorphic character state, because it is present in all three tribes of Cladomorphinae sensu stricto, and readily distinguishes Pterinoxylini n. trib. from Hesperophasmatini and Haplopodini. Females are furthermore characterised by the conspicuous tympanal area (= stridulatory organ) in the basal portion of the alae (see discussion below), which seems to be an autapomorphy of the tribe because it is unique within the entire Cladomorphinae. Males are well recognized by the colouration of the anal region of the alae, which is grey to black and furnished with a variable number of transparent oval spots (Fig. 38). The eggs readily differ from those of Hesperophasmatini and Haplopodini by their remarkable size, very elongate and alveolar capsule which is at least 3x longer than wide, as well as the hollow, peripheral or crest-like extensions of the polar-area and operculum (Fig. 44). In addition to characters mentioned above the new tribe also differs from Hesperophasmatini by the much larger size as well as the considerably more elongate and slender body (♀♀ in particular). Sensory areas on the probasisternum and profurcasternum are also present in most Hesperophasmatini (→ 4.2.4), but the evaginations are much shorter, rather cone-shaped and have a deep central pit. Comments: The striking genus Pterinoxylus Audinet-Serville, 1838 was previously included in Haplopodini by Günther (1953) or Hesperophasmatini by Bradley & Galil (1977) and all subsequent authors. Zompro (2004: 139) disregarded the genus in his newly proposed arrangement of the New World anareolate Phasmatodea. Detailed investigation of Pterinoxylus and comparison with Hesperophasmatini has revealed several characters that clearly separate the genus from this tribe. Consequently, Pterinoxylus is here removed from Hesperophasmatini and placed in a tribe of its own, the Pterinoxylini n. trib.. A detailed comparison and distinction from Hesperophasmatini and Haplopodini is provided in Table 2 below. Distribution: Northern half of South America, lower Central America and Lesser Antilles (Dominica, Martinique & Saint Lucia). The following three interesting and taxonomically important key features warrant a more detailed recognition: 1. Stridulatory organ. If disturbed, live ♀♀ produce a stridulating sound, which is produced by scraping the rough basal portion of the alae by the down-curving posterior margin of the tegmina. The basal half of the prominent radial vein of the alae is dorsally covered with numerous very minute rasp-like tubercles, and together with the median and cubital vein forms a conspicuous more or less oval, sub-basal membrane or typhanal area, which is of a considerably thinner diameter than the rest of the costal region. In scraping these rasps by the sharp and hard posterior edge of the tegmina, a rustling sound is produced that is presumably increased by vibration of the tympanal area of the alae. In addition, the convex and dome-like shape of the tegmina may serve to increase the sound by resonance. This stridulating sound has so far only been reported to be produced by ♀♀, but if the alae of ♂♂ are spread a similar but much more slight sound can be observed. As in ♀♀ the radial vein of the alae is very prominent in its basal portion and covered with the same rasp-like tubercles, but interestingly no tympanal area is developed, perhaps the reason why the sound is by far not as loud as in ♀♀. 2. Alae of ♀♀: In addition to exhibiting a stridulatory organ in the costal region of the alae, ♀♀ of Pterinoxylus are typical for having the anal region of a conspicuously convex contour. This makes the conventional spreading of the wings in preserved specimens almost impossible, due to the outer margin curves increasingly downward as the wing is opened. This has already been pointed out by Rehn (1957: 188, footnote 9). 3. Sensory areas. The probasisternum and profurcasternum of both sexes show rough sensory areas very similar in structure to those seen in the closely related Hesperophasmatini, and a priori are homologous to the Oriental Heteropterygidae and certain Neotropical Pseudophasmatidae: Xerosomatinae: Xerosomatini. The probasisternum bears two sensory areas, one each at the excavation of the procoxae (Figs. 41, 43), and the profurcasternum has one central sensory area, which is much larger than those of the probasisternum and easily seen without magnification (Fig. 42). On the probasisternum they are formed by a conspicuous, convex and slightly transverse, white hump which dorsally bears numerous minute, cone-like evaginations each of which has a central pit or hollow. These evaginations are circular in cross-section, pale to mid brown basally and dark brown at the



31

apex. Occasionally, there are two further, much smaller sensory areas interior of the two large outer ones. The prominent sensory area of the profurcasternum is of a slightly different structure, being formed by an oval, central cluster of rounded, whitish granules each of which bear a cluster of 2–6 of the same cone-like evaginations seen on the sensory areas of the probasisternum (Fig. 42). The function of these sensory areas is still unknown, but due to the central hollow seen in each of the numerous small evaginations, they appear to serve as tactile or olfactory organs. An ultrastructural study of these organs appears necessary for any broader discussion (Bradler, 2009: 32). Genus included: 1. Pterinoxylus Audinet-Serville, 1838: 226. Type-species: Pterinoxylus difformipes Audinet-Serville, 1838: 227 (= Haplopus eucnemis Burmeister, 1838: 577), by monotypy.

TABLE 2. Comparison of Haplopodini, Pterinoxylini n. trib. and Hesperophasmatini. Haplopodini

Pterinoxylini n. trib.

Hesperophasmatini **

Body length (♀♀)

50.0–197.0 mm

135.0–189.0 mm

36.0–93.0 mm

Body length (♂♂)

60.0–126.0 mm

94.0–118.0 mm

31.0–87.0 mm

Cylindrical to oval (triangular*)

± cylindrical

Oval, dorsoventrally compressed

± glabrous; mostly colourful insects

Not glabrous & strongly sculptured; brown

Not glabrous & strongly sculptured; mostly brown

No

Yes

No

Cylindrical

In basal portion with a longitudinal carina ventrally

Cylindrical to oval

Oval

Strongly dorsoventrally flattened and dilated

Dorsoventrally flattened

Probasisternum

No sensory areas

With paired sensory areas (Figs. 41 & 43)

Sometimes with paired sensory areas (Fig. 29)

Profurcasternum

No sensory area

With a central sensory area (Fig. 41–42)

Mostly with a central sensory area (Figs. 28 & 30)

-

Short and cone-shaped with a deep central pit or hollow

Elongate, papillate or tubelike

Wings (♀♀)

Apterous or brachypterous

Brachypterous; alae with a tympanal region (= stridulatory organ) in basal portion

Apterous, rarely with vestigial, scale-like alae

Colouration of alae (♂♂)

Plain or reticulate; transparent, pink or orange

Dark grey to black with transparent spots (Fig. 38)

Transparent; reticulate or with faint greyish mottling (may be red in Hypocyrtus)

Distinct, ± spinose

Indistinct, smooth or minutely spinose

Indistinct or lacking; may be spinose apically

Protibiae

Slender; sometimes with single, small lobes dorsally

All carinae ± expanded and lamellate or lobate

Slender; sometimes with irregular rounded lobes/ lobules

Meso- and metatibiae

Medioventral carina central

Medioventral carina lamellate and strongly displaced towards anteroventral carina in median portion

Medioventral carina central

Body cross-section (♀♀) Body surface (♂♂) Gula Antennae (cross-section) Scapus (cross-section)

Evaginations of sensory areas

Medioventral carina of meso- and metafemora



HENNEMANN ET AL.

TABLE 2. (Continued) Haplopodini

Pterinoxylini n. trib.

Hesperophasmatini **

Probasitarsus (♂♂)

Slender and furcate dorsally; the two dorsal carinae clearly separated

Carinate dorsally; the two dorsal carinae melted with another a forming a raised, rounded lobe

Slender and furcate dorsally; the two dorsal carinae clearly separated

Egg (Shape of capsule)

Ovoid to barrel-shaped; < 2x longer than wide

Alveolar with a hollow, peripheral polar extension; > 3x longer than wide (Fig. 44)

Almost spherical to bulletshaped; < 2.5x longer than wide (Figs. 24–27)

Egg (Operculum)

With a conical or knob-like central elevation or tuberculose with a raised circular rim of tubercles

With a prominent, hollow peripheral or crest-like extension on outer margin (Fig. 39)

With a circle of hair-like structures

Distribution

West Indies, Florida Keys & Honduras (Fig. 52)

Northern half of South America, lower Central America & Lesser Antilles

West Indies & Central America

* Paracranidium Brock, 1998 ** The distinction of this tribe can only be preliminary and is solely based on the few genera currently known (à see comments on Hesperophasmatini above)

5. REVISION OF THE TRIBE HAPLOPODINI GÜNTHER, 1953 Haplopodini Günther, 1953 rev. stat. Type-genus: Haplopus Burmeister, 1838: 576. Haplopodini Günther, 1953: 557 (in part). Bacteridae (Sectio: Bacteriae), Brunner v. Wattenwyl, 1893: 80, 83 (in part). Bacteriinae, Kirby, 1904a: 431 (in part). Bacterinae, Rehn, 1904: 61 (in part). Cladoxerinae, Karny, 1923: 130 (in part). Cranidiini, Zompro, 2005: 135 (in part). Otte & Brock, 2005: 32 (in part). Hesperophasmatini Bradley & Galil, 1977: 188 (in part). Phibalosomini (Sectio: Phibalosomata), Redtenbacher, 1908: 399. Phibalosominae, Shelford, 1908: 343, 355 (in part). Phibalosomatinae, Moxey, 1971: 44 (in part).

Description: ♀♀, ♂♂: Small to very large (body lengths: ♀♀ incl. subgenital plate 50.0–197.0 mm, ♂♂ 60.0–126.0 mm), moderately slender to very robust and massive (♀♀ in particular) Cladomorphinae with long but thickened antennae. ♀♀ either various shades of brown and grey or bright green, ♂♂ often very colourful insects. Body cylindrical in ♂♂ and cylindrical to oval in ♀♀; in one case triangular in cross section with a prominently raised longitudinal median keel along entire dorsal body surface. Body often ± glabrous (♂♂ in particular). ♀♀ mostly brachypterous or with vestigial wings, rarely apterous, ♂♂ with scale-like tegmina and alae either well developed or shortened but rarely apterous. Head indistinctly longer than wide with the vertex ± convex and bispinose or bi-cornute; rarely flat and unarmed. No gula. No ocelli. Antennae long and projecting over mesothorax (♀♀) or median segment (♂♂); cylindrical and consisting of 55–80 segments. Antennomeres shortened indistinctly longer than wide and increasingly perlamorph; scapus slightly compressed dorsoventrally. Pro- and mesothorax to a variable degree armed with granules, tubercles or spines, rarely unarmed. Pronotum longer than wide and narrower than head. Probasisternum or profurcasternum without sensory areas. Mesothorax elongate and cylindrical (♂♂) or ± widened towards the posterior (♀♀). Mesonotum often with a cluster of tubercles or spines near anterior margin; in one case with a prominently semi-circularly raised longitudinal median keel. Mesopleurae



33

of ♀♀ often ± expanded and armed with a longitudinal marginal row of tubercles, teeth or spines. Mesosternum simple. Median segment longer than metanotum (only in one case slightly shorter than metanotum). Abdomen longer than head and thorax combined. Tergites mostly unarmed but II–IV in ♀♀ may bear longitudinal carinae or single tooth- or crest-like lobes. Abdomen of ♀♀ ± swollen and broadened basally or pre-medially, segments II–VII of ± uniform width in ♂♂. Praeopercular organ of ♀♀ indistinct and formed by a longitudinal ridge or tubercle near posterior margin of sternum VII; in one case formed by a cluster of three granules (Figs. 326–343). Anal segment of ♂♂ slightly tectiform with the posterior margin rounded or roundly indented medially; posterior margin ventro-medially with a variable number of small in-curving denticles. Cerci at best equal in length to anal segment, cylindrical to oval in cross-section and ranging from straight to slightly in-curving. Vomer well developed and sclerotized, either roundly triangular or with the base semi-circular and the single terminal hook strongly elongated and papillate (Figs. 344–364). Poculum of ♂♂ convex, cup-like to conical and often with a ± distinct basal swelling or spiniform projection. Gonapophyses VIII of ♀♀ short and fully hidden under anal segment; not or hardly longer than gonapophyses IX. Gonoplacs lacking. Epiproct small, either hidden under anal segment or slightly projecting; shape ranging from triangular to semi-circular. Subgenital plate of ♀♀ ± projecting over apex of abdomen, often lanceolate or naviculate and keeled longitudinally. Legs rather short and stocky (♀♀) to moderately long and slender (♂♂), at least with small sub-apical ventral spines on the meso- and metafemora but often with additional teeth, lobes or expansions. Profemora curved and compressed basally; cross-section slightly triangular with the anterodorsal carina ± raised. Meso- and metafemora and all tibiae trapezoidal in cross-section. Medioventral carina of meso- and metafemora distinct and usually armed with a longitudinal row of spines; slightly misplaced towards anteroventral carina in profemora. Tibiae without an area apicalis. Tarsi simple and slender, probasitarsus occasionally with a rounded lobe dorsally and with the two dorsal carinae melted with another. Meso- and metabasitarsus sulacte dorsally with the two dorsal carinae well separated. All basitarsi at best as long as following three tarsomeres combined.

FIGURE 52. Map showing the distribution of the tribe Haplopodini Günther, 1953

Eggs: Medium-sized to large, capsule ovoid or barrel-shaped. Capsule surface finely punctured to strongly coriaceous, rugulose or tuberculose. Micropylar plate at best ½ as long as capsule, shape variable. Internal


HENNEMANN ET AL.

micropylar plate open and with a widely triangular posteromedian notch, no median line (Fig. 45). Operculum either flat and with a circular rim of tubercles or ± conically raised to knob-like. Differentiation (Table 2): Closely related to Pterinoxylini n. trib. and Hesperophasmatini with which it shares the lack of gonoplacs and the short gonapophyses VIII of ♀♀, both characters of which are likely synapomorphies of these three tribes. On this basis and as discussed above Haplopodini possibly forms a monophyletic clade together with these two tribes within present Cladomorphinae sensu lato. The lack of sensory areas on the probasisternum and profurcasternum, increasingly shortened but greater number (55–80) of antennomeres and increasingly thickened, perlamorph antennae suggest Haplopodini is the sister-taxon of Pterinoxylini n. trib. + Hesperophasmatini (Fig. 409). While the lack of sensory areas most certainly is a plesiomorphic character, the morphology of the antennae is likely to be an autapomorphy of Haplopodini. The lack of a gula is here estimated as a synapomorphy of Haplopodini + Hesperophasmatini. A confirmed definition of these character states however deserves a comprehensive phylogenetic analysis of the subfamily Cladomorphinae and supposedly closely related taxa. Furthermore, Haplopodini differs from both tribes by the distinct medioventral carina of the profemora which is slightly displaced towards the anteroventral carina, distinct and often spinose medioventral carina of the mesoand metafemora of both sexes, as well as the ± glabrous body surface, mostly pretty colouration and transparent orange to pink anal region of the alae of ♂♂ (either plain or reticulate). From Pterinoxylini n. trib. it also differs by: the lack of a tympanal region in the basal portion of the alae (= stridulatory organ) of ♀♀; less distinctly triangular cross-section of the profemora; not dilated or lamellate carinae of the protibiae and ovoid to barrel-shaped, much smaller eggs which lack hollow, peripheral extensions on the polar-area and operculum (→ Table 2). From Hesperophasmatinim, with which they share the lack of a gula, members of Haplopodini may also be distinguished by the more decidedly triangular cross-section of the profemora, longer often lanceolate or naviculate subgenital plate of ♀♀ and lack of conspicuous hairy structures in the eggs. Also, Haplopodini on average are larger and less stocky than members of Hesperophasmatini. Comments: Günther (1953: 557) originally established the tribe Haplopodini to include some of the neotropical genera of Redtenbacher's Phibalosomatini: Sectio Phibalosomata (1908: 399) and Phibalosomatini: Sectio Eurycanthae (1908: 339), this is: Haplopus Burmeister, 1838, Diapherodes Gray, 1835, Pterinoxylus Audinet-Serville, 1838, Hesperophasma Rehn, 1904 and Lamponius Stål, 1875. Bradley & Galil (1977: 188) renamed Günther's tribe Hesperophasmatini and added some not closely related genera (Agamemnon Moxey, 1971, Taraxippus Moxey, 1971 and Aploploides Rehn & Hebard, 1838). Zompro (2004: 139) transferred Hesperophasmatini from Cladomorphinae to Pseudophasmatidae: Xerosomatinae and placed the genera Haplopus, Aploploides, Diapherodes and Paracranidium in the distinctive South American tribe Cranidiini Günther, 1953. However, these four Antillean genera differ significantly from Cranidium Westwood, 1843, the type-genus of Cranidiini, and are here removed from that tribe (→ 4.2.3, Table 1). Haplopodini Günther, 1953 is therefore reinstated to contain Haplopus, Aploploides, Diapherodes and Paracranidium as well as four newly described genera (rev. stat.). These share a good number of morphological characters, which sufficiently distinguish them from Cranidiini, Hesperophasmatini or Pterinoxylini n. trib. and obviously form a monophyletic clade, the tribe Haplopodini (→ Table 2). Distribution (Fig. 52): West Indies (Greater and Lesser Antilles, Bahamas), Florida Keys, Cayman Islands, Santanilla & Central America (Honduras). Most of the Haplopodini are distributed throughout the West Indies, with only one genus (Apteroplopus n. gen., Fig. 377) represented in Central America (eastern Honduras), hence this is a principally Antillean taxon. Haplopus is restricted to the Greater Antilles, Virgin Islands, St. Thomas, St. Croix, Florida Keys, Dry Tortugas and Bahamas and one species is found also on the Cayman Islands and Swan Islands (= Santanilla, Fig. 376). Parhaplopus n. gen. only contains three species, one of which is endemic to Cuba and the other two known from Hispaniola and Gonaive Island (Fig. 378). Cephaloplopus n. gen. is represented by three species on Hispaniola and has one species on Cuba (Fig. 377). Diapherodes is distributed throughout almost the complete Lesser and Greater Antilles, but has not yet been recorded from Hispaniola, Cuba or the Bahamas (Figs. 375, 381). Paracranidium is restricted to the mountainous regions of East-Jamaica (Fig. 378) and the two monotypical genera Aploploides and Venupherodes n. gen. are endemic to Cuba (Fig. 374).



35

Genera included: 1. Aploploides Rehn & Hebard, 1938: 49. Type-species: Aploploides stenocephalum Rehn & Hebard, 1938: 49, by original designation of Rehn & Hebard, 1938: 49. 2. Apteroplopus n. gen. Type-species: Dyme grosse-tuberculata Brunner v. Wattenwyl, 1907: 323, by present designation. 3. Cephaloplopus n. gen. Type-species: Cephaloplopus pulchellus n. sp., by present designation. 4. Diapherodes Gray, 1835: 13, 33. Type-species: Mantis gigantea Gmélin, 1789: 2055, by subsequent designation of Kirby, 1904a: 362. 5. Haplopus Burmeister, 1838: 576. Type-species: Cyphocrana micropterus St. Fargeau & Audinet-Serville, 1825: 445, by monotypy. = Aplopus Gray, 1835: 13, 34. 6. Paracranidium Brock, 1998: 28. Type-species: Diapherodes (Cranidium) pumilio Westwoood, 1843: 50, pl. 61: 2, by original designation of Brock, 1998: 28. 7. Parhaplopus n. gen. Type-species: Haplopus cubensis Saussure, 1868: 68, by present designation. 8. Venupherodes n. gen. Type-species: Platycrana venustula Audinet-Serville, 1838: 242, by present designation.

Keys to the genera of Haplopodini Günther, 1953 ♀♀* 1. 2. 3. 4. -

5. -

6.

-

Wings present . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4 Apterous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 Body length > 6 cm; dorsal body surface without a prominent keel nor tectiform; Cuba . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 Small insects (body < 6 cm); dorsal body surface strongly tectiform; mesonotum with a prominent, semicircularly raised longitudinal median keel (Fig. 287); Jamaica (Fig. 378) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Paracranidium Massive insects with thorax and abdomen considerably broadened (Fig. 314); head globose and indistinctly longer than wide; vertex bi-tuberculate (Fig. 318). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Venupherodes n. gen. Slender insects (Fig. 53); head elongate and about 2x longer than wide; vertex flat and unarmed (Fig. 58). . . . . . . Aploploides Slender insects with body cylindrical in cross-section; colour mostly brown or grey; mesothorax > 2.0x longer than head and pronotum combined; anal region of alae reticulate, with dark radial and transverse veins; not Lesser Antilles or Jamaica . . . . 5 Robust to very massive insects, with meso-, metathorax and basal portion of abdomen broadened; mostly bright green; body oval in cross-section; mesothorax < 2x longer than head and pronotum combined; anal region of alae plain pink or violet; Greater & Lesser Antilles (Fig. 375). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Diapherodes Large insects (body length > 10 cm); pair of spines on vertex extending < 2/3 the height of head capsule and with a single point; abdominal tergum VII at best gently expanded or with a small posterolateral lobe. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6 Smaller insects (body < 10 cm); pair of spines of the vertex extending by > 2/3 the height of head capsule, crenate or multituberculate; abdominal tergum VII prominently dilated, lateral lobes extending > ½ of body width; Cuba & Hispaniola (Fig. 377) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Cephaloplopus n. gen. Mesothorax slender and ± parallel-sided; mesonotum granulose or tuberculose; apex of subgenital plate narrow and ± pointed; probasitarsus slender; Greater Antilles, Dry Tortugas, Florida Keys, Bahamas, Cayman Islands and Swan Islands (Fig. 376) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Haplopus Mesothorax swollen medially; mesonotum armed with large, blunt spine-like tubercles; apex of subgenital plate rounded or broadly truncate; probasitarsus with a lobe dorsally; only Cuba & Hispaniola (Fig. 378). . . . . . . . . . . . . . Parhaplopus n. gen.

*♀♀ of Apteroplopus n. gen. are not known

♂♂* 1. 2. 3. 4. 5. 6.

Pterous or brachypterous; median segment longer than metanotum; West Indies . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 Apterous (Fig. 53); median segment less than half the length of the metanotum; Honduras (Fig. 377) . . . Apteroplopus n. gen. Head globose and indistinctly longer than wide; vertex armed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 Head elongate, about 2x longer than wide; vertex flat and unarmed; Cuba (Fig. 374) . . . . . . . . . . . . . . . . . . . . . . . Aploploides Spines of vertex extending < 2/3 the height of head capsule, with a single point; abdominal tergum VII parallel-sided or with a small posterolateral lobe . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4 Spines of the vertex extending by > 2/3 the height of head capsule, often crenate or multi-tuberculate; abdominal tergum VII strongly expanded with lateral lobe extending by > ½ of body width; Cuba & Hispaniola (Fig. 377) . . .Cephaloplopus n. gen. Mesothorax > 2x longer than head and pronotum combined; not Lesser Antilles or Jamaica . . . . . . . . . . . . . . . . . . . . . . . . . . . 5 Mesothorax < 2x longer than head and pronotum combined; Greater & Lesser Antilles (Figs. 367) . . . . . . . . . . . .Diapherodes Body surface dull; cerci slender, distinctly shorter than anal segment and laterally compressed; anal region of alae plain . . . . 6 Body surface glabrous; cerci obtuse, ± as long as anal segment and ± cylindrical in cross-section; anal region of alae reticulate (Fig. 247); Greater Antilles, Dry Tortugas, Florida Keys, Bahamas, Cayman Islands and Swan Islands (Fig. 376) . . . Haplopus Body length < 7 cm; head globose; pronotum with a pair of anterior spines (Fig. 319); mesonotum only with 2–4 paired spines near anterior margin; Cuba (Fig. 374). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Venupherodes n. gen.


HENNEMANN ET AL.

-

Body length > 8 cm; head with vertex at best gently rounded; pronotum unarmed; complete mesonotum armed with spines; Cuba & Hispaniola (Fig. 378) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Parhaplopus n. gen.

*♂♂ of Paracranidium Brock, 1998 are not known

Eggs* 1. 2. 3. 4. -

Capsule surface strongly sculptured; no raised central capitulum . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 Capsule surface minutely punctured or rugulose; operculum with a raised conical or knob-like central capitulum . . . Haplopus Micropylar plate < 1/3 the length of capsule. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 Micropylar plate > 1/3 the length of capsule. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4 Micropylar plate longer than wide; heart or drop-shaped; capsule irregularly granulose and covered with fine, raised ridges (Figs. 312-313) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Parhaplopus n. gen. Micropylar plate wider than long; shield-shaped; capsule very prominently and enevenly sculptured (Figs. 324–325). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Venupherodes n. gen. Large (capsule length > 3.5 mm). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Diapherodes Small (capsule length < 3.5 mm) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Cephaloplopus n. gen.

* eggs of Aploploides Rehn & Hebard, 1938, Apteroplopus n. gen. and Paracranidium Brock, 1998 are not known

5.1. Genus Aploploides Rehn & Hebard, 1938 Type-species: Aploploides stenocephalum Rehn & Hebard, 1938: 49, by original designation of Rehn & Hebard, 1938: 49. Aploploides Rehn & Hebard, 1938: 49. Bradley & Galil, 1977: 188. Moxey, 1972: 54 (in litt.). Bragg, 2001: 628. Zompro, 2004: 305. Otte & Brock, 2005: 49.

Description: ♀♀, ♂♂ (Figs. 53–54). Rather small, elongate and slender Haplopodini (body lengths: ♂♂ 73.0–78.0 mm, ♀♀ 87.6–95.0 mm), ♂♂ with fully developed alae, ♀♀ apterous. Body subcylindrical with entire surface smooth and shiny (except for very minute granules on the vertex, pronotum and along lateral margins of the mesonotum). General colouration greenish. Head elongate, 2x longer than wide, vertex flat and smooth; cheaks gently widened (Fig. 58). Eyes small, subcircular. Antennae filiform, longer than head and complete thorax combined in ♂♂, shorter than head, pro and mesonotum combined in ♀♀. Scapus dorsoventrally compressed, pedicellus subcylindrical. Pronotum slightly shorter and narrower than head, rectangular. Mesothorax elongate, about 1.5x longer than head and pronotum combined. Mesonotum parallel-sided and with a fine longitudinal median carina. Mesosternum simple. Metanotum rectangular. Meso- and metapleurae unarmed. Tegmina of ♂♂ small and scale-like, alae reaching about half way along tergum VI. Anal region transparent. Median segment rectangular and slightly longer than metanotum. Abdomen elongate. Segments II–VI of ♂♂ parallel-sided, of equal width and about 3x longer than wide; of ♀♀ just slightly longer than wide. Segment VII of ♀♀ longer than previous and slightly narrowing towards the posterior, VIII–X shorter than previous and tapering to apex. Sternum VII of ♀♀ with a faint praeopercular organ, formed by a median tubercle close to posterior margin. Anal segment of ♂♂ with a faint posteromedian indentation, of ♀♀ bilobate. Epiproct very small, triangular. Cerci slender, elongate, very weakly incurving and tapered towards the apex (Figs. 55–56). Poculum of ♂♂ indistinct and scooplike, just reaching posterior margin of tergum IX (Fig. 57), posterior with a slight median indentation. Vomer broadly triangular, somewhat narrowed in apical half and with a short and blunt terminal hook (Fig. 344). Subgenital plate of ♀♀ elongate, keeled and extending beyond apex of abdomen by about 2/5 of its total length, apex slightly rounded (Figs. 55–56). Legs of moderate length in ♂♂, and comparatively short in ♀♀ the profemora being shorter than the mesonotum and hind legs (incuding tarsi) reaching to posterior of tergum VI. Profemora strongly compressed and curved basally, with the medioventral carina slightly displaced towards anteroventral carina. Medioventral carina of meso- and metafemora minutely spinulose, anteroventralcarina bispinose subapically. Basitarsi short, at most as long as combined length of following two tarsomeres, simple. Eggs unknown.



37

Differentiation: Easily distinguished from all other genera of Haplopodini by the following characters: head elongate 2x longer than wide vertex flat and smooth; antennae of ♀♀ shorter than head and thorax combined; body surface entirely smooth and glabrous (except for very minute granules on the vertex, pronotum and along lateral margins of the mesonotum); legs very weakly armed; ♀♀ apterous. The lack of wings in ♀♀ is shared with the Cuban Venupherodes n. gen. and Jamaican Paracranidium Brock, 1998, but the first genus differs at once by the globose and bi-cornute head, and the latter by the triangular crosssection of the body and prominent, semicircularly raised median keel of the mesonotum. Distribution (Fig. 374): Northwest Cuba (Pinar del Rio Province), endemic.

FIGURES 53–58. Aploploides stenocephalum Rehn & Hebard, 1938. 53. ♀ PT: NW-Cuba, Pinar del Rio [ANSP]; 54. ♂ PT: NW-Cuba, Pinar del Rio [ANSP]; 55. ♀ apex of abdomen, lateral view; 56. ♀ apex of abdomen (dorsal view); 57. ♂ apex of abdomen (lateral view); 58. ♀ PT head in lateral view: Cuba, Pinar del Rio [ANSP].


HENNEMANN ET AL.

Species included: 1. Aploploides stenocephalum Rehn & Hebard, 1938: 49, pl. 4: 18–21 (♂, ♀).

Aploploides stenocephalum Rehn & Hebard, 1938 (Figs. 53–58, 334, 352, 374) Aploploides stenocephalum Rehn & Hebard, 1938: 49, pl. 4: 18–21 (♂, ♀). HT, ♂: Cuba, Pinar del Rio, Pinar del Rio Province, Sept. 14’13 F.E. Lutz, five kilometers distand on road to Coloma, river edge, area of bamboo, shrubs and Weeds; Aploploides stenocephalum R + H, Holotype [USNM]; AT, ♀: Cuba, 12 ½ K. S. of Pinar Rio, Sept. 23’13, F.E. Lutz, region of pines and palmettos, taken from pine, Aploploides stenocephalum R + H, Allotype [USNM]; PT, 1 ♀: Cuba, 12 ½ K. S. of Pinar Rio, Sept. 23’13, F.E. Lutz, region of pines and palmettos, taken from pine, Aploploides stenocephalum R + H, Paratype [USNM]; PT, ♂: Cuba, 12 ½ K. S. of Pinar Rio, Sept. 23’13, F.E. Lutz, region of pines and palmettos, taken from pine, Aploploides stenocephalum R + H, Paratype [USNM]; PT, ♀: Cuba, 12 ½ K. S. of Pinar Rio, Sept. 12–23’13, Aploploides stenocephalum R + H, Paratype, w0090, ANSP [ANSP]; PT, ♂: Aploploides stenocephalum R + H, Paratype Hebard CLN, Cuba /3., K. S. of Pinar Rio, Sept. 12–23’13, w0091, ANSP [ANSP]; PT, ♂: Aploploides stenocephalum R + H, Paratype Hebard CLN, Cuba /3., K. S. of Pinar Rio, Sept. 12–23’13, w0092, ANSP [ANSP]. Moxey, 1972: 55, fig. 11 (♀ anal segment; in litt.). Otte & Brock, 2005: 49.

Further material [1 ♂, 3 nymphs]: CUBA: 1 ♂: Col. F. de Zayas, Pinares de Vinales, V.1960, Prov. P.R. CUBA [USNM]; 1 ♂ (penultimate instar), 2 nymphs: Cuba, Cerro de Cabras ca. 400 ft., Pinar del Rio Province, Sept. 11’13, C.W. Leng, beaten from pine in a dense forest [USNM].

Comments: The original descriptions of Rehn & Hebard (1938) are very detailed and along with the illustrations provided (pl. 4, figs. 18–24) serve well to recognize this distinctive species. Furthermore, the diagnosis of the genus presented above is sufficient enough for differentiation. Hence, no descriptions are provided. The holotype has the terminal abdominal segments destroyed by pests. Distribution (Fig. 374): Western Cuba (Pinar del Rio Province: Pinar del Rio [ANSP; USNM]; Cerro de Cabras [USNM] & Pinares de Viñales [USNM]). Number of specimens examined: 11 TABLE 3. Measurements of Aploploides stenocephalum Rehn & Hebard, 1938 [mm]. ♂, HT [USNM]

♀, AT [USNM]

♂♂

♀♀

-

91.5

-

88.0–95.0

Body

73.0

87.6

70.0–78.0

81.5–88.0

Pronotum

2.7

4.2

2.7–2.8

4.0–4.4

Mesonotum

10.8

15.9

11.4–14.0

15.0–16.3

Metanotum

5.4

7.2

5.2–5.7

6.5–7.5

Median segment

7.6

6.1

7.1–7.9

5.6–6.5

Subgenital plate

-

20.2

-

19.0–21.0

Tegmina

9.1

-

9.1–10.0

-

Alae

28.0

-

35.0–40.5

-

Profemora

13.8

13.7

14.1–14.6

12.9–14.6

Mesofemora

10.5

11.5

10.0–11.3

10.8–12.0

Metafemora

15.0

16.8

14.0–16.2

15.2–17.0

Protibiae

-

-

11.8–13.0

9.8–11.0

Mesotibiae

-

-

8.5–9.4

8.7–9.5

Metatibiae

-

-

12.0–13.0

13.0–14.3

Antennae

-

-

32.0–36.0

> 25.0

Body (incl. sg. pl.)



39

5.2. Genus Apteroplopus n. gen. Type-species: Dyme grosse-tuberculata Brunner v. Wattenwyl, 1907: 323, by present designation. Dyme, Brunner v. Wattenwyl, 1907: 318 (in part—not Dyme Stål, 1875). Bacteria, Otte & Brock, 2005: 64 (in part).

Description: ♂♂ (Fig. 59). Large (body length 115.4 mm) and very slender, apterous Haplopodini. Body surface strongly shiny and smooth except for some paired tubercles on the mesonotum and minute granules on the mesoand metapleurae. Head indistinctly longer than wide and gently narrowing towards the posterior, vertex very gently convex and with two small tubercles. Eyes prominent, circular and projecting hemispherical. Antennae conspicuously thickened and longer than head and complete thorax combined; > 50 antennomeres. Scapus 1.5x longer than wide, oval in cross-section. Pronotum rectangular and unarmed, slightly longer and as broad as head. Mesothorax very elongate, more than 3.5x longer than head and pronotum combined. Mesonotum parallel-sided and slightly broadened at the posterior, dorsally armed with several irregularly paired, blunt tubercles. Metanotum half the length of mesonotum, slightly broadened at anterior margin. Meso- and metasternum and pleurae simple, the latter very minutely granulose. Abdomen about as long as head and complete thorax combined. Median segment about ¼ the length of the metanotum. Abdominal segments II–VII slightly decreasing in length, at least 2.5x longer than wide and parallel-sided. Tergum VIII 2/3 the length of VII, swollen and roundly dilated subapically; distinctly broader than II–VII and broadest segment. Posterior margin of anal segment with a wide concave excavation, the posterolateral angles thickened (Fig. 61). Cerci of moderate size, laterally compressed, slender, slightly incurving and tapered towards the apex; projecting over apex of anal segment (Figs. 60–61). Vomer distinct, triangular with a single terminal hook. Poculum moderately convex, scoop-like and with a slight posteromedian indentation; just reaching to posterior margin of tergum IX (Fig. 60). Legs long and slender, profemora and mesofemora shorter than mesothorax, hind legs almost reaching apex of abdomen. Profemora strongly compressed and curved basally, the meso- and metafemora very slightly constricted towards the base. Anterodorsal carina of meso- and metafemora slightly elevated sub-apically. Medioventral carina of profemora very indistinctly displaced towards anteroventral carina, with two minute teeth apically. Outer ventral carinae of meso- and metafemora with 2–4 sub-apical spines. Medioventral carina of meso- and metafemora with 3–4 spines sub-apically. Tibiae unarmed. Tarsi elongate, probasitarsus as long as following three tarsomeres combined with the dorsal carina slightly elevated towards the apex. Meso- and metabarsitarsi as long as following two tarsomeres combined. ♀♀ and eggs unknown. Differentiation: ♂♂ are easily distinguished from all other genera of Haplopodini by lacking tegmina and alae, very elongate mesothorax which is more than 3.5x longer than the head and pronotum combined, and having the median segment considerably shortened, being less than 1/3 the length of the metanotum. Furthermore, this is the only representative of Haplopodini to be known from Central America. From the closely related Haplopus Burmeister, 1838 ♂♂ may additionally be distinguished by the smaller and more slender, laterally compressed and apically tapered cerci. Comments: Dyme sensu Brunner v. Wattenwyl (not Stål, 1875) is a highly polyphyletic grouping and examination of his specimen of D. grossetuberculata has shown this clearly not to be a member of Dyme Stål, 1875. The glabrous body surface, conspicuously thickened antennae, apically spinose ventral carinae of the femora, slightly displaced medioventral carina of the profemora, tuberculate mesonotum as well as features of the genitalia place this species in the tribe Haplopodini. Features such as the complete lack of tegmina or alae, short median segment and very elongate mesothorax clearly show it to form a distinct generic unit, here named Apteroplopus n. gen.. Within Haplopodini the new genus appears to be close to Haplopus Burmeister, 1838, showing strong general resemblance to ♂♂ of this genus. Distribution (Fig. 377): Honduras. Etymology: Masculine. A combination of “Apterus” (lat. = wingless) and the ending “-plopus” to indicate the close relation to Haplopus Burmeister, 1838. Species included: 1. Apteroplopus grosse-tuberculatus (Brunner v. Wattenwyl, 1907: 323) [Dyme].


HENNEMANN ET AL.

FIGURES 59–61. Apteroplopus grossetuberculatus (Brunner v. Wattenwyl, 1907) n. gen., n. comb.. 59. ♂ HT: Honduras [NHMW, No. 647]; 60. ♂ HT, apex of abdomen (lateral view) [NHMW, No. 647]; 61. ♂ HT, apex of abdomen (dorsal view) [NHMW, No. 647].

Apteroplopus grossetuberculatus (Brunner v. Wattenwyl, 1907) n. gen., n. comb. (Figs. 59–61, 377) Dyme grosse-tuberculata Brunner v. Wattenwyl, 1907: 323. HT, ♂: Coll. Br. v. W.; Honduras, E. Wittkugel; det. Br. v. W. Dyme grossetuberculata; 18.563 [NHMW, No. 647]. Bacteria grossetuberculata, Brock, 1998a: 32. Otte & Brock, 2005: 64.

Description: ♂ (Fig. 53). The following description is based on the unique ♂ HT in NHMW, which is complete except for lacking the left hind claw and the terminal antennomeres of both antennae. Large (body length 115.4 mm) and very slender insect. General colouration plain mid to olive green, the abdomen slightly darker (brown in the HT due to preservation). Pronotum pale brown, the terminal three abdominal segments greyish brown. Antennae pale brown dorsally and black ventrally. Apices of all femora and tibiae dark brown. Tarsi reddish brown. Points of the mesonotal tubercles and leg armature bright green. Eyes dark reddish brown. Head: Subcylindrical, vertex very gently convex and unarmed except for two minute tubercles. Eyes large, circular, their length only about 1.3x in that of cheeks. Scapus 1.5x longer than wide, slightly dorsoventrally compressed and oval in cross-section. Pedicellus cylindrical and 2/3 the length of scapus. Third antennomere almost twice as long as pedicellus; IV very short, V 2x longer than VI. Following six antennomeres elongate and increasing in length (X longest), the remaining gradually decreasing in length towards tip of antennae. Thorax: Pronotum about 1.4x longer than wide and unarmed, transverse median depression distinct, slightly curved and almost reaching lateral margins of segment. Mesonotum parallel-sided and slightly broadened at the posterior. Dorsal surface armed with about 16 irregularly paired, blunt tubercles which decrease in size towards the posterior. Meso- and metapleurae with a longitudinal row of small, rounded granulose. Meso- and metasternum simple. Abdomen: Median segment about 1.7x longer than wide and gently constricted medially. Tergum VIII dorsally with three almost parallel, longitudinal carinae. IX about 2/3 the length of VIII, distinctly narrower, decidedly constricted medially and with a blunt longitudinal dorsal keel. Anal segment less than ¾ the length of IX, gently widened towards the posterior and with a wide roughly triangular median excavation; dorsally with a fine longitudinal median carina (Fig. 61). Legs: As for the genus. Comments: ♀♀ and eggs unknown. So far only known from the unique ♂ holotype (Fig. 55). REVISION OF HAPLOPODINI


41

Distribution (Fig. 377): Honduras [NHMW]. Number of specimens examined: 1 TABLE 4. Measurements of Apteroplopus grossetuberculatus (Brunner v. Wattenwyl, 1907) n. gen., n. comb. [mm]. ♂ HT [NHMW] Body

115.4

Pronotum

4.2

Mesonotum

30.2

Metanotum

15.7

Median segment

4.0

Profemora

24.1

Mesofemora

20.0

Metafemora

24.0

Protibiae

25.3

Mesotibiae

18.0

Metatibiae

22.9

Antennae

> 57.0

5.3. Genus Cephaloplopus n. gen. Type-species: Cephaloplopus pulchellus n. sp., by present designation.

Description: ♂♂, ♀♀. Small to medium-sized (body length including subgenital plate ♀♀ 84.5–88.9 mm, ♂♂ 58.2–72.4 mm) and moderately elongate Haplopodini. ♀♀ brachypterous, ♂♂ brachypterous or with fully developed alae. Body ± cylindrical (♂♂) or sub-cylindrical (♀♀) in cross-section. Colouration of ♀♀ various shades of brown, straw and grey. ♂♂ colourful and multi-colourous insects, mostly pale to dull green with body surface glabrous. Head indistinctly longer than wide and globose, vertex roundly convex and distinctly bi-cornute. Horns extending by > 2/3 the height of head capsule, sometimes with a single point only (certain ♂♂) but usually crenate or multi-tuberculose; the dextral horn larger than the sinistral. Antennae filiform in ♀♀ and moderately thickened and robust in ♂♂; in ♀♀ almost as long as head and complete thorax combined, in ♂♂ reaching at least to abdominal segment III. Pronotum slightly shorter than the head and with a ± distinct pair of anterior tubercles or spines (rarely with an additional posterior pair); otherwise smooth (♂♂) or tuberculose (♀♀). Mesothorax elongate and ± parallel-sided, at least 2x longer than head and pronotum combined. Mesonotum in ♀♀ rather irregularly and to a various degree armed with granules and spiniform tubercles, in ♂♂ armed with 6–10 ± distinct and pointed spines. Mesopleurae smooth in ♂♂ and with a longitudinal row of acute granules in ♀♀. Meso- and metasternum irregularly and sparsely granulose in ♀♀, the metasternum smooth in ♂♂. Tegmina broadly oval and with a moderately distinct central hump; ± reaching to (♀♀) or projecting over posterior margin of metanotum (♂♂). Alae of ♀♀ ± as long as tegmina; of ♂♂ variable, either indistinctly longer than tegmina or as long as reaching to abdominal segment VI. Anal region reticulate with distinct brown to black radial and transverse veins in ♀♀, translucent pink in ♂♂ and with only the marginal portion reticulate. Abdomen considerably longer than head and thorax combined. Median segment longer than metanotum, smooth. Segments II–VII longer than wide (more distinct in ♂♂). Tergites unarmed in ♂♂, II–IV often with a ± prominent pair of spines or crenate posterior lobes in ♀♀. VII with lateral margins widely expanded and forming a prominent lobe which laterally extends by > ½ the width of body; posterior margin irregularly dentate in ♀♀. VIII–X slightly narrower (♀♀) or roughly equal in width (♂♂) to previous segments. Sternites II–VII smooth in ♂♂ and with a few small granules in ♀♀. Praeopercular organ of ♀♀ formed by a distinct elongate, wart-like median tubercle close to posterior margin of sternum VII. Anal segment with a longitudinal median carina, the posterior margin tapered and with a slight


HENNEMANN ET AL.

median notch. Epiproct very small, roundly triangular and hardly projecting over posterior margin of anal segment. Vomer of ♂♂ well developed, the basal portion roundly broadened and the terminal hook of moderate length. Cerci tapered towards the apex and laterally compressed, very small in ♀♀ and about 2/3 the length of anal segment in ♂♂. Poculum of ♂♂ convex, cup-like and longitudinally carinate. Subgenital plate of ♀♀ rather moderately elongate, projecting over the apex of abdomen by more than the length of tergum X (anal segment) and with the apex rounded. Legs of moderate length, profemora shorter (♀♀) or slightly longer than mesothorax (♂♂), mesofemora shorter (♀♀) or about as long as mesothorax, and hind legs not (♀♀) or ± reaching to apex of abdomen (♂♂). Two outer ventral carinae of meso- and metafemora each with 1–3 sub-apical spines or teeth, the medioventral carina of these femora with a longitudinal row of 4–6 ± distinct spines. The outer ventral carinae of ♀♀ sometimes elevated into rounded lobe (occasionally dentate). Dorsal carinae of all femora unarmed, but sometimes with a ± distinct, roughly triangular sub-apical tooth on the posterior carina in (♂♂). ♀♀ with a ± prominent sub-apical lobe on posterodorsal carina, the meso- and metafemora each with a ± prominent sub-apical and sub-basal lobe (the sub-basal lobes may be very shallow). Tibiae unarmed in ♂♂ and with 1–4 rounded dorsal lobes in ♀ (the basal lobe always present and largest). Coxae spinose. All basitarsi elongate and longer than following three tarsomeres combined; probasitarsus of ♀♀ with a rounded lobe dorsally. Eggs: Small (capsule length 3.1–3.4 mm), barrel-shaped and cylindrical in cross-section; capsule about 2.0x longer than wide. Dorsal surface of capsule considerably more convex than ventral surface. Polar-area ± impressed. Capsule surface strongly coriaceous and covered with irregular raised tubercles and ridges. Micropylar plate variable in shape, basal portion broadened and < ½ the length of capsule; sculptured like capsule. Posterior end of micropylar plate with a distinct triangular gap. Median line short but distinct. Operculum flat, circular and in the centre structured like capsule; no conspicuous central capitulum. Colouration brown. Differentiation (Table 5): Closely related to Haplopus Burmeister, 1838 but readily distinguished by: the considerably smaller size (body lengths: ♀♀ < 100 mm, ♂♂ < 70 mm); very prominent horns of the head which extend by > 2/3 the height of the head capsule; shorter mesothorax; large lateral lobes of abdominal tergum VII of both sexes; more prominently lobate legs and lobed probasitarsus of ♀♀; smaller, laterally compressed and tapered cerci of ♂♂. The eggs are remarkably different by having the capsule surface decidedly tuberculose, the capsule indented at the polar-area and lacking a knob-like capitulum. For a more detailed comparison and differentiation see Table 5 below. Distribution (Fig. 377): Hispaniola & Cuba. Etymology: Masculine. The generic name is a combination of “Cephalus” (lat. = head), which draws emphasise to the prominent head armature, and the ending “-plopus” to indicate the close relation to Haplopus Burmeister, 1838. Species included: 1. Cephaloplopus alope n. sp. [Distribution: Cuba] 2. Cephaloplopus euchlorus n. sp. [Distribution: Hispaniola] 3. Cephaloplopus laetus n. sp.[Distribution: Hispaniola] 4. Cephaloplopus pulchellus n. sp.[Distribution: Hispaniola]

TABLE 5. Comparison and differentiation of Cephaloplopus n. gen. and Haplopus Burmeister, 1838. Cephaloplopus n. gen.

Haplopus


< 70 mm

> 85mm

Body length incl. subgen. pl. (♀♀)

< 100 mm

> 110 mm

Slender and filiform

Robust

Very prominent; spine-like, conical, crenate or multi-tuberculate and extending by >2/3 the height of head capsule

Small to moderately sized, with a single point; extending by < 2/3 the height of head capsule

About 2x longer than head and pronotum combined

> 2.2x longer than head and pronotum combined

Antennae (♀♀) Spines of the vertex

Mesothorax (♀♀)




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TABLE 5. (Continued) Cephaloplopus n. gen.

Haplopus

Abdominal tergum VII

With very large posterolateral lobes, which extend by more than ½ the body width

At best with small posterolateral lobes

Cerci (♂♂)

Laterally compressed and tapered towards apex; at best 2/3 the length of anal segment

Obtuse and ± cylindrical; about as long as anal segment

With a rounded dorsal lobe

Slender

With a ± prominent sub-apical lobe on posterodorsal carina

Unarmed dorsally

With a ± prominent sub-basal and sub-apical lobe on dorsal carinae (basal ones may be indistinct)

At best with a slight sub-apical tooth on posterodorsal carina

With 1–4 rounded dorsal lobes

Not distinctly lobed

Barrel-shaped

Ovoid

Distinctly coriaceous / tuberculose and covered with irregular ridges

Minutely granulose / rugulose to almost smooth

Impressed

Convex and rounded

At best ½ the length of capsule

> ½ the length of capsule

Flat and with an irregularly raised circle of tubercles

± convex; with a conical or knob-like central capitulum

Hispaniola & Cuba (Fig. 377)

Hispaniola, Jamaica, Puerto Rico, Virgin Islands, Bahamas, Florida Keys, Cayman Islands 6 Swan Islands (Fig. 376)

Probasitarsus (♀♀) Profemora (♀♀) Meso- and Metafemora (♀♀)

Tibiae (♀♀) Egg (capsule) Egg (capsule surface) Egg (polar area) Egg (micropylar plate) Egg (operculum) Distribution

Note: The distinguishing features define to adult insects only. Nymphs of Haplopus spp. may have the body and leg armature considerably stronger developed than the corresponding adults, thus resembling Cephaloplopus n. gen.

Keys to the species of Cephaloplopus n. gen. ♀♀* 1. 2. -

Subgenital plate long and lanceolate, extending over abdomen by at least combined length of tergites IX–X; horns of the head flattened apically; Hispaniola . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 Subgenital plate extending over abdomen by slightly more than length of anal segment (Figs. 64–65); horns of head distinctly conical (Fig. 63); Cuba . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . alope n. sp. Pronotum with two pairs of spines (Fig. 88); meso- and metafemora with prominent dorsal lobes sub-basally; tibiae with one rounded pre-medial lobe dorsally . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .pulchellus n. sp. Pronotum with one anterior pair of tubercles (Fig. 69); meso- and metafemora lacking dorsal sub-basal lobes; tibiae with at least two lobes dorsally . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . euchlorus n. sp.

* ♀♀ of C. laetus n. sp. are not known

♂♂* 1. 2. -

Alae well developed, exceeding abdominal tergum V . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 Brachypterous, alae reaching only half way along median segment (Figs. 80, 82) . . . . . . . . . . . . . . . . . . . . . . . . . . laetus n. sp. Pronotum with two pairs of tubercles/spines (Fig. 89); general colouration dull green, head unicolorous (Fig. 89); costal region of alae with a longitudinal row of 6–8 distinct, elliptical pale spots (Fig. 86) . . . . . . . . . . . . . . . . . . . . . . . . . . .pulchellus n. sp. Pronotum with one anterior pair of tubercles (Fig. 70); general colouration pale green to cream; head pale green anteriorly, white posterolaterally and with a pale brown, longitudinal median stripe (Fig. 70); costal region of alae without a row of pale spots (Figs. 67–68) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . euchlorus n. sp.

* ♂♂ of C. alope n. sp. are not known


HENNEMANN ET AL.

FIGURES 62–65. Cephaloplopus alope n. gen., n. sp.. 62. ♀ HT: Cuba Camagüey [ANSP]; 63. ♀ HT head in lateral view: Cuba Camagüey [ANSP]; 64. ♀ HT, apex of abdomen (dorsal view); 65. ♀ HT, apex of abdomen (lateral view).

Cephaloplopus alope n. gen., n. sp. (Figs. 62–65, 335) Diapherodes alope Moxey, 1972: 92 (in litt.). [Nomen nudum] HT, ♀: Camagüey, Cuba, July 30. 1923, J. Acuña Col., E.E.A. Cuba; Ento No. 10660, w0179, Diapherodes alope Moxey Type (MS) [ANSP]. PT, ♀ (nymph): E.E.A. Cuba, Ento No. 10680, w0184, Diapherodes alope Moxey Paratype (MS) [ANSP]. PT, ♀ (nymph): E.E.A. Cuba, Ento No. 10681, w0182, Diapherodes alope Moxey Paratype (MS) [ANSP]. PT, ♀ (nymph): E.E.A. Cuba, Ento No. 10682, w0181, Diapherodes alope Moxey Paratype (MS) [ANSP]. PT, ♀ (nymph): Camagüey, Cuba, Col. J. Acuña, Julie 21. 1923, E.E.A. Cuba, Ento No. 10660, w0183, Diapherodes alope Moxey Paratype (MS) [ANSP]. PT, ♀ (nymph): Cangrejeras, Prov. Habana, May 30, ´31, S.C. Bruner, w0180, Diapherodes alope Moxey Paratype (MS) [ANSP].

Diagnosis: This, the only Cuban species of the genus, is distinguished from the three other known members by the distinctly conical horns of the head, which are almost circular in cross-section (Fig. 63), and much shorter subgenital plate, which projects over the apex of the abdomen by only a little more than the length of the anal segment (Figs. 64–65). Furthermore, the lobes of the legs are comparatively less developed. Etymology: “Alope” was the beautiful daughter of King Cercyon, and is a noun in apposition to the generic name Haplopus. It is the manuscript name supposed for this species by Moxey (1972: 92, in litt.) and here selected. Description: ♀ (Fig. 62). Of moderate size (body length incl. subgenital plate 85.0–88.9 mm) and rather slender for the genus, with a pair of very prominent conical cephalad horns, a greatly triangularly dilated abdominal tergum VII and comparatively short subgenital plate. General colouration of body and legs reddish to ochraceous mid brown with a greyish wash and all over furnished with pale grey to whitish speckles and markings,



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the legs with irregular dark greyish brown markings. Eyes dull red, antennae except scapus reddish brown. Tegmina and costal region of alae very dark brown and mottled with some pale green. Anal region of alae red with darker veins. Tarsi chestnut brown. Head (Fig. 63): Strongly globose, vertex convex and armed with an extremely prominent pair of large conical horns (the dextral one about 3x larger than the sinistral), which extend by as much as the height of head capsule; horns cylindrical basally carinate apically and tipped with a minute spine at the apex. Remainder parts of head with scattered, minute tubercles. Between the bases of the antennae with a slightly raised and smooth oval area. Eyes rather small, circular and conatined about 2.5x in length of cheeks. Antennae at least as long as complete thorax and median segment combined (broken in the HT). Scapus oval in cross-section, rectangular, 2x longer than wide. Pedicellus cylindrical and less than half the length of scapus. Thorax: Pronotum considerably shorter and narrower than head, longer than wide and slightly narrowing posteriorly. Transverse median sulcus indistinct and short, not reaching lateral margins of segment. Anterior half with a pair of strong spine-like tubercles (Fig. 63), posterior margin with four regularly-spaced tubercles, remaining parts of pronotum very minutely granulose. Mesothorax about 2x longer than head and pronotum combined. Mesonotum covered with numerous scattered small tubercles and four conspicuous tuberculate spines clustered close to the anterior margin (Fig. 63) as well as a few slightly larger tubercles medially. Meso- and metapleurae as well as meso- and metasternum with a few scattered small tubercles. Tegmina oval, scale-like and narrowed towards the base, just reaching to posterior margin of metanotum; central protuberance shallow. Alae shorter than tegmina and reaching less than half length along median segment. Abdomen: Median segment roughly 2x longer than wide and widened towards the posterior margin; unarmed. Segments II and III of equal length and width, parallel-sided. IV–VI slighly tapering and decreasing in length, VI shortest and narrowest, VII as long as II. II–VI on average about 2x longer than wide. Tergites II and III each with a pair of retrorse posteromedial spines and on each side with a prominent, foliaceous, triangular elevation (largest on II). IV with four minute retrorse spines at posterior margin, V and VI unarmed. VII with four irregular, raised longitudinal carinae and greatly dilated towards the posterior, forming a roughly triangular, foliaceous lateral expansion, which laterally projects by more than 2/3 of body width (Fig. 64). Sternites II–VII unarmed. Praeopercular organ formed by two gently curved and somewhat converging carinae at posterior margin of sternum VII (Fig. 335). Tergum VIII 2/3 the length of VII, distinctly narrower, strongly convex, parallel-sided and 1.3x longer than wide; anterior margin broader than posterior margin; IX 2/3 the length of VIII. Anal segment very slightly shorter than IX, tapered towards the apex, posterior margin with a shallow concave excavation (Fig. 64). Epiproct distinct, triangular, tectiform and projecting over posterior margin of anal segment (Fig. 64). Cerci short, oval in cross-section and tapered towards the apex, finely setose. Subgenital plate fairly short, slightly keeled with the apex rounded and projecting over apex of abdomen by slightly more than length of anal segment (Figs. 64–65). Legs: Profemora ¾ length of mesothorax, mesofemora slightly longer than metathorax and metafemora reaching about 1/3 the way along abdominal segment IV. Medioventral carina of profemora unarmed, of mesofemora with 2–3 and of metafemora armed with 3–5 spines. Antero- and posteroventral carinae of meso- and metafemora each with two small triangular sub-apical teeth. Dorsal carinae of meso- and metafemora with a minute and blunt submedial tooth and a more distinct tooth sub-apically (that of the anterior carina considerably larger). All tibiae unarmed except for a minute, rounded lobe about ¼ the way along anterodorsal carina. Basitarsi slightly longer following three tarsomeres combined, dorsal carina of probasitarsus just gently rounded. Nymphs: These are similar in appearance and colour to the adult insects but have the armature of the thorax, abdomen and legs much more prominently developed. Comments: Moxey (1972: 92) described Diapherodes alope in manuscript but never published this name. The suggested type-material and specific name are taken from Moxey. The ♀ from Alianza in ANSP and considered a PT by Moxey is not traced. Distribution: Cuba (Camagüey & Province Habana: Cangrejeras) [ANSP]. Number of specimens examined: 6


HENNEMANN ET AL.

TABLE 6. Measurements of Cephaloplopus alope n. gen., n. sp. [mm] Measurements [mm]

♀, HT [ANSP]

Body (incl. subgen. pl.)

88.9

Pronotum

3.9

Mesonotum

16.8

Metanotum

4.1

Median segment

6.3

Tegmina

5.2

Alae

4.0

Profemora

13.7

Mesofemora

11.7

Metafemora

15.0

Protibiae

12.6

Mesotibiae

11.2

Metatibiae

15.2

Antennae

> 29.0

Cephaloplopus euchlorus n. gen., n. sp. (Figs. 66–79, 336, 353, 379) HT, ♂: Dominican Republic, RD-063 Sierra Prieta, Santo Domingo Prov., 18°38.315’N 69°58.302’W, 20.XI.2002, D. Perez, R. Bastardo, B. Hierro [USNM]. PT, ♀: Dominican Republic, RD-063 Sierra Prieta, Santo Domingo Prov., 18°38.315’N 69°58.302’W, 20.XI.2002, D. Perez, R. Bastardo, B. Hierro [USNM]. PT, egg (ex abdomen): Dominican Republic, RD-063 Sierra Prieta, Santo Domingo Prov., 18°38.315’N 69°58.302’W, 20.XI.2002, D. Perez, R. Bastardo, B. Hierro [USNM]. PT, ♂: Dominican Republic, RD-041, Sierra Prieta, Villa Mella, Santo Domingo Prov., 18°38.939’N 69°58.373’W, 8.VII.2002, 500 ft., D. Perez, B. Hierro, R. Bastardo [USNM]. PT, ♂: Dominican Republic, RD-219 Sierra Prieta, Villa Mella, Santo Domingo Prov., 142 m, 18°38.925’N 69°58.303’W, 12.IV.2004, D. Perez, B. Hierro, R. Bastardo. (n) [USNM]. PT, ♀ (penultimate instar): Dominican Republic, RD-091, Sierra Prieta, Santo Domingo Prov., 133 m, 18°39.010’N 69°58.409’W, 18.III.2003, D. Perez, R. Bastardo, B. Hierro (night) [USNM]. PT, ♀ (nymph n3): Dominican Republic, RD-063, Sierra Prieta, Santo Domingo Prov., 18°38.315’N 69°58.302’W, 20.XI.2002, D. Perez, R. Bastardo, B. Hierro [USNM]. PT, ♀ (nymph n4): Dominican Republic, DR-062 ~ 5 km W Las Américas Airport, Santo Domingo, 18°28.158’N 69°43.601’W, 19.XI.2002, D. Perez, R. Bastardo, B. Hierro. (night) [USNM]. PT, ♀ (nymph n2): Dominican Republic, DR-060 Monte Rio, ~ 4 km W of beach, 70 m, dry scrub, 18°22.972’N 70°43.036’W, 17.XI.2002, D. Perez, R. Bastardo, B. Hierro. (night) [USNM].

Diagnosis: Females differ from those of the the Cuban C. alope n. sp. by: the longitudinal apical keel of the cephalad horns (Fig. 62); much longer subgenital plate, which projects over the abdomen by almost 1.5x the combined length of abdominal tergites IX–X (Fig. 68); dorsally multi-lobate protibiae and presence of a distinct dorsal sub-apical lobe on the meso- and metatibiae. Males are very similar to those of C. laetus n. sp. but at once differ by the well developed alae (Fig. 67, hardly longer than the tegmina in laetus), but also by the more slender and brownish instead of green legs, more distinct sub-apical tooth of the dorsal carinae of the meso- and metafemora and shape of the anal segment (Fig. 70). From C. pulchellus n. sp. it differs by the shorter horns of the head (Figs. 62–63), lacking a posterior pair of tubercles on the pronotum and smaller lateral lobes of abdominal tergum VII of both sexes. ♀♀ also differ by: the longer alae; smaller crest-like lobes of abdominal tergum II (Fig. 65); lack of the lateral sub-basal lobes of the



47

subgenital plate; lack of sub-basal dorsal lobes on the meso- and metafemora and presence of a rounded, dorsal sub-apical lobe on the meso- and metatibiae. Males furthermore differ by: the less angular anal segment (Fig. 70); pale green general colouration; pale brown legs and lack of distinct pale spots along the anterior margin of the costal region of the alae. The eggs can be distinguished from those of C. pulchellus n. sp. by: the less distinctly sculptured capsule surface and narrow anterior extension of the micropylar plate (Figs. 71–72).

FIGURES 66–68. Cephaloplopus euchlorus n. gen., n. sp.. 66 ♀ PT: Dominican Republic, Santo Domingo Province, Sierra Prieta [USNM]; 67. ♂ HT: Dominican Republic, Santo Domingo Province, Sierra Prieta [USNM]; 68. ♂ PT: Dominican Republic, Santo Domingo Province, Sierra Prieta [USNM].

Etymology: The specific name “euchlorus” (lat. = beautifully coloured) refers to the pretty colouration of the ♂♂, which consists of bright apple green, pale and dark brown, yellow and white. Description: ♀ (Fig. 66): Of moderate size for the genus (body length including subgenital plate 87.5 mm) and rather slender, with a prominent pair of crenulate horns on the head, a moderately dilated abdominal tergum VII and comparatively long subgenital plate. General colouration of body and legs mid brown with a greyish wash and all over furnished with irregular whitish speckles and markings. Head with cheeks mostly white and a small, elongate black spot above eyes, a conspicuous dark brown marking near posterior margin. Eyes dark reddish brown, antennae greyish brown with indistinct blackish annulations. Tegmina and costal region of alae very dark


HENNEMANN ET AL.

brown with a green wash in central portion; the main longitudinal veins black. Anal region of alae bright red with all longitudinal veins broadly marked with black. Head: Strongly globose, vertex convex and armed with a pair of prominent, apically compressed and crenulate cephalad horns (the dextral one almost 2x larger than the sinistral); both lateral surfaces of the horns with several spiniform tubercles (Fig. 69). Posterior of vertex with a pair of moderate tubercles and a pair of minute granules on the frons. Eyes rather small, circular and contained a little more than 2x in length of cheeks. Antennae reaching to posterior margin of mesonotum and consisting of about 45 segments. Scapus compressed dorsoventrally, roundly rectangular and about 2x longer than wide. Pedicellus cylindrical and roughly half the length of scapus. Thorax: Pronotum about as long but narrower than head, 1.2x longer than wide and gently constricted medially with the anterior margin widened and posterior margin rounded. Transverse median sulcus distinct and almost straight. In anterior portion with a pair of short spines (Fig. 69) and the surface otherwise with irregularly disposed small tubercles. Mesothorax about 2.2x longer than head and pronotum combined. Mesonotum with a fine longitudinal median carina and irregularly tuberculate; a cluster of four short but strong spines is present at anterior margin as well as two pairs of low spines in the median portion of the dorsal surface. The mesopleurae with a longitudinal row of about ten, the metapleurae with five small tubercles. Meso- and metasternum sparsely tuberculose. Tegmina oval, strongly broadened medially, narrowed towards the base and very slightly projecting over posterior margin of metanotum; central protuberance rather distinct, rounded. Alae reaching about 3/5 the way along median segment. Abdomen: Median segment roughly 2x longer than wide and widened towards the posterior; unarmed. Segments II–V roughly of equal length, VI slightly shorter than previous, III–VI gently narrowing; IV 3.3x longer than wide. Tergum VII slightly shorter than VI and with the lateral margin in posterior half dilated into a prominent rounded lobe, which laterally extends by about 2/3 the width of segment (broken in the unique adult ♀ available but described from a large ♀ nymph). All tergites with a few scattered granules and a longitudinal row of about five granules along lateral margins, II and III a pair of irregular, crenulate carinae posteromedial, which are less distinct on III (Fig. 72). These merely represented as a pair of tubercles on IV–IX. Sternites II–VI with a pair of short carinae near posterior margin and praeopercular organ represented by an elongate wart-like structure on VII (Fig. 336). Tergites VIII–X distinctly narrower than previous and roughly of uniform width; VIII ¾ the length of VII, almost 2x longer than wide and gently constricted medially, IX 2/3 the length of VIII. Anal segment shorter than IX, longitudinally carinate dorsally and with a concave lateral emargination; posterior portion strongly tapered towards a narrow and medially notched apex. Epiproct very small and with a posteromedian notch. Cerci short, conical and with a rather acute apex. Subgenital plate elongate, naviculate, keeled longitudinally and projecting over abdomen by more than the combined length of tergites IX–X (Fig. 75); apex rounded. Legs: Profemora about ¾ length of mesothorax, mesofemora about as long as metathorax and metafemora slightly projecting over posterior margin of abdominal segment III. Medioventral carina of profemora with three spines, of meso- and metafemora with 4–5 rather strong spines. Posteroventral carina of meso- and metafemora with one, the anteroventral carina with two sub-apical spines; both carinae gently expanded and minutely dentate in basal portion. The dorsal carinae each with a roundly triangular lobe sub-apically. Anterodorsal carina of tibiae with a distinct rounded sub-basal and sub-apical lobe. The anterodorsal carina of profemora strongly raised, slightly undulate and with a small rounded lobe sub-apically; the same carina in protibiae furnished with 3–4 rounded lobes. Probasitarsus with a distinct rounded dorsal lobe and a little longer than following two tarsomeres combined (Fig. 71). Meso- and metabasitarsi slender and as long as following three tarsomeres combined. ♂ (Figs. 67–68): Of moderate size (body length 63.8–72.0 mm) and rather slender for the genus with long alae (length 32.3–34.0 mm) which reach to abdominal segment VI; body surface glabrous. General colour of body pale creamish brown, the ventral body surface and three terminal abdominal segment pale apple green. Head creamish mid brown with the lower portions of cheeks white and the frons pale apple green. Antennae ochre with a greyish wash and with some grey annulations in apical portion. Eyes dark greyish brown with sepia mottling. Pronotum with the lateral margins broadly green. Lower portion of metapleurae and lateral margins of abdominal sternites II–V white. Spines of the mesothorax ochre with dark tips, lateral lobes of abdominal tergum VII dark brown, cerci creamish mid brown. Tegmina dark brown with the median portion of the anterior margin first pale green, then with a fine white stripe, the innermost portion pale apple green (Fig. 73). Costal region of alae with a similar green and interiorly white marking in basal portion, the anterior margin otherwise dull brown with slight pale mottling, then green; anal region transparent pale pink with a greyish wash along outer margin (Fig. 74). Legs and tarsi cream to mid brown with the apex of all femora slightly darker. REVISION OF HAPLOPODINI


49

FIGURES 69–79. Cephaloplopus euchlorus n. gen., n. sp.. 69. Head of ♀ PT [USNM]; 70. Head and pronotum of ♂ PT [USNM]; 71. Protarsus of ♀ PT [USNM]; 72. Abdominal tergum II of ♀ PT showing crenulate lobes [USNM]; 73 Metathorax and tegmina of ♂ PT (lateral view) [USNM]; 74. Right tegmen and ala of ♂ PT [USNM]; 75. ♀ PT, apex of abdomen (lateral view) [USNM]; 76. ♂ PT, apex of abdomen (lateral view) [USNM]; 77. ♂ PT, apex of abdomen (lateral view) [USNM]; 78. Egg PT (dorsal view) [USNM]; 79. Egg PT (lateral view) [USNM].


HENNEMANN ET AL.

Head: General shape as in ♀♀ but with the cephalad horns even longer but not crenulate, apex compressed, carinate and with a single pointed spine; dextral horn projecting by > 2/3 the height of head capsule (Figs. 70). Surface otherwise smooth except for a posterior pair of tubercles. Eyes prominent, circular and projecting hemispherically, no more than 1.5x length of cheeks. Antennae ± reaching posterior margin of abdominal segment II; consisting of about 50 antennomeres. Thorax: Pronotum about as long but narrower than head, general shape as in ♀♀. Surface smooth except for a pair of short spines in the anterior portion (Fig. 70). Mesothorax about 2.3x longer than head and pronotum combined. Armature of mesonotum variable; usually with four distinct, pointed spines near anterior margin (Fig. 70) and 2–3 pairs of pointed spines in the median portion. Meso- and metapleurae unarmed. Mesosternum with a few irregularly disposed granules, metasternum smooth. Tegmina oval and reaching about 1/3 along median segment, central protuberance distinct and roundly conical (Fig. 73). Alae reaching about 1/3 along abdominal segment VI. Abdomen: Segments II–V of equal length and about 3.2x longer than wide; VI–VII slightly decreasing in length with VII no more than 2.3x longer than wide. All tergites and sternites smooth, sternites IV–VI with two low carinae near posterior margin. VII with lateral margins in posterior half expanded into a ± distinct roundly triangular and retrorse lobe (Figs. 76–77); this either indistinct or extending laterally by as much as 2/3 of body width. VIII slightly shorter than VII and gently constricted in basal portion; IX shorter than VIII and constricted medially. Anal segment with a faint longitudinal median carina and narrowed posteriorly, the posterior margin rounded on both sides of the triangular median emargination (Fig. 77), slightly swollen and on ventral surface armed with several small, black in-curving denticles. Vomer with base broadly semi-circular and the terminal hook rather long, papillate and gently up-curving (Fig. 353). Cerci elongate, a little longer than anal segment, slightly laterally compressed and gently incurving. Poculum moderately convex, cup-like, longitudinally carinate and with a blunt basal protuberance (Fig. 76). Legs: Profemora a little longer and mesofemora slightly shorter than mesothorax, metafemora reaching about half way along abdominal segment IV. Medioventral carina of profemora with 1–2 small sub-apical spines, of mesofemora with four and metafemora with five prominent, pointed spines. Dorsal carinae of meso- and metafemora with a ± distinct triangular sub-apical tooth or lobe (may be very faint). Basitarsi about as long as following three tarsomeres combined. Nymphs: Even young nymphs show some of the characteristic features of this species, e.g. the crenulate cephalad horns or dorsal lobes of the tibiae. As in other species of the tribe, these have the whole body and leg armature much more prominently developed. The colouration includes various shades of brown furnished with a variable degree of whitish markings and speckles. Variation: In ♂♂ some variation is seen in the size of the cephalad horns, number and size of the mesonotal spines, size of the lateral lobe of abdominal tergum VII and size of the sub-apical tooth or lobe on the dorsal carinae of the meso- and metafemora. Egg (Figs. 78–79): Capsule 2x longer than wide, surface sparsely covered with irregular tuberculate structures, sub-glabrous. Polar-area with a distinct indentation. Micropylar plate slightly less than half the length of capsule, almost 2x longer than wide with the anterior 1/3 strongly narrowed and the posterior portion rounded. Centre of micropylar plate with arounded, wart-like protuberance. Colour plain reddish mid brown. Measurements [mm]: Length 3.4, width 1.7, height 1.9, length of micropylar plate 1.6. Distribution (Fig. 379): Hispaniola, S-Dominican Republic (Santo Domingo Province: Sierra Prieta & Azua Province: Monte Rio) [USNM]. Number of specimens examined: 8 TABLE 7. Measurements of Cephaloplopus euchlorus n. gen., n. sp. [mm] ♂, HT [USNM]

♂♂, PT [USNM]

♀, PT [USNM]

-

-

87.5

Body

72.0

63.8–65.0

78.5

Pronotum

2.9

2.4–2.8

3.8





51

TABLE 7. (Continued) ♂, HT [USNM]

♂♂, PT [USNM]

♀, PT [USNM]

Mesonotum

13.5

11.9–12.4

17.0

Metanotum

-

4.7

5.0

Median segment

-

6.0

5.3

Tegmina

7.7

7.8–8.2

6.0

Alae

34.0

32.3–33.0

5.0

Profemora

14.9

12.6–12.7

12.5

Mesofemora

12.3

11.8

10.9

Metafemora

16.2

14.2–14.5

13.2

Protibiae

14.8

12.4–13.0

11.0

Mesotibiae

10.2

9.9–10.2

9.6

Metatibiae

14.4

12.6–12.8

12.3

Antennae

ca. 35.0

> 25.0

25.5

Cephaloplopus laetus n. gen., n. sp. (Figs. 80–84, 379) HT, ♂: Dominican Republic, RD-153, La Poza de Agua Nueva, El Curro, Sierra Martín García, Azua Prov., 18°18.324’N 70°57.176’W, ~800 m, 15–16.VII.2003, D. Perez, R. Bastardo, B. Hierro. (day/night) [USNM]. PT, ♂: Dominican Republic, Sierra Martín García, Azua Prov., El Curro, near caseta parques, 18°22.372'N 71°01.724'W, 1,350 m, 22–23/viii/2014, D. Perez, C. Marte, K. Rodríguez, A. Sánchez [IIBZ].

Diagnosis: This new species is very close to C. euchlorus n. sp. but at once distinguished from this and the other known species in the genus by the shortened alae, which are hardly longer than the tegmina (Fig. 82). From C. euchlorus n. sp. ♂♂ (the only sex known) furthermore differ by the more robust and green legs which have only the knees brown, less distinct sub-apical tooth of the dorsal carinae of the meso- and metafemora and shape of the anal segment (Fig. 84). Etymology: The specific name “laetus” (lat. = pleasant) refers to the nice and bright colouration of ♂♂. Description: ♂ (Fig. 73): The colouration is described from a photo of the live holotype taken at the typelocality by the third author. Of moderate size (body length 60.8–67.0 mm) and rather robust for the genus with conspicuously shortened alae (length 4.8 mm) which are hardly longer than the tegmina; body surface glabrous. General colour pale apple green, the mesonotum, mesopleurae and lateral surfaces of the abdomen with a cream brown wash. Head creamish mid brown with the lower portions of cheeks white and the frons pale apple green (Fig. 81). Antennae ochre and becoming dark brown in the apical portion. Eyes very dark reddish brown. Pronotum brown with the lateral margins broadly green. Lower portion of metapleurae white. Spines of the mesothorax ochre, lateral lobes of abdominal tergum VII dark brown, cerci creamish mid brown. Tegmina with anterior margin bright yellow, then with a longitudinal brown streak, which is narrow in basal half and rather wide but weakly defined in posterior half, the innermost portion pale apple green (Fig. 82). Costal region of alae similar in colouration to tegmina but the longitudinal median stripe less distinct; anal region plain red. Legs green with the apical ¼ of all femora and tibiae mid brown; front legs with a brownish wash. Basitarsi green, the remaining tarsomeres chestnut brown. Head: Globose, vertex rather convex and armed with a pair of prominent, pointed cephalad horns; the larger dextral one almost projecting by height of head capsule. Apex of the horns strongly narrowed, pointed and slightly directed towards the anterior (Fig. 81). Two pairs of small granules are present in the posterior portion. Eyes almost circular, projecting hemispherically and their length contained less than 2x in that of cheeks. Antennae moderately robust, very long and reaching to posterior of abdominal segment III; consisting of 54 segments.


HENNEMANN ET AL.

FIGURES 80–84. Cephaloplopus laetus n. gen., n. sp.. 80. ♂ HT, Dominican Republic, Azua Province, La Poza de Agua Nova [USNM]; 81. Head and pronotum of ♂ HT in alteral view [USNM]; 82. Tegmina and alae of ♂ HT [USNM]; 83. ♂ HT, apex of abdomen (dorsal view) [USNM]; 84. ♂ HT, apex of abdomen (lateral view) [USNM].

Thorax: Pronotum about equal in length but narrower than head, very slightly narrowed towards the posterior and with the lateral margins gently rounded; surface smooth except for a pair of blunt tubercles in centre of anterior portion (Fig. 81). Transverse median sulcus moderately distinct. Mesothorax about 3x longer than head and pronotum combined. Mesonotum armed with four rather strong but short spines at anterior margin (Fig. 81), a prominent pre-medial pair of spines and a further pair of much smaller spines some 3/5 the way along dorsal surface. A row of very indistinct granules along lateral margins. Mesopleurae with a longitudinal row of minute granules, metapleurae unarmed. Mesosternum with very fine transverse sculpturing and sparsely set with a few small granules; metasternum smooth. Tegmina oval and just not reaching posterior margin of metanotum, central protuberance rather longitudinal and conspicuously displaced towards the posterior. Alae hardly longer than tegmina reaching about half way along median segment (Fig. 82). Abdomen: Segments II–V roughly of equal length and about 2.8x longer than wide; very gently constricted medially with surface smooth. VI only about ¾ the length of previous. VII even shorter and with the lateral margins in posterior half expanded into a large, roundly trinagular lobe; this laterally projects by almost half the width of



53

segment. All sternites smooth. VIII and IX just slightly shorter than VII, IX constricted medially. Anal segment shorter than IX, constricted in posterior portion and with a deep but narrow posteromedian incision (Fig. 84); on ventral surface armed with several small, black in-curving denticles. Vomer broadly triangular with the lateral margins strongly swollen; terminal hook rather short and slightly up-curving. Cerci elongate, about as long as anal segment, laterally compressed and gently in-curving. Poculum moderately convex, cymbiform, with an acute basal protuberance and carinate longitudinally (Fig. 83); posterior margin angulate. Legs: Profemora about equal in length mesothorax, the mesofemora slightly shorter, metafemora projecting a little over posterior margin of abdominal segment IV. Medioventral carina of profemora with two small spines in apical portion, of meso- and metafemora armed with four rather strong spines. Anteroventral carina of meso- and metafemora with two, the posteroventral carina with one sub-apical spine; both dorsal carinae very slightly expanded sub-apically. Basitarsi longer than following three tarsomeres combined. Comments: ♀♀ and eggs unknown. Distribution (Fig. 379): So far only known from the type-localites in the Sierra Martín García, Azua Province, SE-Dominican Republic: La Poza de Agua Nueva (800 m) [USNM] and El Curro (1350 m) [IIBZ] Number of specimens examined: 2 TABLE 8. Measurements of Cephaloplopus laetus n. gen., n. sp. [mm] ♂, HT [USNM]

♂, PT [IIBZ]

Body

67.0

60.8

Pronotum

2.9

2.7

Mesonotum

15.8

13.3

Metanotum

3.6

3.5

Median segment

5.1

4.9

Tegmina

4.1

4.1

Alae

4.8

3.7

Profemora

15.7

15.3

Mesofemora

12.3

13.0

Metafemora

15.4

16.1

Protibiae

16.1

15.3

Mesotibiae

12.8

12.3

Metatibiae

16.4

16.7

Antennae

ca. 40.0

39.2

Cephaloplopus pulchellus n. gen., n. sp. (Figs. 85–99, 337, 354, 379) HT, ♂: Dominican Republic, RD-034 Boca de la Cañada, ~ 15 km S Oviedo, Pedernales Prov., 4.VII.2002, 17°54.901’N 71°30.067’W, B. Hierro, R. Bastardo, D. Perez [USNM]. PT, ♀: Dominican Republic, RD-034 Boca de la Cañada, ~ 15 km S Oviedo, Pedernales Prov., 4.VII.2002, 17°54.901’N 71°30.067’W, B. Hierro, R. Bastardo, D. Perez [USNM]. PT, 3 eggs: Dominican Republic, RD-034 Boca de la Cañada, ~ 15 km S Oviedo, Pedernales Prov., 4.VII.2002, 17°54.901’N 71°30.067’W, B. Hierro, R. Bastardo, D. Perez [USNM]. PT, ♂: Dominican Republic, Parque Nacional Jaragua, Boca de la Cañada, 18 km S. Oviedo, 22.–23.IX.2000, D. Perez, R. Bastardo, B. Hierro [USNM]. PT, ♂ (nymph n4): Dominican Republic, RD-135, ~ 7 km Rd. to Caseta 1, Parque Nacional Sierra de Bahoruco, Independencia Prov., 18°17.711’N 71°34.335’W, 777 m, VII.2003, D. Perez, R. Bastardo, B. Hierro (day). [USNM]. PT, ♂ (nymph n5): Dominican Republic, RD-219, Sierra Prieta, Villa Mella, Santo Domingo Prov., 142 m, 18°38.925’N 69°58.303’W, 12.IV.2004, D. Perez, B. Hierro, R. Bastardo. (n) [USNM].


HENNEMANN ET AL.

Diagnosis: Females differ from those of all other known species by: the very prominent, apically carinate and crenulate horns of the head (Figs. 87–88); very large, retrorse crest-like lobes of abdominal tergum II (Fig. 91) and presence of such structures on IV; very long and slender, foliaceous lateral lobes of tergum VII, which posteriorly reach half way along tergum VIII (Figs. 93–94); presence of a sub-basal lateral lobe of the subgenital plate (Fig. 97); presence of only a single prominent sub-basal lobe on the posterodorsal carina of the profemora, and very prominent lobes of the mid and hind legs. The pretty ♂♂ are even more distinctive and at once differ from all other representatives of the genus by the dull green colouration and longitudinal row of 7–8 oval yellow markings along the anterior margin of the costal region of the alae (Fig. 86). They are furthermore well characterized by: the very long and slender cephalad horns (Fig. 89); presence of a distinct posterior pair of tubercles on the pronotum (Fig. 89); very large, slender and acutely triangular lateral lobes of abdominal tergum VII, which posteriorly reach half way along tergum VIII (Figs. 95–96) and presence of two sub-apical spines on the posteroventral carina of the meso- and metafemora. The eggs can be distinguished from those of C. euchlorus n. sp., the only other eggs known from the genus, by the more distinctly sculptured capsule surface and shorter, shield-shaped micropylar plate (Fig. 98). Etymology: The name “pulchellus” (lat. = beautiful) refers to the very pretty and distinctive colouration of ♂♂. Description: ♀ (Fig. 78). Rather small (body length including subgenital plate 84.5 mm) and moderately slender for the genus, with a pair of very large crenulate horns on the head, very large foliaceous lateral lobes on abdominal tergum VII and very characteristic body and leg armature. General colour of body and legs pale creamish grey, all over furnished with whitish markings and speckles as well as dark brown mottling along the extreme margins of all body and leg appendages; abdominal segments I–IV dark brown in the unique adult ♀ at hand and most certainly caused by preservation. Head with a pair of small diverging dark brown markings on frons and a large rather obscure dark brown marking on back of vertex; above the eyes with a short and narrow dark line directed antero-posteriad. Eyes creamish mid brown, antennae greyish brown with indistinct blackish annulations. Large tubercles or spines of the thorax with brown tips and the lateral granules dark olive. Abdominal sternites II–VII broadly dark brown posteriorly. Tegmina and costal region of alae very dark brown with the latter washed with white in basal portion; anal region of alae bright red with longitudinal veins black. Head: Globose, vertex strongly convex and armed with a pair of very large, apically compressed and crenulate cephalad horns, which project by more than height of head capsule (the dextral one slightly larger than the sinistral); both lateral surfaces of the horns with several spiniform tubercles (Figs. 87–88) and apex pointed medially. Posterior of vertex with several acute tubercles of variable sizes. Eyes rather small, circular and contained a little less than 2x in length of cheeks. Antennae reaching to posterior margin of metanotum and consisting of 35 segments. Scapus dorsoventrally compressed, roundly rectangular and about 2x longer than wide. Pedicellus cylindrical less than half the length of scapus. Thorax: Pronotum slightly shorter and considerably narrower than head, 1.2x longer than wide, roughly rectangular and with the posterior margin rounded. Transverse median sulcus distinct, almost straight but not reaching lateral margins of segment. Surface with two pairs of strong but rather short spines, one in anterior portion and one in posterior portion (Fig. 88); otherwise with a few small, scattered granules towards the lateral margins in particular. Mesothorax about 2x longer than head and pronotum combined. Mesonotum with a fine longitudinal median carina and with a pair of very strong but short anterior and pre-medial spines (the posterior pair larger, Fig. 85); surface otherwise irregularly tuberculate. Meso- and metapleurae each with a longitudinal row of rounded tubercles; meso- and metasternum irregularly tuberculose. Tegmina broadly ovate and projecting slightly over posterior margin of metanotum; central protuberance very prominent, roundly conical and conspicuously displaced towards the posterior margin (Fig. 83). Alae slightly shorter than tegmina and reaching no more than ¼ the way along median segment. Abdomen: Median segment roughly 2x longer than wide and gently widened towards the posterior; a few small granules in posterior portion. Segments II–IV roughly equal in length, V–VI slightly decreasing in length and shorter than previous; IV 3.2x longer than wide. Tergites II–VI covered with a few scattered granules. II and IV each with a posterior pair of very prominent and retrorse, crenulate foliaceous lobes; these considerably larger and with the outer margin dentate on II (Fig. 91). III, V and VI merely with two very low carinae posteromedially. Tergum VII slightly shorter than VI and with the lateral margin in posterior half protruded into a large and elongate,



55

retrorse foliaceous lobe which laterally projects by almost the bod y width and reaches as far back as the middle of tergum VIII; posterior margin irregularly dentate (Figs. 93–94). Sternites II–VI with a few scattered granules and a pair of short carinae near posterior margin; praeopercular organ represented by an elongate wart-like structure on VII (Fig. 337). Tergites VIII–X distinctly narrower than previous and roughly of uniform width; VIII ¾ the length of VII, 2x longer than wide and constricted medially, IX 3/5 the length of VIII. Anal segment shorter than IX, longitudinally carinate dorsally and with a blunt triangular tooth medio-laterally; posterior portion strongly tapered towards a narrow and medially notched apex (Fig. 93). Epiproct very small and roundly triangular. Cerci short, conical and with a rather acute apex. Subgenital plate elongate, naviculate, keeled longitudinally and projecting over abdomen by more than the combined length of tergites IX–X, apex rounded (Fig. 93). Lateral surfaces subbasally with a conspicuous rounded, lobe-like expansion (Fig. 97). Legs: Profemora and mesofemora shorter than mesothorax, metafemora reaching about 1/3 along abdominal segment IV. Medioventral carina of profemora with two very small apical spines, of mesofemora armed with 4–5 and of metafemora with six rather strong spines. Anterodorsal carina of profemora with a distinct, tri-dentate apical lobe and in protibiae with a roundly triangular lobe some 1/3 off the base. Posteroventral carina of meso- and metafemora with one, the anteroventral carina with two sub-apical spines; both carinae roundly expanded in basal portion. Both dorsal carinae with a triangular lobe pre-medially and sub-apically; very large and dentate on the posterior carina but much smaller and smooth on the anterior carina. Posterodorsal carina of meso- and metatibiae with a small, roundly triangular lobe-like tooth some 1/3 off the base. Probasitarsus with a distinct rounded dorsal lobe and a little longer than following three tarsomeres combined (Fig. 92). Meso- and metabasitarsi slender and longer than three tarsomeres combined. ♂ (Fig. 86). Rather small (body length 56.9–60.6 mm) and rather slender for the genus with long alae (length 28.6–29.5 mm) which reach to abdominal segment VI and a very characteristic colouration; body surface subglabrous. General colour of body dull green; ventral body surface, lateral surfaces of pro- and mesonotum and great parts of mesopleurae pale creamish brown with faint whitish mottling. Head green with a washed yellowish spot behind eyes, the lower anterior portion brown and the apices of the cephalad horns black (Fig. 89); eyes mid brown, antennae dark ochre and with a greenish wash in apical portion. Spines of the thorax with black tips. Mesopleurae with an elongate triangular green marking near posterior margin, metapleurae green. Abdominal tergites II–VI dark greyish laterally and with an obscure white posterolateral marking; lateral lobes of tergum VII dark brown. Tegmina dull green with anterior margin broadly blackish brown and in basal half interiorly bordered by an elongate, pale yellow marking. Costal region of alae dull green, the anterior margin with a broad blackish brown longitudinal stripe and furnished with a row of 7–8 oval pale yellow markings (Fig. 86). Anal region transparent pink with a greyish wash along outer margin. Legs dull green with the ventral surfaces and apex of all tibiae reddish mid brown; tarsi chestnut brown. Head: General shape as in ♀♀ but with the cephalad horns very long, slender and almost cylindrical near the base; apex compressed and with one or two pointed spines; dextral horn slightly larger and projecting by more than height of head capsule (Fig. 89). Surface otherwise smooth except for two pairs of tubercles on posterior of vertex. Eyes very prominent, circular and projecting hemispherically, contained no more than 1.3x in length of cheeks. Antennae ± reaching half way along abdominal segment IV; consisting of about 52 antennomeres. Thorax: Pronotum shorter and narrower than head and about 1.3x longer than wide; armature as in ♀♀ but the posterior pair of spines considerably larger than the anterior pair (Fig, 89). Mesothorax about 2.3x longer than head and pronotum combined. Mesonotum with a slight longitudinal median carina and armed with three pairs of short spines, the median pair largest; surface otherwise with very fine transverse sculpturing and some small granules along lateral margins. Mesopleurae granulose, metapleurae smooth. Meso- and metasternum very sparsely granulose, the mesosternum with very fine transverse sculpturing. Tegmina oval and reaching about 1/3 the along median segment, central protuberance distinct and acutely conical. Alae reaching about ¼ along abdominal segment VI. Abdomen: Segments II–V of uniform length and about 3x longer than wide; VI–VII slightly decreasing in length with VII no more than ¾ the length of II. All tergites and sternites smooth. Tergum VII with lateral margins in posterior half protruded into a very large, acutely triangular and retrorse lobe; this extending laterally by slightly more than body width (Figs. 95–96). VIII slightly shorter than VII and very gently widened in posterior portion; IX shorter than VIII and constricted towards the posterior. Anal segment with a faint longitudinal median carina,


HENNEMANN ET AL.

constricted at the base than distinctly expanded; the posterior margin broadly triangular with a small median indentation (Fig. 95) and on ventral surface armed with several small, black in-curving denticles. Vomer with base broadly triangular and terminal hook of moderate length and gently up-curving (Fig. 354). Cerci elongate, a little shorter than anal segment, slightly laterally compressed, tapered towards the apex and gently incurving. Poculum conically convex, cup-like, longitudinally carinate and with the posterior margin rounded (Figs. 96, 346). Legs: Profemora slightly longer, mesofemora roughly as long as mesothorax, metafemora almost reaching posterior margin of abdominal segment IV. Medioventral carina of profemora with 1–2 small sub-apical spines, of meso- and metafemora armed with four prominent, pointed spines. Both ventral carinae of meso- and metafemora with two distinct sub-apical spines, the anterodorsal carina with a triangular tooth sub-apically. Probasitarsus slightly longer, meso- and metabasitarsus about equal in length to remaining tarsomeres combined; all slender. Nymphs: Even rather young nymphs are easily recognized by some of the characteristic features of this species, e.g. the very long crenulate cephalad horns or single dorsal lobe of the tibiae. As in other species of the tribe, the appendages of the body and legs are even more prominently developed than in adults and ♂ nymphs also have a pair of crenulate lobes on abdominal tergum II which are missing in the alate adult insects. The colouration is similar to that of adult ♀♀. Egg (Figs. 98–99): Capsule 2x longer than wide, surface densely and unevenly tuberculate and irregularly pitted; a few enlarged tubercles roughly form some irregular ridge-like structures. A rather distinct longitudinal bulge running from the micropylar plate towards the polar-area. Polar-area merely with a very shallow indentation. Micropylar plate only about 1/3 the length of capsule, shield-shaped and about as long as wide. Colour yellowish mid brown, the outer margin of the micropylar plate dark brown. Measurements [mm]: Length 3.1, width 1.6, height 1.7, length of micropylar plate 1.0. Distribution (Fig. 379): Hispaniola, S-Dominican Republic (Pedernales Province: Parque Nacional Jaragua; Independencia Province: Parque Nacional Sierra de Bahoruco & Santo Domingo Province: Sierra Prieta) [USNM]. Number of specimens examined: 5 TABLE 9. Measurements of Cephaloplopus pulchellus n. gen., n. sp. [mm] ♂, HT [USNM]

♂, PT [USNM]

♀, PT [USNM]

-

-

84.5

Body

60.6

56.9

77.1

Pronotum

2.5

2.3

3.5

Mesonotum

12.0

10.2

15.6

Metanotum

-

-

5.0

Median segment

-

-

5.6

Tegmina

6.7

6.4

4.9

Alae

29.5

28.6

4.0

Profemora

14.0

13.0

13.9

Mesofemora

11.9

11.3

12.4

Metafemora

15.6

14.9

14.8

Protibiae

14.3

14.0

13.2

Mesotibiae

6.1

11.6

12.6

Metatibiae

15.9

16.0

16.2

Antennae

40.0

38.0

ca. 29.5




57

FIGURES 85–92. Cephaloplopus pulchellus n. gen., n. sp.. 85. ♀ PT: Dominican Republic, Pedernales Province, Boca de la Cañada [USNM]; 86. ♂ HT: Dominican Republic, Pedernales Province, Boca de la Cañada [USNM]; 87. Head of ♀ PT (caudal view) [USNM]; 88. Head and pronotum of ♀ PT (lateral view) [USNM]; 89. Head and pronotum of ♂ HT in lateral view [USNM]; 90. Tegmina and alae of ♀ PT [USNM]; 91. Abdominal tergum II of ♀ PT showing the large crenulate lobes [USNM]; 92. Protarsus of ♀ PT [USNM].


HENNEMANN ET AL.

FIGURES 93–99. Cephaloplopus pulchellus n. gen., n. sp.. 93. ♀ PT, apex of abdomen (dorsal view) [USNM]; 94. ♀ PT, apex of abdomen (lateral view) [USNM]; 95. ♂ HT, apex of abdomen (dorsal view) [USNM]; 96. ♂ HT, apex of abdomen (lateral view) [USNM]; 97. Basal portion of subgenital plate of ♀ PT showing the lobe-like appendages (lateral view) [USNM]; 98. Egg PT (dorsal view) [USNM]; 99. Egg PT (lateral view) [USNM].

5.4. Genus Diapherodes Gray, 1835 Type-species: Mantis gigantea Gmélin, 1789: 2055, by subsequent designation of Kirby, 1904a: 362. Diapherodes Gray, 1835: 33 (in part). Burmeister, 1838: 560, 573 (in part). Westwood, 1859: 84 (in part). Saussure, 1872: 185 (in part). Stål, 1875: 31. Bolivar, 1888: 140 (in part). Kirby, 1904a: 362 (in part). Redtenbacher, 1908: 434 (in part). Rehn, 1909: 200. Wolcott, 1922: 23. Wolcott, 1936: 35. Rehn & Hebard, 1938: 53, pl. 4: 22 (♀). Wolcott, 1948: 50.



59

Wolcott, 1951: 50. Moxey, 1972: 85 (in litt.; in part). Langlois & Lelong, 1997: 43, fig. 20 (♂, ♀). Brock, 1998c: 33. Langlois, Lelong, Rastel, Polidori & Dorel, 2000: 46, fig. 19-g (♂, ♀) & photos 10–12 (egg). Lelong, Langlois, Rastel & Dorel, 2003: 48, fig. 19a–g (♂, ♀) & photos 11–13 (egg). Lelong & Langlois, 2005: 264. Otte & Brock, 2005: 120 (in part). Langlois, Lelong & Rastel, 2006: 42, figs. 19 a–g (♂, ♀, nymph), photos 5–6 (♀) & 27–29 (egg). Aplopus, Redtenbacher, 1908: 429 (in part). Rehn, 1904: 68 (in part). Rehn, 1909: 200. Wetmore, 1916: 58. Wolcott, 1923: 23. Wolcott, 1936: 35. Wolcott, 1948: 50. Wolcott, 1951: 50. Otte & Brock, 2003: 301 (in part). Aplopus (?), Rehn & Hebard, 1938: 52. Cyphocrana, St. Fargeau & Audinet-Serville, 1825: 445. Audinet-Serville, 1831: 61. Haplopus, Westwood, 1859: 85 (in part). Kirby, 1904a: 363 (in part). Mantis, Drury, 1773: 88, 89, pl. 44: 1 & 50 (in part). Fabricius, 1775: 274. Fabricius, 1787: 227. Gmélin, 1789: 2054-2055. Olivier, 1792: 634. Fabricius, 1793: 13. Phasma, Fabricius, 1798: 187, 188. St. Fargeau & Audinet-Serville, 1825: 101. Westwood, 1859: 34, pl. 22: 7, 7a, 7b (♀). Phasma (Diapherodes), de Haan, 1842: 109. [Not: Diapherodes, Saussure, 1872: 183. à Venupherodes n. gen.]

Description: ♀♀, ♂♂. Medium-sized to very large (body lengths: ♀♀ 92.0–197.0 mm, ♂♂ 60.0–126.0 mm) and moderately slender to very robust Haplopodini. ♀♀ in particular, often broad to very broad and massive insects with the body distinctly oval in cross-section and the thorax and abdomen conspicuously broadened; brachypterous. ♂♂ moderately slender, body cylindrical or sub-cylindrical in cross-section, with scale-like tegmina and well developed alae. Body surface of ♀♀ ± shiny, colouration usually plain bright green (rarely brown or with brown markings), ♂♂ brown or green. Head globose, indistinctly longer than wide. Vertex roundly convex, tuberculose and armed with a pair of ± prominent tubercles or blunt spines. Antennae with all antennomeres moderately thickened and longer than head and thorax combined. Scapus dorsoventrally flattened, longer than pedicellus. Pronotum tuberculose, with or without a ± prominent pair of blunt spines or tubercles in anterior half. Mesothorax at best slightly more than 2x the length of head and pronotum combined, in ♂♂ slender and roughly parallel sided, in ♀♀ ± strongly widened towards the posterior and constricted anteriorly. Mesonotum anteriorly armed with 2–6 distinct spiniform tubercles or spines; surface otherwise smooth, sparsely granulose or tuberculose. Meso- and metapleurae of ♀♀ ± strongly expanded and laterally projecting over mesonotum; armed with a longitudinal marginal row of ± prominent tubercles, teeth or spines. Meso- and metasternum either smooth, both granulose, or only the mesosternum with a few scattered granules. ♀♀ with scale-like tegmina which at best slightly project over posterior margin of metanotum. Alae at best slightly longer than the tegmina, but usually much shorter and may be vestigial. Tegmina of ♂♂ oval, longer than metanotum and with a ± distinct conical central hump. Alae at least reaching to posterior margin of abdominal tergum V. Anal region of alae transparent pink or violet (♀♀), plain transparent white, pink or orange (♂♂). Abdomen distinctly longer than head and complete thorax combined; unarmed in ♂♂, tergites sometimes finely multi-carinate in ♀♀. Abdomen of ♀♀ ± broadened and swollen basally or sub-basally, with segments II and/or III the broadest, then gradually narrowing towards the apex. Abdomen of ♂♂ slender and cylindrical with the terminal three segments slightly broader than


HENNEMANN ET AL.

previous. In ♀♀ segments II–IV transverse to quadrate, V to VII slightly longer than wide; in ♂♂ II–VII parallelsided and 1.5–2.5x longer than wide. Tergum VII of ♀♀ with lateral margins ± rounded and/or expanded, or with a triangular posterolateral lobe; in ♂♂ parallel-sided or with a small lobe posterolaterally. Praeopercular organ of ♀♀ indistinct; formed by a single posteromedian granule or wart-like tubercle on sternum VII. Cerci very small, oval in cross-section and tapered towards the apex; slightly in-curving in ♂♂, often hidden underneath anal segment in ♀♀. Epiproct very small and mostly hidden underneath anal segment. Vomer short and broad; roughly triangular. Subgenital plate of ♀♀ keeled, narrowed towards a ± pointed apex and extending considerably beyond apex of abdomen. Poculum of ♂♂ strongly convex and cup-like. Legs of moderate length and rather broad. Profemora at best about as long as mesothorax, hind legs not reaching to apex of abdomen. Medioventral carina of femora armed with a longitudinal row of spines (often indistinct on profemora). Both dorsal carinae of meso- and metafemora occasionally with a sub-apical lobe, the ventral carinae each with 1–2 distinct and pointed sub-apical spines. Some carinae of mid and hind legs may be ± elevated and ledge-like in ♀♀ of certain species. Basitarsi short, at best slightly longer than following two tarsomeres combined. Eggs: Medium-sized to large (capsule length 3.7–5.6 mm), ovoid and cylindrical in cross-section; capsule 1.5–2.0x longer than wide. Polar-area ± impressed in lateral aspect. Capsule surface strongly coriaceous and covered with irregular raised tubercles and ridges to a variable degree. Micropylar plate generally drop or heartshaped, narrowed towards anterior end and broadened basally, at least 1/3 the length of capsule; central portion sculptured like capsule. Posterior end of micropylar plate with a wide triangular gap. Median line distinct. Operculum flat to slightly convex, circular and in the centre structured like capsule; no conspicuous central capitulum. Colouration irregularly grey or brown. Differentiation: Closely related to the Cuban Venupherodes n. gen. and Haplopus Burmeister, 1838. Slender representatives of the jamaicensis-group in particular resemble stockier representatives of Haplopus. However, Diapherodes is absent in Cuba and easily distinguished from Venupherodes n. gen. by: the larger size (♀♀ in particular); unicolourous head of ♂♂; elevated and prominently armed mesopleurae which laterally project over the mesonotum; mesothorax which is gradually broadened towards the posterior; lack of a longitudinal median keel of the mesonotum; presence of tegmina and alae; basally or sub-basally broadened abdomen; simple praeopercular organ and longer subgenital plate of ♀♀. The eggs at once differ by the less strongly sculptures capsule surface and longer, elongate micropylar plate (→ Table 33). From Haplopus it differs by: the broader and more robust body (♀♀ in particular); relatively shorter mesothorax (at best a little more than 2x the length of head and pronotum combined); shorter basitarsi and plain anal region of the alae of both sexes; oval body cross-section and more decidedly broadened meso- and metathorax of ♀♀, as well as the smaller and much more slender cerci ♂♂. Furthermore, ♀♀ are mostly plain bright green, whereas ♀♀ of Haplopus are brown or grey. The eggs clearly differ from those of Haplopus by having the capsule surface much more prominently sculptured and lacking a distinct hat- or knob-like central capitulum. Comments: The classification of Diapherodes has varied through the years, but in the way treated by Redtenbacher (1908: 434) the genus was paraphyletic and as interpreted by Moxey (1972: 85 ff., in litt.) included five generic units. Several features of the insects and eggs indicate two species-groups within the genus, here recognized as the gigantea-group and jamaicensis-group. These two groups are geographically rather well separated, members of the gigantea-group being restricted to the Lesser Antilles, and the three known species of the jamaicensis-group occuring only on the Greater Antillean islands of Jamaica and Puerto Rico (→ 6.). The systematic level of sub-genera was considered for these two groups of species in Diapherodes, but due to the lack of satisfying distinctive features in the eggs in particular, interpreting these solely as species-groups appears more appreciable. For a better understanding of the present arrangement a detailed comparison of these two species-groups here proposed appears warranted and is presented below (→ Table 10). Distribution (Fig. 375): Lesser Antilles (Guadeloupe, Dominica, Martinique, Saint Lucia, Grenada, & Saint Vincent → gigantea-group, Fig. 381) and Greater Antilles (Puerto Rico & Jamaica → jamaicensis-group). Species & subspecies included: I. Gigantea-group 1. Diapherodes angulata (Fabricius, 1793: 13) [Mantis]. rev. stat. = Haplopus grayi Kaup, 1871: 36. n. syn. [Distribution: Guadeloupe]



61

2. Diapherodes dominicae (Rehn & Hebard, 1938: 53). n. comb. [Distribution: Dominica] 3. Diapherodes gigantea gigantea (Gmélin, 1789: 2055) [Mantis]. = Mantis gigas Drury, 1773: 89, pl. 50 (♀). = Cyphocrana cornuta St. Fargeau & Audinet-Serville, 1825: 445. [Distribution: Saint Vincent & Grenada] 4. Diapherodes gigantea saintluciae n. ssp. [Distribution: Saint Lucia] 5. Diapherodes martinicensis Lelong & Langlois, 2005: 264. [Distribution: Martinique]

II. Jamaicensis-group 6. Diapherodes achalus (Rehn, 1904: 68) [Aplopus]. n. comb. = Diapherodes longiscapha Redtenbacher, 1908: 435. n. syn. [Distribution: Puerto Rico] 7. Diapherodes jamaicensis (Drury, 1773: 88, pl. 44: 1 (♂)) [Mantis]. = Mantis bispinosa Fabricius, 1775: 274. = Haplopus christopheri Westwood, 1859: 84, pl. 33: 4, 4a (♀). n. syn. = Diapherodes glabricollis Gray, 1835: 33. n. syn. = Haplopus murinus Redtenbacher, 1908: 429. n. syn. = Dipaherodes pulverulentus Gray, 1835: 34. n. syn. [Distribution: Jamaica] 8. Diapherodes laevicollis Redtenbacher, 1908: 434. [Distribution: Jamaica]

TABLE 10. Comparison of the gigantea-group and jamaicensis-group of Diapherodes Gray, 1835. gigantea-group

jamaicensis-group


64.0–126.0 mm

61.5–76.5 mm


92.0–197.0 mm

104.0–164.5 mm

With a pair of prominent anterior spines

No anterior spines

Mesonotum (♂♂)

Granulose or tuberculose

Smooth

Mesonotum (♀♀)

No median carina; surface armed with several blunt tubercles or spines

With an indistinct longitudinal median carina; surface ± granulose or tuberculose

Meso- and metapleurae (♂♂)


Smooth

Meso- and metapleurae (♀♀)

Strongly deflexed and armed with a marginal row of prominent teeth or spines

Gently expanded and with a marginal row of granules or acute tubercles

Meso- and metasternum (♂♂)

Granulose

Smooth

Metasternum (♀♀)


Smooth

Vestigial; < half the length of tegmina

± as long as tegmina

Lateral margins gently rounded

With a small posterolateral lobe or distinctly deflexed laterally

Very small and mostly fully concealed by anal segment

Large triangular to shield-shaped and projecting over anal segment

4.1–5.6 mm

3.5–3.8 mm

Elongate; ± ½ as long as capsule

About as long as wide; ± 1/3 the length of capsule

Lesser Antilles (Fig. 381)

Greater Antilles (Puerto Rico & Jamaica)

Pronotum

Alae (♀♀) Abdominal tergum VII (♀♀) Epiproct (♀♀) Egg (capsule length) Egg (micropylar plate) Distribution


HENNEMANN ET AL.

Keys to the species and subspecies of Diapherodes ♀♀* 1. 2. 3. -

4. 5. 6. -

7. -

Meso- and metapleurae strongly expanded and with a prominent marginal row of teeth or spines; pronotum with a distinct pair of anterior spines; alae vestigial, < ½ the length of tegmina; Lesser Antilles . . . . . . . . . . . . . . . . . . . . . . . . 2. (gigantea-group) Meso- and metapleurae tuberculose; no anterior spines on pronotum; alae ± as long as tegmina; Greater Antilles (Jamaica & Puerto Rico). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6. (jamaicensis-group) Body length > 11 cm; mesonotum > 1.5x longer than head and pronotum combined; > 50 antennomeres. . . . . . . . . . . . . . . . . 3 Body length < 10 cm; mesonotum only 1.3x longer than head and pronotum combined (Fig. 144); 37 antennomeres; Martinique (Fig. 381) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . martinicensis Body length < 15 cm; mesonotum with numerous spiniform tubercles or spines; carinae of mid and hind legs not conspicuously expanded and dorsal carinae not dentate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4 Very large, body length > 15 cm (Figs. 120–122); mesonotum unarmed or at best with six enlarged spines in anterior portion; mid and hind legs with carinae ± strongly expanded and acutely dentate (Fig. 132); Saint Vincent & Grenada (Fig. 381) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . gigantea gigantea Small, body length < 14 cm; head bi-tuberculate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5 Large, body length > 14 cm, head prominently bi-cornute (Fig. 107); Guadeloupe (Fig. 381) . . . . . . . . . . . . . . . . . . . . angulata Spines of meso- and metapleurae considerably longer than width of base, acute; tegmina large and overlapping (Figs. 136–137); Saint Lucia (Fig. 381) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . gigantea saintluciae n. ssp. Spines of meso- and metapleurae small and blunt (Figs. 109–110); tegmina small and leaving a distinct gap between each other (Figs. 109–110); Dominica (Fig. 381). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .dominicae Slender insects; mesothorax slightly widened towards the posterior, > 2.5x longer than head and pronotum combined; abdominal tergum VII gently broadened and indistinctly broader than VI . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7 Very massive insects (Figs. 183–184); mesothorax strongly broadened towards the posterior, only about 2.1x longer than head and pronotum combined; abdominal tergum VII prominently laterally expanded; almost 2x broader than VI (Fig. 186); Jamaica . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .laevicollis Slender insects (Figs. 167–170); head globose and distinctly bi-cornute (Fig. 173–174); mesonotum granulose and with 2–4 anterior spines, not keeled; abdomen slender; Jamaica. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .jamaicensis Broader insects (Fig. 157); head longer than wide and faintly bi-tuberculate (Fig. 159); mesonotum smooth and with a longitudinal median carina; abdomen distinctly broadened sub-basally; Puerto Rico. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . achalus

* body lengths given are including the subgenital plate

♂♂ 1. 2.

3. 4. 5. 6. 7. -

Mesonotum irregularly granulose, tuberculose or spinose; metasternum and pleurae granulose; Lesser Antilles (Fig. 381) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .2 (gigantea-group) Mesonotum unarmed (may have 2–8 anterior spines), metasternum and pleurae unarmed; Greater Antilles (Jamaica & Puerto Rico) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6 (jamaicensis-group) Mesonotum tuberculose or spinose; colouration of body, tegmina and costal region of alae brown . . . . . . . . . . . . . . . . . . . . . . 3 Mesonotum sparsely granulose; body, tegmina and costal region of alae green (Figs. 101–102); Guadeloupe (Fig. 381) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . angulata Head distinctly bi-cornute; anterior margin of tegmina white or pale yellowish . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4 Cephalad pair of tubercles obsolete (Fig. 113); anterior margin of tegmina bright yellow and green with a dark green stripe interiorly (Fig. 117); Dominica (Fig. 381) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .dominicae Mesothorax elongate > 1.7x longer than head and pronotum combined; alae at best reaching to posterior margin of abdominal tergum VI; ventral portion of tegmina without distinct colouration . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5 Mesothorax short, only 1.5x longer than head and pronotum combined; alae reaching to posterior margin of abdominal tergum VII (Figs. 146–147); central portion of tegmina bright purple; Martinique (Fig. 381) . . . . . . . . . . . . . . . . . . . . . . martinicensis Body length > 9 cm; tubercles of pro- and mesonotum dull green (Fig. 125); alae at best reaching half way along abdominal tergum VI; Saint Vincent & Grenada (Fig. 381). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . gigantea gigantea Body length < 9 cm; tubercles of pro- and mesonotum black; alae reaching to posterior margin of abdominal tergum VI; Saint Lucia (Fig. 381). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . gigantea saintluciae n. ssp. Mesonotum with 2–8 distinct paired anterior spines; alae at best reaching to abdominal tergum VI; tegmina plain green; Jamaica . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7 Mesonotum unarmed, at most with some minute granules; alae projecting over tergum VII; tegmina with a brown longitudinal median stripe (Fig. 158); Puerto Rico . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . achalus Slender; mesonotum 2x longer than head and pronotum combined; alae reaching to posterior of abdominal tergum VI (Figs. 171–172); scapus and pedicellus greenish or brown (Fig. 175).. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . jamaicensis More robust; mesonotum 1.8x longer than head and pronotum combined; alae reaching to posterior of abdominal tergum VII (Fig. 185); scapus and pedicellus black . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .laevicollis



63

5.4.1. The gigantea-group Characteristics: Members of this species-group are restricted to the Lesser Antilles (Fig. 381) and are characterised by their large to very large size (body lengths: ♂♂ > 60 mm, ♀♀ > 90 mm) and robust habitus in both sexes (refers to ♀♀ in particular). ♀♀ usually plain bright green (rarely brownish), ♂♂ green or brown; ventral body segments often whitish. Pronotum with a distinct anterior pair of tubercles or spines. ♀♀ with a decidedly broadened mesothorax and a strongly elevated longitudinal marginal row of prominent, pointed spines or teeth on the meso- and metapleurae. Meso- and metapleurae of ♂♂ with a longitudinal row of tubercles. Mesonotum with single prominent spiniform tubercles or spines, and at least in ♂♂ irregularly set with small tubercles and granules. Meso- and metasternum granulose (♂♂) to tuberculose (♀♀). ♀♀ with moderately sized tegmina but very small vestigial alae, these no more than half the length of tegmina. Abdomen of ♀♀ broadened basally, segments II and III broadest segments, following gradually tapering. Tergum VII of ♀♀ with lateral margins gently rounded. Epiproct of ♀♀ very small and mostly fully concealed by anal segment. Most carinae of the legs usually ± granulose or denticulate. Eggs medium-sized to large (capsule > 4.0 mm), micropylar plate generally drop-shaped, longer than wide and about half as long as the capsule. A detailed comparison with the jamaicensis-group is presented in Table 10.

Diapherodes angulata (Fabricius, 1793) rev. stat. (Figs. 100–108, 346, 365, 381) Mantis angulata Fabricius, 1793: 13. Type(s), ♀ (♀): Insula Guadeloupe Dom. De Badier (not traced—presumed lost). [Unnecessary replacement name for Mantis gigas Drury, 1773]. NT (by present designation), ♀: Guadeloupe Le Moult [MNCN]. Phasma angulata, Fabricius, 1798: 187. Phasma angulatum, Lichtenstein, 1802: 11. Cyphocrana angulata, Audinet-Serville, 1831: 61. Diapherodes angulata, Burmeister, 1838: 547. Rehn, 1903: 136. Brock, 1998c: 33. Haplopus grayi Kaup, 1871: 36, pl. 2: 1 (♀) & 20, pl. 1: 20 (eggs). HT, ♀: Haplopus Grayi Kaup Mol. v. Rosenberg, Holotypus [HLDH]. n. syn. Moxey, 1971: 99 (in litt.). [Listed as a synonym of Diapherodes gigas (Drury, 1773)—in error] Diapherodes grayi, Kirby, 1904a: 363. Otte & Brock, 2003: 305. Diapherodes gigantea, Redtenbacher, 1908: 434 (in part). Moxey, 1972: 99 (in litt.; in part). Diapherodes gigas, Moxey, 1971: 98 (in litt.; in part—only specimens from Guadeloupe). Langlois & Lelong, 1997: 43, figs. 20a (♀) & 20 b (♂). Lelong & Langlois, 2001: 242, figs. 1 (♀), 2 (♂). [Not: Phasma angulata Stoll, 1813: 61, pl. 21: 77 (♀), = Haplopus micropterus (St. Fargeau & Audinet-Serville, 1825: 445] Further material [14 ♀♀, 21 ♂♂, 6 nymphs, eggs]: GUADELOUPE: 5 ♀♀, 5 ♂♂, 2 ♂♂ (nymphs): Guadeloupe Le Moult [MNCN]; 1 ♀, 2 ♂♂: Guadeloupe Trois Rivières; D.Vitrac Dedit.; Collection A. Finot, Diapherodes gigas, Drury [MNHN, coll. Finot, box no. 294]; 1 ♀: Guadeloupe Trois Rivières; D.Vitrac Dedit.; Collection A. Finot, Diapherodes gigas, Drury [MNHN]; Eggs (ex abdomen): Diapherodes gigas Drury, Trois Rivières, Guadeloupe, 69.321 [MNHN]; 1 ♀, 1 ♂, 1 ♂ (penultimate instar), 2 ♀♀ n5: Guadeloupe, Gourbeyre 1896 Diapherodes gigas Drury [MNHN]; 1 ♂: Vernou, 15–30-VI-19771; Museum Paris Guadeloupe, A. Villiers [MNHN]; 5 ♂♂: M. Delfosse Elevage ♂ 1993; Diapherodes gigantea gigantea (Gmélin, 1789), E. Delfosse Dét. 23/02/03 [MNHN]; 1 ♀: M. Delfosse, Guadeloupe ♀ Elevage 1994; Diapherodes gigantea gigantea [MNHN]; 1 ♂: M. Delfosse, Basse-Terre, Guadeloupe 90; Diapherodes gigantea gigantea 04/05 [MNHN]; 1 ♀: M. Delfosse, Elevage, Guadeloupe, 1993; Diapherodes gigantea gigantea (Gmélin, 1789) E. Delfosse Dét. 04/05 [MNHN]; 2 ♂♂: M. Delfosse Guadeloupe, 6/1993; Elevage, Diapherodes gigantea [MNHN]; 1 ♂: M. Delfosse, Guadeloupe, 07/1993 [MNHN]; 2 ♂♂: M. Delfosse Guadeloupe ♂ 04/1993; Elevage, Diapherodes gigantea [MHNH]; 1 ♀: Guadeloupe, Antilles ♀; Diapherodes gigas Drury [MHNG]; 1 ♀: Antilles; Diapherodes gigas Drury [MHNG]; 1 ♀: 3/33 Guadeloupe, Mr. H. d. Sauss.; Diapherodes gigas Drury [MHNG]; 1 ♀ (nymph): Larve, Guadeloupe, Mr. H. d. Sauss; Diapherodes gigas Drury [MHNG]. NO DATA: 1 ♀: no data [NHMUK]; 1 ♂: no data [MNHN].


HENNEMANN ET AL.

FIGURES 100–102. Diapherodes angulata (Fabricius, 1793). 100. ♀: Guadeloupe, Trois Rivères [MNHN]; 101. ♂: Guadeloupe [MNHN]; 102. ♂: Guadeloupe [MNHN].

Diagnosis: Distinguished from the type-species D. gigantea gigantea (Gmélin, 1789) by: the smaller size and prominently bi-spinose vertex of both sexes; more numerous but smaller spiniform tubercles of the mesonotum; shorter and more numerous spines of the marginal row of the meso- and metapleurae; more slender legs and much less prominently dentate, not considerably expanded carinae of the mid and hind legs of ♀♀ (Fig. 106). From D. dominicae (Rehn & Hebard, 1938) ♀♀ differ by: the larger size; more robust legs; much more prominent cephalad



65

pair of spines (Fig. 107); less numerous but more pronounced tubercles of the mesonotum; larger anterior pair of spines on the pronotum; larger and more pointed spines of the meso- and metapleurae (Fig. 100) and larger, broader tegmina which overlap interiorly (Fig. 100). ♂♂ are at once distinguished from all other representatives of the gigantea species-group by the green body, tegmina and costal region of the alae and very sparsely granulose mesonotum (Figs. 101–102).

FIGURES 103–108. Diapherodes angulata (Fabricius, 1793). 103. ♀ NT, apex of abdomen (lateral view) [MNCN]; 104. ♀ NT, apex of abdomen (dorsal view) [MNCN]; 105. Apex of ♂ abdomen (lateral view) [MNHN]; 106. Right hind leg of ♀ NT (posterolateral view) [MNHN]; 107. Head and pronotum of ♀ NT [MNHN]; 108. Head and pronotum of ♂ [MNHN].

Description: ♀ (Fig. 100). Medium-sized to large (body length including subgenital plate 141.0–162.5 mm) and moderately broad for the genus. General colouration usually plain pale to mid green, the ventral body surface with a whitish wash; occasionally olive or pale to mid brown specimens may occur. Ventral surface of meso- and metapleurae white, spines of the longitudinal marginal row dull orange to very dark red. Dorsal spines of the head and thorax orange with ochre bases. Tegmina and alae of same colouration as body. Antennae pale brown, the two basal segments greenish. Tarsi greenish pale to mid brown. Eyes dull orange to reddish mid brown. Head: Globose, vertex strongly convex and armed with two prominent, blunt to fairly acute spines; a few small granules are present in the posterior portion (Fig. 107). Eyes circular, convex and of moderate size; their length contained about 2.3x in that of cheeks. Between the bases of the antennae with two rather distinct, oval impressions. Antennae longer than fore legs and laid back, projecting over posterior margin of median segment; with more than 60 segments. Scapus dorsoventrally flattened, 2x longer than wide and gently narrowed towards the base. Pedicellus cylindrical, slightly constricted towards the apex and about half as long as scapus. Thorax: Pronotum slightly shorter and narrower than the head; indistinctly longer than wide and gently widened towards the posterior. In front of the very prominent and well defined, slightly curved transverse median sulcus armed with a large pair of rather long, blunt spines; posterior half with a more or less decided pair of tubercles close to posterior margin and one small tubercle at each posterolateral angle (Fig. 107). Mesothorax 1.6–1.7x longer than head and pronotum combined, and decidedly constricted anteriorly. Mesonotum dorsally armed with two pairs of spiniform tubercles close to anterior margin (Fig. 107) and two or three further pairs of slightly enlarged tubercles in the anterior half of the dorsal surface; otherwise irregularly set with a few small tubercles. Along lateral margins with a longitudinal row of small, rounded granules. Metanotum unarmed, about 2/ 5 the length of mesonotum and about as long as wide. Spines of the meso- and metapleurae very distinct and acute;


HENNEMANN ET AL.

the longest being about 2x as long as its basal width. Mesopleurae with 17–22, metapleurae with 13–17 spines. Meso- and metasternum each with a variable number of pointed tubercles, which are more decided and larger on the metasternum. Tegmina rather large for the genus and usually slightly projecting over posterior of metanotum (length 11.9–13.0 mm) with posterior margin roundly angulate; overlapping or leaving a small space (up to 3 mm) inbetween them. Alae very small, projecting underneath tegmina by only about 0.5–2.0 mm. Abdomen: Median segment slightly longer than metanotum and trapezoidal, being gently narrowed towards the anterior. Segments II–V roughly of equal length, VI and VII slightly shorter. II–IV about 1.6–1.7x wider than long, VI about as long as wide. VII narrower and slightly longer than VI, parallel-sided. Praeopercular organ formed by an elongate, longitudinal swelling near posterior margin of sternum VII (Fig. 346). Tergum VIII slightly about 2/3 the length and narrower than VII, decidedly more convex than previous and gently widened towards the posterior. IX about half as long as VIII and slightly wider than long. Anal segment longer than IX, gently narrowed towards the posterior and with a faint longitudinal median keel dorsally (Fig. 104). Posterior margin broadly rounded and with a very shallow median indentation, the lateral margins with a concave excavation at bases of cerci. Epiproct very small and hidden under anal segment. Subgenital plate long and gradually narrowed towards a pointed tip; projecting over apex of abdomen by at least the combined length of tergites IX and X (Figs. 103–104). Legs: All of moderate length, profemora about as long as mesothorax and metatibiae reaching about halfway along abdominal tergum VIII. All carinae densely and sharply but very minutely granulate or denticulate (more decided on mid and hind legs). Mid and hind legs decidedly carinate (Fig. 106). Medioventral carina of meso- and metafemora blunt but low and with 4–6 stout spines which decrease in size towards the base of femur. Posteroventral carina with one, anteroventral carina with two rather small pointed sub-apical spines. Anterodorsal carina of meso- and metatibiae very weakly rounded apically. Basitarsus about 1.3x longer than second tarsomere. ♂ (Figs. 101–102). Medium-sized to large (body length 93.5–110.0 mm) and stocky for the genus, with long alae (53.0–56.0 mm) and a moderately, dorsally sparingly granulose mesothorax. General colouration pale yellowish green to mid green, the tegminal and costal region of the alae with distinctly darker longitudinal veins. Ventral thoracal segments and abdominal sternum II dull green, abdominal sternites III–VII whitish. Ventral surfaces of meso- and metapleurae white. Largest tubercles of the head and dorsal surface of the thorax tipped with dull orange. Tegmina and costal region of same colouration as body, the tegmina with a broad longitudinal and very pale creamish green stripe along the anterior margin which is continued in the basal quarter of the alae. Bases of alae pale red, anal region hyaline. Antennae pale to mid brown and becoming gradually darkler brown towards the apex, two basal segments greenish. Eyes dark orange to reddish brown. Head: Generally as in ♀♀. Vertex strongly convex and smooth except for two blunt but well decided, conical tubercles; posterior portion usually with two small granules (Fig. 108). Eyes large, cylindrical and projecting hemispherically; their length contained about 2x in that of cheeks. Antennae reaching halfway along abdominal tergum III, otherwise structured like in ♀♀. About 70 antennomeres. Thorax: Pronotum shorter and slightly narrower than head, about as long as wide with the anterior margin slightly widened. Otherwise generally as in ♀♀ but armature less distinct (Fig. 108). Mesothorax rather elongate, about 2x longer than head and pronotum combined; mesonotum almost 6x longer than wide. Anterior of mesonotum with a well decided pair of blunt tubercles, dorsal surface otherwise irregularly set with a few low tubercles and granules; a longitudinal row of small granules along lateral margins. Meso- and metapleurae each with a longitudinal row of minute granules. Meso- and metasternum sparsely granulose. Alae reaching halfway along tergum VI. Abdomen: Tergum VII with posterolateral angles very gently expanded and rounded. Tergum VIII 2/3 of VII and gradually widened towards the posterior, broadest segment. IX ¾ the length of VIII and very weakly narrowed towards the posterior. Posterior half of tergum VII, as well as VIII and IX with two parallel, longitudinal carinae dorsomedially. Anal segment with a very faint longitudinal median keel, the posterior portion abruptly narrowed and with a broadly triangular median indentation. Sternum VIII with an acute longitudinal median carina. Vomer triangular with basal portion strongly broadened and almost equal to overall length, terminal hook short and slender (Fig. 365). Poculum granulose, convex and cup-like (Fig. 105) with the posterior margin broadly triangular (Fig. 365); indistinctly projecting over posterior margin of tergum IX. Legs: All of moderate length for the genus, profemora slightly longer than mesonotum, metatibiae almost reaching posterior margin of abdominal tergum VII. All carinae densely but very minutely granulate to denticulate. Posteroventral carina of meso- and metafemora with one, anteroventral carina with two, slender and pointed subapical spines. Medioventral carina broad and flat, armed with 5–6 short but strong spines which decrease in size



67

towards the base of femur. Tarsi rather stout, slightly less than half the length of corresponding tibia. Basitarsi about 1.3x longer than second tarsomere. Comments: This species was first described by Fabricius (1793: 13) who stated the type locality to be “Insula Guadeloupe”. The original ♀ type specimen(s) is not traced in ZMUC nor in any other likely collection and hence believed lost (Brock, 1998c: 33). Fabricius (1793) based his description on a specimen(s) from Guadeloupe and introduced Mantis angulata as a replacement name for Mantis gigas Drury, 1773, which had however already been replaced by Mantis gigantea Gmélin, 1798. The name of Fabricius is therefore an unnecessary replacement name and a junior objective homonym of Mantis gigantea Gmélin. Diapherodes gigantea (Gmélin, 1789) is endemic to Grenada and Saint Vincent and detailed examination of several specimens, including ♂♂, from Guadeloupe in MNCN and MNHN has shown these to be a distinct species. As the type specimen(s) on which Fabricius (1793: 13) based his description of Mantis angulata were from Guadeloupe and represent a distinct species, Mantis angulata Fabricius is not a synonym of Mantis gigantea and becomes an available name (rev. stat.). Therefore the specific name of Fabricius (1793) is here re-introduced and used for the species from Guadeloupe. As the original type-specimen(s) is lost, an almost perfect ♀ in MNCN from a series of specimens with the data “Guadeloupe Le Moult” is here selected as the neotype of Mantis angulata Fabricius, 1793 in order to ensure stability of the name. Further specimens from the same source are in MNHN. Kaup (1871: 36, pl. 2: 1) described and figured Haplopus grayi from a ♀ wrongly labelled “Molukken”, but there can however be no doubt his specimen originated from the Lesser Antilles. Examination of Kaup's holotype in HLDH has proven this to be clearly distinct from Diapherodes gigantea but to match very well with typical ♀♀ of D. angulata from Guadeloupe. Comparison has proven the synonymy of Kaup's species and suggests this specimen to have originated from Guadeloupe. Moxey (1971: 99) erroneously synonymised Haplopus grayi with Diapherodes gigantea. Diapherodes angulata appears to have been moderately abundant in certain localities in Guadeloupe, which is seen in the good series of specimens collected by Le Moult (now in MNCN and MNHN). It has however become apparently rare and was not encountered during recent extensive collections conducted on the island in the late 1990's (Langlois & Lelong, 1997: 43). D. angulata is endemic in Guadeloupe and all records of D. gigantea from this island are based on misidentifications. Distribution (Fig. 381): Guadeloupe (Basse-Terre [MNHN]; Trois Riviéres [MNHN]), endemic. Number of specimens examined: 43 TABLE 11. Measurements of Diapherodes angulata (Fabricius, 1793) [mm] ♀, NT [MNCN]

♀♀ [MNCN]

♂♂ [MNCN]


146.0

141.0–162.5*

-

Body

135.7

129.5–149.0*

93.5–110.0

Pronotum

7.1

6.1–7.3

3.8–4.0

Mesonotum

24.5

24.6–25.5

15.9–18.8

Metanotum

9.0

8.1–9.0

7.2

Median segment

9.2

9.2–10.1

7.6

Tegmina

13.0

11.9–12.1

14.2–14.8

Alae

4.8

4.5–5.0

53.0–56.0

Profemora

24.5

25.0–27.2

18.0–19.3

Mesofemora

22.7

23.0–23.2

15.8–18.4

Metafemora

32.5

33.5–34.0

21.2–24.3

Protibiae

24.9

23.8–26.2

16.0–18.0

Mesotibiae

21.6

21.2–24.2

13.9–15.2

Metatibiae

29.8

31.0–35.1

20.0–22.0

Antennae

64.0

58.0

> 46.0–53.0

* ♀♀ in MHNG measure 120.0–121.0 mm (126.0–134.0 mm including the subgenital plate)


HENNEMANN ET AL.

FIGURES 109–112. Diapherodes dominicae (Rehn & Hebard, 1938). 101. ♀ HT: Dominica [ANSP]; 102. ♀ Dominica, Croix [coll. ASPER]; 103. ♀ apex of abdomen (lateral view): Guadeloupe, Croix [coll. ASPER]; 104. ♀ apex of abdomen (dorsal view): Guadeloupe, Croix [coll. ASPER].

Diapherodes dominicae (Rehn & Hebard, 1938) n. comb. (Figs. 109–119, 347, 366, 381) Diapherodes gigantea dominicae Rehn & Hebard, 1938: 53, pl. 4: 22 (♀). HT, ♀: Dominica, B. W. I., II.1927 James Bond, Diapherodes gigantea dominicae, Type, R & H, ANSP 5539, WO188, = Diapherodes gigas (Drury), det. C.F. Moxey 1972 [ANSP, No. 5539]. Otte & Brock, 2003: 304. Diapherodes gigas, Moxey, 1971: 98, 99 (in litt.; in part—only specimens from Dominica) Diapherodes gigantea, Langlois & Lelong, 2010: 60, figs. 19 (in part—only records from Dominica). Diapherodes sp. Lelong, Langlois, Rastel & Dorel, 2003: 48, figs. 19a–g (♀, ♂, nymph) & REM-photos 11–13 (egg). Further material [6 ♀♀, 4 ♂♂, eggs]: DOMINICA: 1 ♀: Dominica 1937, Walter H. Hodge; USNM; ANSP; w0187 [USNM]; 1 ♂: Dominica, Clarke Hall, 7.II.1965; J.F.G. Clarke Thelma M Clarke; Bredin-Archold-Smithsonian Bio. Surv. Dominica; USMNH; w0189 [USNM]; 1 ♀: Dominique, Croix, alt. 808 m, 15°20’49’’N 61°18’23’’E, 26X.2000 [coll. ASPER, No. DOM00-101]; 2 ♂♂, 2 ♀♀, eggs: Dominique, Bellevue Mountain, alt. 455–533 m, 15°37’22’’N 61°26’03’’E, 29.X.2000 [coll. ASPER, No’s DOM00-106 & 192 to 194]; 1 ♀: Dominique, Morne Macaque, alt. 762–855 m, 15°20’31’’N 61°18’38’’E, 20.X.2000 [coll. ASPER, No. DOM00-190]; 1 ♂: Dominique, Picard River track, alt. 26–146 m, 15°33’14’’N 61°27’06’’E, 30.X.2000 [coll. ASPER, No. DOM00-191].



69

FIGURES 113–119. Diapherodes dominicae (Rehn & Hebard, 1938). 113. ♂ head and pronotum: Guadeloupe, Picard River track [coll. ASPER]; 114. ♀ HT, head and pronotum [ANSP]; 115. ♂ apex of abdomen (lateral view) [coll. ASPER]; 116. ♂ apex of abdomen (dorsal view) [coll. ASPER]; 117. Tegmina of ♂ [coll. ASPER]; 118. Egg (lateral view), reproduced from REM-photograph by Lelong et al. (2003: appendix II, fig. 11); 119. Egg (dorsal view), reproduced from REM-photograph by Lelong et al. (2003: appendix II, fig. 12).

Diagnosis: Differing from all other members of the gigantea species-group of Diapherodes by the fairly obsolete cephalad spines, which are merely represented as rounded tubercles (Fig. 114). Females are similar to D. angulata (Fabricius, 1793) from Guadeloupe but differ by: the smaller size; comparatively more slender legs; more numerous but smaller tubercles of the mesonotum; much smaller anterior pair of tubercles on the pronotum (Fig. 114); comparatively smaller spines of the longitudinal marginal row of the meso- and metapleurae and smaller not overlapping tegmina (Fig. 109–110). Males strongly resemble those of D. gigantea saintluciae n. ssp. but differ by: the broader, laterally rounded and apically less acuminate vomer (Fig. 359); bright green legs (at best with a slight greenish hue in gigantea); greenish brown antennae and distinctive colouration of the anterior margin of the


HENNEMANN ET AL.

tegmina and alae, which is bright yellow and then gradually becomes dull green towards the interior (Fig. 117). From D. angulata ♂♂ at once differ by the brown body, tegmina and alae as well as the strongly tuberculate mesonotum. Description: ♀ (Figs. 109–110). Fairly small members of the gigantea species-group (body length including subgenital plate 116.0 to ca. 126.0 mm) and moderately robust for the genus (maximum body width at abdominal segment II 15.0–16.8 mm), with fairly small and slender tegmina (9.0–10.0 mm). General colouration plain apple green, the tubercles and granules of the head and thorax ochre to orange, the larger ones tipped with dull red or mars brown. Antennae straw and becoming mid brown towards the apex; two basal segments with a greenish hue. Ventral body surface dull green with a whitish hue. Head: Vertex rounded, globose and armed with two ± decided, blunt tubercles, the posterior portion set with about ten small granules (Fig. 114). Eyes fairly small, circular in outline and their length contained almost 3x in that of cheeks. Antennae with 51 antennomeres and roughly reaching to posterior margin of metanotum. Thorax: Pronotum a little shorter but about as wide as head, the transverse median sulcus very pronunced and almost reaching to lateral margin of segment. A pair of low blunt spines in anterior portion and a pair of lower rounded tubercles at posterior margin; otherwise with a few granules in posterior half (Fig. 114). Mesothorax fairly short and roughly 1.5x longer than head and pronotum combined; posterior margin more than 2x the width of anterior margin, the anterior portion gradually widened and the posterior half almost parallel-sided. Most parts of mesonotum irregularly set with fairly small spines and tubercles, which decrease in size towards the posterior; usually two pairs of somewhat enlarged but blunt spines close to anterior margin (Fig. 114) and 2–4 somewhat enlarged spines pre-medially. The lateral portions unarmed. Metanotum with a few very low granules in anterior portion. Spines of the marginal row of the meso- and metapleurae rather small and blunt, the largest spine not twice as long as its basal width; mesopleurae with 17–20 and metapleurae with about 10–12 spines (Figs. 109–110). Meso- and metasternum sparsely granulose. Tegmina fairly small and comparatively slender, hardly reaching posterior margin of metanotum, not overlapping and leaving a great space inbetween each other (Figs. 109–110). Alae much smaller and at best projecting underneath tegmina by 2 mm. Abdomen: Segment II about equal in length to median segment, widest part of body and very slightly widened towards the posterior; about 1.6x wider than long. III–VII gradually narrowing and roughly equal in length; VII with the lateral margins sometimes gently rounded. Sternites smooth and with two fine longitudinal carinae laterally. Praeopercular organ on sternum VII formed by a fairly decided tubercle and a C-shaped carina (Fig. 347). Tergum VIII narrowed medially and about 1.2x longer than wide. Anal segment narrowed towards the apex, hardly longer than wide and with posterior margin broadly rounded (Fig. 112). Epiproct very small and fully concealed by anal segment. Subgenital plate long and gradually narrowed towards a pointed tip; projecting over apex of abdomen by roughly the combined length of tergites IX and X (Figs. 111–112). Legs: All of moderate length and fairly slender for the gigantea species-group. Meso- and metafemora with the outer carinae minutely granulose; medioventral carina with 6–8 fairly short but acute spines, two outer ventral carinae each with a single sub-apical spine. Posteroventral carina of meso- and metatibiae minutely denticulate. Anterodorsal carina with a low triangular lobe sub-basally and slightly triangularly expanded sub-apically. Basitarsi almost as long as following two tarsomeres combined. ♂. Of moderate size (body length 82.0–98.0 mm) and fairly robust for the genus, with long alae (47.0–52.0 mm). Colouration of the body pale to mid brown, the ventral body surface dull ochre with a whitish wash; abdominal sternites slightly greenish and II–VII each with two black spots post-medially. Head dull ochre (Fig. 113), antennae greenish ochre. Legs mid green with the apex of all femora and tibiae dull green to mid brown. Thoracic armature dull green, the largest tubercles and spines tipped with very dark reddish brown. Tegmina and costal region of alae pale to mid brown with all veins dull green. Anterior margin of tegmina bright yellow, then gradually becoming green and the green section interiorly bordered by a dull bluish green longitudinal stripe (Fig. 117). Central hump dull brown. Basal portion of anterior margin of alae coloured like tegmina. Anal fan of alae transparent orange. Head: About 1.2x longer than wide, the cheeks almost parallel-sided. Vertex gently rounded and with a pair of very low blunt tubercles, otherwise irregularly scattered with small, rounded granules (Fig. 113). Eyes circular in outline, projecting hemispherically and their length contained about 1.5x in that of cheeks. Antennae robust and consisting of 68 antennomeres. Scapus compressed dorsoventrally, 1.3x longer than wide with lateral margins decidedly deflexed and rounded.



71

Thorax: Pronotum about equal in length to head, the anterior margin decidedly wider than posterior margin and the transverse median sulcus very pronounced. Anterior portion with a pair of fairly distinct, blunt spines, the posterior portion with a pair of low tubercles behind transverse sulcus and a pair of slightly more distinct tubercles at posterior margin (Fig. 113). Mesothorax about 1.6x longer than head and pronotum combined. Mesonotum all over with irregularly scattered blunt tubercles, which decrease in size towards the posterior; two pairs of blunt spines near anterior margin (Fig. 113) and a longitudinal row of rounded tubercles along lateral margins. Meso- and metapleurae with a marginal row of differently sized blunt tubercles. Meso- and metasternum irregularly scattered with small granules and various low, rounded tubercles. Tegmina with central hump fairly pronounced and convex (Fig. 117). Alae reaching about half way along abdominal tergum VI. Abdomen: Segments II–VII very slightly gradually narrowing and decreasing in length; tergum VII with the posterolateral angles gently deflexed. Tergum VIII strongly widened towards the posterior and trapezoidal in dorsal view; VIII–X with a longitudinal median carina, which is blunt on VIII and IX but fairly fine on X. Anal segment with lateral margins almost parallel-sided, the posterior margin with a broad and shallow median indentation (Fig. 116). Cerci with apical half round in cross section and the apex blunt. Vomer broad, rounded and with the apical portion gradually narrowing towards a fairly short apical hook (Fig. 366). Poculum with basal portion roundly convex (Fig. 115), otherwise fairly flat and with the posterior margin broadly rounded. Legs: All of moderate length and fairly robust; metafemora reaching about half way along abdominal segment IV and metatibiae reaching to posterior margin of abdominal segment VI. All outer carinae of mid and hind legs very minutely denticulate. Medioventral carina of mesofemora only with two fairly prominent and acute spines in apical portion in metafemora with five prominent spines, that decrease in size towards the base. Two outer ventral carinae each with a single sub-apical spine. Basitarsus about 1.3x longer than second tarsomere. Egg (Figs. 118–119): Of moderate size (capsule length 4.78 mm), capsule about 1.7x longer than wide. Polar area slightly impressed if seen in lateral aspect. Capsule surface to various degrees covered with irregular, distinctly raised areas covered with clusters of sponge-like structured tubercles and granules. These form a conspicuous longitudinal ridge from the posterior end of the micropylar plate to the polar-area. Opercular collar decidedly marginated. Micropylar plate rather large and about half as long as capsule. Entire surface, except for a stripe along the outer margin, covered with irregular granules and raised ridges. Centre with an elongate, impressed region. Micropylar cup distinct and placed in a very deep central gap of posterior margin of plate. Operculum in the centre with a prominently raised circular rim and a conical tubercle in the centre. General colouration pale brown. Outer margin of micropylar plate and micropylar cup darker brown. Measurements according to Lelong, Langlois, Rastel & Dorel (2003: 51) [mm]: Length 4.78, width 2.72, height 3.05, diameter of operculum 1.70. Comments: Rehn & Hebard (1938: 53) described Diapherodes gigantea dominicae based on a single ♀ in ANSP, which was collected by the leading American ornithologist James Bond. They distinguished it from D. gigantea (Gmélin, 1789) by the smaller size, smaller spines of the mesothorax (a very variable feature in gigantea), smaller spines of the meso- and metapleurae and unarmed carinae of the legs. But in fact, all these characters of the ♀♀ place it much closer to D. angulata (Fabricius, 1793) from neighbouring Guadeloupe rather than D. gigantea from Grenada and St. Vincent. Males however differ from D. angulata by a good number of obvious features and rather resemble those of D. gigantea or D. martinicensis Lelong & Langlois, 2005. Consequently, detailed examination of Dominican specimens has shown these to represent a separate and valid species (n. comb.). Moxey (1971: 99) erroneously synonymised D. gigantea dominicae Rehn & Hebard with Diapherodes gigantea. Lelong, Langlois, Rastel & Dorel (2003) provided detailed descriptions and illustrations of the adult ♀ and ♂, nymphs and eggs as well as information on the habitats and biology of Diapherodes dominicae in Dominica, their specimens being provisionally identified as “Diapherodes sp.”. They stated this species to be not particularly abundant with only single specimens found in each prospected locality. In their natural habitats, which include montane and lowland hygrophilic forests as well as cultivated zones in xero-mesophilic environments, these insects are usually found at heights of 1.5–3 metres off the ground and feed on guava (Psidium guajava, Myrtaceae) and mapou-baril (Sterculia caribea, Sterculiaceae). In captivitiy in Europe Lelong, Langlois, Rastel & Dorel (2003: 48) reported it to accept bramble (Rubus fruticosus, Rosaceae) as an alternative food-plant. Distribution (Fig. 381): Dominica (Croix, 808 m; Bellevue Mountain 455–533 m & Picard River track, 26–146 m) [USNM; coll. ASPER]. Endemic. Number of specimens examined: 11


HENNEMANN ET AL.

TABLE 12. Measurements of Diapherodes dominicae (Rehn & Hebard, 1938) n. comb. [mm] ♀, HT dominicae [ANSP]

♀♀ *

♂♂ *

> 123.3

116.0–120.0

-

121.8

-

82.0–98.0

Pronotum

7.0

5.5–6.0

3.5–4.0

Mesonotum

24.7

20.5–21.0

13.0–17.0

Metanotum

8.3

17.0–17.5**

12.0–13.5**

Median segment

9.1

-

-

Tegmina

9.7

9.0–10.0

10.0–12.5

-

-

47.0–52.0

Profemora

22.0

20.0–21.0

12.0–18.0

Mesofemora

18.5

17.0

11.0–12.5

Metafemora

26.2

23.0–24.0

18.5–20.5

Protibiae

20.1

-

-

Mesotibiae

16.0

-

-

Metatibiae

24.7

-

-

Antennae

> 33.5

46.0–47.0

45.0–56.0

Body (incl. sg. pl.) Body

Alae

* according to Langlois & Lelong, 2010: 60 ** including the median segment

Diapherodes gigantea gigantea (Gmélin, 1789) (Figs. 120–135, 348, 368, 381, 392–393, 396, 406–408) Mantis gigantea Gmélin, 1789: 2055. HT, ♀: St. Vincent; BMNH(E) #878212 [NHMUK]. [Replacement name for Mantis gigas Drury, 1773]. Diapherodes gigantea, Kirby, 1904a: 362. Rabaey, 2004: 10, figs. 1, 2 & plate figs. 1 (adult couple) & 2. Diapherodes gigantea gigantea Rehn & Hebard, 1938: 53. Otte & Brock, 2005: 120. [„Type lost“] Mantis gigas Drury, 1773: 89, pl. 50 (♀). HT, ♀: St. Vincent; BMNH(E) #878212 [NHMUK]. [Junior homonym of Gryllus (Mantis) gigas Linné, 1758] Diapherodes gigas, Gray, 1835: 33. Westwood, 1859: 84. Saussure, 1871–1872: 184. Brunner v. Wattenwyl & Redtenbacher, 1892: 208. Brunner v. Wattenwyl, 1893: 606. Redtenbacher, 1908: 434, pl. 20:1 (♀). Moxey, 1971: 98 (in litt.; in part—only specimens from Saint Vincent & Grenada). Bradley & Galil, 1977: 189. [Mantis gigas Drury, 1773 removed from homonymy] Phasma (Diapherodes) gigas, Westwood, 1873: 100, pl. 50 (♀). Cyphocrana cornuta St. Fargeau & Audinet-Serville, 1825: 445. HT, ♀: Diapherodes gigas Drury, Saint Vincent [NHMUK]. [Unnecessary replacement name for Mantis gigas Drury, 1773 → Junior objective synonym of Mantis gigantea Gmélin, 1789] [Not: Gryllus (Mantis) gigas Linné, 1758: 425; → Valid name: Phasma gigas (Linné, 1758), Phasmatinae: Phasmatini] [Not: Haplopus grayi Kaup, 1871: 36, erroneous synonym of Moxey, 1971: 99] [Not: Diapherodes gigantea dominicae Rehn & Hebard, 1938: 53, erroneous synonym of Moxey, 1971: 99] [Not: Diapherodes gigas, Langlois & Lelong, 1997: 43, figs. 20a (♀) & 20 b (♂), misidentification à Specimens from Guadeloupe are Diapherodes angulata angulata (Fabricius, 1793)].



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[Not: Diapherodes gigas, Lelong & Langlois, 2001: 242, figs. 1 (♀), 2 (♀), misidentification → Specimens from Guadeloupe are Diapherodes angulata angulata (Fabricius, 1793)]. [Not: Langlois, Lelong & Dorel, 2006: 42, figs. 19 a–g (♂, ♀, nymph), photos 5–6 (♀) & 27–29 (egg, REM-photos) à here described as D. gigantea saintluciae n. ssp.]. [Not: Langlois & Lelong, 2010: 69 à records from Dominica are D. dominicae (Rehn & Hebard, 1938)]. [Not: Langlois & Lelong, 2010: 69 à records from Saint Lucia are D. gigantea saintluciae n. ssp.]. Further material [50 ♂♂, 52 ♀♀, 9 nymphs, eggs]: SAINT VINCENT: 1 ♀: Saint Vincent, W. I., H.H. Smith, Collectio Br. v. W., det. Br. V. W. Diapherodes gigas Drury, 97. Said to be found principally in high trees: one specimen was obtained on a guava-tree. Second specimen wet in Nov. and one in May. In living specimens the ground colour is a pea-green, spines above brown, those at the legs black. [NHMW, No. 838]; 1 ♀: St.Vincent, W. I., H.H. Smith, Collectio Br. v. W., det. Br. V. W. Diapherodes gigas Drury, 18.737 [NHMW, No. 838]; 1 ♀: Saint Vincent, W. I., H.H. Smith, 9b, Saint Vincent W. I., 95–206 (NHMUK); 1 ♂: 321, Saint Vincent, W. I., H.H. Smith, Edge of Fonst. Sunfriese [Sufriére] Volcano, Leeward side 1200 ft., Jan. 5 on leaf of Heliconia or “Wild Plantain”, Diapherodes gigas [NHMUK]; 1 ♂: 40, 4-2, 1936, Diapherodes G. R. Gray, Diapherodes gigas (Drury), Saint Vincent [wings spread] [NHMUK]; 1 ♀: Saint Vincent, W. Indies, 1907-296, Pres. By Miss Patterson [NHMUK]; 1 ♀: Sharp’s Valley, Saint Vincent, B.W.I., VIII.25.1941, R. S. Fennah [ANSP]; 1 ♀: Saint Vincent, W. Indies, H.H. Smith; A.N.S.P., Ex Carn. Mus, Bruner Cln. [ANSP]. GRENADA: 1 ♀: St. Georges, Grenada, July 1969, Frank E. Miller; Diapherodes gigas (Drury), det. A Gurney 1969 [USNM]; 1 ♀: Grenada, W. I., H.H. Smith, Collectio Br. v. W., det. Br. V. W. Diapherodes gigas Drury, 19.660, May 15. Purchased. This species is often seen on the Cashew-tree (Anacardiaceaee) [NHMW, No. 838]; 1 ♂ (penultimate instar): Vendome Est. (Leeward side), Grenada, W. I., H.H. Smith, Vendome Estate Leeward, 1200 ft. open place, Sept. 8, on Guava bush [NHMUK]; 1 ♂: Grenada W. I., H.H. Smith, Grenada W. I., 95-206, St. Catharinés Rictory (Winward), 500 ft. June 10, “Left hind wing used for drawing” Ragge [NHMUK]; 1 ♂ (nymph n3): West Indies: Grenada, Par. St. Andrew, Birch Grove, 8-VI-1990, coll. M.C. Thomas [FSCA]; 1 ♀: Grenada, B.W.I. [ANSP]; 1 ♀: Grenada, W. Indies, H.H. Smith; A.N.S.P., Ex Carn. Mus, Bruner Cln. [ANSP]; 1 ♂: Diapherodes gigantea (Gmélin, 1789). Elevage Grenade Don Alexandre Lerch 2007, E. Delfosse Dec.2007 [MNHN]; 1 ♂: M. Delfosse, Grenade, ♂ 3/12/2012 [MNHN]; 1 ♂: M. Delfosse, Grenade, ♂ 23/11/2012 [MNHN]; 1 ♀: M. Delfosse, Grenade, Elevage ♀ 22/07/2007 [MNHN]; 2 ♀♀: ex Zucht: K. Rabaey (Belgien), urspr.: Grenada, III.–VIII. 2004 [coll. FH, No’s 0359-1 & 2]; 2 ♂♂: ex Zucht: F. Hennemann, urspr.: Grenada, III.–VIII. 2004 [coll. FH, No’s 0359-3 & 4]; 1 ♀, 2 ♂♂, 1 ♀ (subadult), 4 ♀♀, 1 ♂ (nymphs), 1 Gynandromorph (subadult), eggs: ex Zucht: F. Hennemann, urspr.: Grenada, 2005 [coll. FH, No’s 0359-5 to 14 & E]; 7 ♂♂, 7 ♀♀: ex Zucht: F. Hennemann 2007/08, Herkunft: Grenada, leg. T. & P. James, PSG No. 260 [coll. FH, No’s 035915 to 30]; 5 ♂♂, 2 ♀♀: ex Zucht: F. Hennemann 2009, Herkunft: Grenada, leg. T. & P. James, PSG No. 260 [coll. FH, No’s 0359-31 to 38]; 30 ♂♂, 26 ♀♀, eggs: ex Zucht. O. Conle, Herkunft: Grenada [coll. OC]. TRINIDAD [in error ?]: 1 ♀: 89–69: Trinidad [NHMUK]. NO / UNPRECISE DATA: 1 ♀: no data [wings spread] [NHMUK]; 1 ♀: 40, 4-2, 1329 [presumably from Saint Vincent] [NHMUK]; 1 nymph (n1): 40, 42, 1366 [presumably from Saint Vincent] [NHMUK]; 1 nymph (n1): 40, 4-2, 1367 [presumably from Saint Vincent] [NHMUK]; 2 eggs: 40, 4-2, 1370 [presumably from Saint Vincent] [NHMUK]; 4 eggs: 40, 4-2, 1369 [presumably from Saint Vincent] [NHMUK]; 1 ♀: 40, 4-2, 1365 [wings spread—presumably from Saint Vincent] [NHMUK]; 1 ♀: 59-57, Vigors Coll. [NHMUK]; 1 ♀: West Indies, H. A. Ballou [ANSP].

Diagnosis: This is the largest and most southward distributed species in the genus. Females are well characterized by the ± expanded and dentate carinae of the mid and hind legs (metatibiae in particular, Fig. 132) and sparse armature of the mesonotum (Fig. 126). From D. angulata (Fabricius, 1793) from Guadeloupe it differs by: the larger size of both sexes; much less numerous spiniform tubercles of the mesonotum; decidedly longer but less numerous spines of the marginal row on the meso- and metapleurae (Figs. 126–127); much broader legs and prominently dentate, ± distinctly expanded carinae of the mid and hind legs of ♀♀ (Fig. 132). Males differ from those of D. angulata by: the brown general colouration (Figs. 123, 380); relatively shorter mesothorax; more prominent spiniform tubercles on the mesonotum and translucent orange anal region of the alae (hyalinous in angulata). Eggs principally differ by the considerably larger dimensions and less strongly sculptured capsule surface (Figs. 134–135). Description: ♀ (Figs. 120–122, 392–393). Large to very large (body length including subgenital plate 153.0–197.0 mm) and broad for the genus (maximum body width at abdominal tergum II 15.0–19.0 mm), mid and hind legs with carinae ± expanded and denticulate (metatibiae in particular). General colouration usually plain bright apple-green, the ventral body surface mid green and with a whitish wash on abdominal sternites II–VII,


HENNEMANN ET AL.

more rarely with a black wash; occasionally olive or slightly brownish green specimens occur. Transverse borders between the abdominal tergites yellow to orange. Spines of the marginal expansion of the meso- and metapleurae dull pink to purple (bluish green in young specimens). Dorsal spines of the thorax with the bases yellow and tips orange. Antennae orange to reddish brown; scapus green. Tegmina and alae bright green. Tarsi yellowish or greenish pale to mid brown. Eyes yellowish to reddish brown. Head: Globose, convex, 1.2x longer than wide and armed with two more or less decided humps, tubercles or blunt spines on vertex (Fig. 124). Back with a variable number of small granules, which are roughly arranged in four parallel, longitudinal rows; anterior portion sometimes set with a few scattered granules. Eyes circular, convex and rather small; their length contained almost 2.5x in that of cheeks. Antennae reaching to abdominal segment III; with about 55 segments. Scapus dorsoventrally flattened, 2.2x longer than wide and gently narrowed towards the base. Pedicellus cylindrical and about 2/5 the length of scapus. Thorax: Pronotum slightly shorter and narrower than the head; indistinctly longer than wide. In front of the very prominent transverse median sulcus armed with a pair of large, blunt spines (Fig. 124); posterior half with a more or less decided pair of tubercles close to posterior margin (sometimes lacking). Mesothorax almost 2x longer than head and pronotum combined, and moderately constricted anteriorly. Mesonotum dorsally armed with one or two pairs of prominent blunt spines close to anterior margin and one or two further pairs of slightly smaller spines in the anterior half of the dorsal surface; sometimes specimens may occur which lack any distinct spines and have the mesonotum with only a few small granules (Figs. 124, 126). Metanotum unarmed, about 2/5 length of mesonotum, gently narrowing towards the posterior and about as long as wide. Spines of the meso- and metapleurae very distinct and acute; the longest being almost 3.5x as long as its basal width (Figs. 126-127). Mesopleurae with 11–17, metapleurae with 11–14 spines. Meso- and metasternum each with a variable number of pointed tubercles (Fig. 127), which are generally more decided and larger on the metasternum. Tegmina fairly large for the genus and usually reaching to posterior margin of metanotum (length 11.6–15.3 mm) with posterior margin roundly angulate; usually leaving a space inbetween them and rarely overlapping. Alae very small and projecting underneath tegmina by about 1.0–3.0 mm. Abdomen: Median segment slightly longer than metanotum and trapezoidal, being gently narrowed towards the margin. Segments II–V of equal length, VI–VII slightly shorter. Lateral margins of III–V gently rounded. II–IV about 1.5–1.8x wider than long, V about as long as wide, VI slightly longer than wide. VII narrower and slightly longer than VI with the lateral margins ± rounded posteriorly. Praeopercular organ on sternum VII formed by a short, elongated carina which terminates in a blunt tubercle (Fig. 348). Tergum VIII about 3/5 the length and decidedly narrower than VII, considerably more convex than previous. IX less than half the length of VIII, roughly quadrate. Anal segment longer than IX, the posterior half gently narrowing towards the apex and with a faint longitudinal median carina dorsally. Posterior margin rounded and with a small, triangular median indentation (Fig. 131). Subgenital plate long and gradually narrowed towards a pointed tip; projecting over apex of abdomen by about the combined length of tergites IX and X (Figs. 130–131). Legs: All of moderate length and strikingly broad, with several of the carinae of the mid and hind legs ± elevated. Profemora about as long as mesothorax and metatibiae reaching about half way along abdominal tergum VII. Posterodorsal carina of profemora with a few minute granules; remaining carinae of front legs unarmed. Mid and hind legs strongly carinate with all carinae densely and ± distinctly denticulate to roughly dentate (Fig. 132). Femora with posterodorsal and anteroventral carina ± strongly expanded and lamellate or ledge-like. Medioventral carina faint and with 5–11 spines, which decrease in size towards the base of femur. Posteroventral carina with one, anteroventral carina with two moderately distinct, pointed sub-apical spines. Anterodorsal carina of tibiae raised and ± elevated and rounded apically. Medioventral carina prominently raised and strongly rounded sub-basally. Armature and expansions considerably more decided on hind legs. Basitarsi indistinctly longer than second tarsomere. ♂ (Figs. 123, 396). Large (body length 92.0–126.0 mm) and moderately stocky for the genus, with long alae (50.0–69.5 mm) and a comparatively short sparsely spinose mesothorax. General colouration of head and body pale yellowish to creamish mid brown; ventral body segments whitish. Thoracic armature mid to dull green (Fig. 125). Tegmina and costal region of alae creamish mid brown, the tegmina with a broad, longitudinal white stripe along the anterior margin, continued in the basal quarter of the alae. Anal region of alae pale translucent orange. Antennae reddish brown, two basal segments brown. Eyes dark reddish brown.



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FIGURES 120–123. Diapherodes gigantea gigantea (Gmélin, 1789). 120. ♀: Saint Vincent [NHMW, No. 838]; 121. ♀: captive reared from Grenada [coll. FH, No. 0359-1]; 122. ♀: captive reared from Grenada [coll. FH, No. 0359-28]; 123. ♂, captive reared from Grenada [coll. FH, No. 0359-33].

Head: Generally as in ♀♀, vertex strongly convex with two ± distinct cephalad tubercles or small spines and a few scattered small granules in posterior portion (Fig. 125). Eyes large, very slightly oval in outline and projecting hemispherically; their length contained slightly less than 2x in that of cheeks. Antennae reaching to posterior margin of abdominal tergum III, otherwise as in ♀♀ and with 54–56 antennomeres.


HENNEMANN ET AL.

FIGURES 124–135. Diapherodes gigantea gigantea (Gmélin, 1789). 124. ♀ head and pronotum [coll. FH, No. 0359-29]; 125. ♂ head and pronotum [coll. FH, No. 0359-33]; 126. ♀ mesothorax (dorsal view) [coll. FH, No. 0359-29]; 127. ♀ mesosternum [coll. FH, No. 0359-29]; 128. ♂ apex of abdomen (lateral view); 129. ♂ apex of abdomen (dorsal view); 130. ♀ apex of abdomen (lateral view); 131. ♀ apex of abdomen (dorsal view); 132. Right hind leg of ♀ from Grenada (posterolateral view) [coll. FH, No. 0359-1]; 133. Right hind leg of ♂ from Grenada (posterolateral view) [coll. FH, No. 0359-33]; 134. Egg (dorsal view) [coll. FH, No. 0359-E] 135. Egg (lateral view) [coll. FH, No. 0359-E].



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Thorax: Pronotum about as long and broad as head and gently narrowed towards the posterior. Transverse median sulcus very prominent, slightly curved and reaching lateral margins of segment. Anterior half armed with a median pair of spiniform tubercles or blunt spines; posterior margin with four small, but acute granules, two median ones and one at each outer angle (Fig. 125). Mesothorax of moderate length, about 1.7–1.8x longer than head and pronotum combined; mesonotum about 5x longer than wide. Dorsal surface irregularly set with a variable number of small tubercles and granules; a longitudinal row of small granules along lateral margins. Anterior of mesonotum with two pairs of rather distinct blunt spines (Fig. 125), and usually there is a further pair of ± enlarged pre-median spiniform tubercles. Meso- and metapleurae with a longitudinal row of minute granules. Meso- and metasternum sparingly granulose. Tegmina with a blunt but well decided hump pre-medially. Alae reaching about half way along abdominal tergum VI. Abdomen: Segments II–VII very slightly tapering gradually, II–IV roughly equal in length and V–VII somewhat decreasing in length. Tergum VII very slightly widened towards the posterior and occasionally with the lateral margins slightly deflexed and rounded posteriorly (Fig. 128). Tergum VIII 2/3 the length of VII and gradually widening towards the posterior. IX ¾ the length of VIII and gently narrowing towards the posterior. Anal segment with a very faint longitudinal median keel, the posterior margin broadly rounded and with a ± distinct median indentation (Fig. 129). Vomer about 1.3x longer than maximum width, basal portion broadly rounded, the terminal hook straight acutely pointed of moderate length and bluntly keeled ventrally (Fig. 368). Poculum granulose, convex and cup-like with a blunt central hump and a small median incision at posterior margin (Fig. 368) and very slightly projecting over posterior margin of tergum IX (Fig. 128). Legs: All of moderate length, profemora almost as long as pro- and mesonotum combined, metatibiae almost reaching to posterior margin of abdominal tergum VI. All carinae densely but very minutely denticulate. Posteroventral carina of meso- and metafemora with one, anteroventral carina with two sub-apical spines. Medioventral carina broad and flat, armed with 4–8 short but strong spines in apical half of femur, which strongly decrease in size and disappear about halfway towards the base (Fig. 133). Tarsi rather stout, slightly less than half the length of corresponding tibia. Basitarsi about 1.3x longer than second tarsomere. Nymphs: Newly hatched nymphs are rather robust and have a body length of ±20.0 mm. The colouration is mid green, the body dorsally marbled with some brown and the pronotum with a brown longitudinal median stripe. Coxae, protibiae and antennae brown. Later instars vary from pale green over straw to dark brown and sometimes specimens may possess bold whitish markings on the thorax and abdomen. The sexes can be distinguishes from 3rd instar onwards, but both have abdominal tergum VII with a rounded lateral lobe throughout their entire development which only disappears with the final ecdysis. Variability: This species shows considerable variability in the size and degree of the body and leg armature. Specimens from the type-locality Saint Vincent are on average larger and have the body and leg armature more prominently developed, particularly in ♀♀. While the mesonotum is unarmed or only bears a few moderately distinct spines in ♀♀ from Grenada, it is armed with 6–8 prominent, blunt spines in specimens from Saint Vincent. The distinct pair of tubercles on the vertex seen in ♀♀ from Saint Vincent is often lacking in the Grenada colony. The degree of expansion of the carinae of the mid and hind legs and the degree of leg armature of ♀♀ generally depends on the general size of the insects. The colouration of ♀♀ is usually bright apple-green, but more rarely olive or slightly brownish green specimens may occur. Egg (Figs. 134–135): The following description is based on a large number of eggs in the first author's collection (coll. FH) laid by captive reared specimens originating from Grenada. Large, barrel-shaped and fairly elongate, capsule almost 2x longer than wide. Polar area very gently impressed if seen in lateral aspect. Capsule surface to a variable degree covered with irregular areas and clusters of slightly sponge-like structured tubercles and raised ridges, as well as numerous single wart-like tubercles. These often form a ± decided longitudinal ridge from the posterior end of the micropylar plate to the polar-area. Opercular collar decidedly marginated. Micropylar plate rather large and slightly less than half the length of capsule. Entire surface, except for a narrow space along the outer margin, covered with the same sponge-like structures seen on the capsule. Micropylar cup distinct. Operculum in the centre with a prominent, raised circular rim, dorsally with ± spiniform tubercles. General colouration pale to dark brown, the raised structures often paler brown than lower parts of capsule. Outer margin of micropylar plate and micropylar cup blackish brown. Measurements [mm]: Length including operculum 4.9–5.8, length 4.7–5.6, width 2.9–3.3, height 2.9–3.2, length of micropylar plate 1.8–2.2.


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Comments: This striking species was first described by Drury (1773: 89) as Mantis gigas. As Drury's species was at that time placed in the same genus as Gryllus (Mantis) gigas Linné, 1758, Gmélin (1789: 2055) was in his rights to introduce Mantis gigantea as a replacement name for Drury's species. Although these two taxa were placed in two distinct genera subsequently, Gmélin's Mantis gigantea is the valid name of the species originally described and figured by Drury. With the exceptions of Kirby (1904a), Rehn & Hebard (1938) and Otte & Brock (2003) Gmélin's valid replacement name was not accepted by subsequent authors who all listed Mantis gigantea Gmélin, 1789 as a junior synonym of Mantis gigas Drury, 1773. Drury's ♀ type-specimen was believed lost, but a ♀ in NHMUK with the data “Diapherodes gigas Drury, Saint Vincent” matches perfectly with the illustration provided by Drury (1773, plate 50) in every aspect, hence is here interpreted as the holotype of Mantis gigas Drury. Haplopus grayi Kaup, 1871 (most certainly from Guadeloupe, → see comments on D. angulata above) and Diapherodes gigantea dominicae Rehn & Hebard, 1938 from Dominica were both erroneously interpreted as synonyms of Diapherodes gigas (Drury, 1773) in the unpublished PhD-Thesis of Moxey (1971: 99). In fact, H. grayi is a synonym of D. angulata (Fabricius, 1793) and D. gigantea dominicae is here shown to represent a separate and valid species (→ see Diapherodes dominicae Rehn & Hebard, 1938 n. comb.). Redtenbacher (1908: 434, pl. 20: 1) provided brief descriptions of both sexes and illustrated a ♀ from Saint Vincent in NHMW. Examination of numerous specimens throughout European and American museum collections has proven D. gigantea is restricted to the two southern Lesser Antillean islands Grenada and Saint Vincent. No specimens have so far been recorded from the intervening Grenadines and a specimen from Trinidad in NHMUK is most certainly mislabelled. All records from Guadeloupe and Dominica were based on misidentified material of either D. angulata (Fabricius, 1793) or D. dominicae (Rehn & Hebard, 1938). The specimens recorded from Saint Lucia by Langlois, Lelong & Dorel (2006: 42) and Langlois & Lelong (2010: 69) are here described as a separate subspecies (D. gigantea saintluciae n. ssp.), since these differ by a good number of morphological characters from specimens from Grenada and Saint Vincent (→ see below). In Grenada this species is known to feed on cinnamon (Cinnamomum spp., Lauraceae) and guava (Psidium guayava, Myrtaceae). Hand-written labels underneath the specimens from Saint Vincent and Grenada in NHMW state these insects to be principally found on high trees and to be commonly observed on cashew-trees (Anacardium occidentale, Anacardiaceaee). Hence, this might be another natural host-plant of D. gigantea gigantea. Based on a record of a museum specimen, Moxey (1972: 10) furthermore mentioned Ixora sp. (Rubiaceae) to be a possible host-plant. Rabaey (2004) provided brief information on the distribution in Grenada, alternative food plants and breeding of D. gigantea. In captivity in Europe nymphs and adults accept eucalyptus (Eucalyptus spp., Myrtaceae), bay tree (Laurus nobilis, Lauraceae), guava (Psidium guajava, Myrtaceae), salal (Gaultheria shallon, Ericaceae), different oaks (Quercus spp., Fagaceae) and bramble (Rubus fruticosus, Rosaceae) as alternative food plants. Adults are fairly robust and long-lived insects, ♀♀ reaching ages of up to one year. Mating is frequent but mainly takes place during the night. On average ♀♀ lay two eggs per day, which are simply flicked away by a rapid movement of the abdomen. The strong and ventrally spinose hind-legs of ♀♀ are frequently used for active defense, whereas ♂♂ flash their large orange wings. This species has been included on the Phasmid Study Group (PSG) culture-list as culture No. 260. Photos of an interesting captive reared gynandromorph have been contributed by Mieke Duytschaever (Belgium). This specimen is a mainly ♂ but has several ♀ characteristics on the right. While there is a fully developed ala on the left, the right ala is only represented as a small scale-like rudiment. All right legs and most parts of the right body surface a bright geen as in ♀♀, the remaining portions of the body being brown as typical for ♂♂ of this species (Figs. 406–407). The genitalia are mostly ♂ with a typical anal segment, but the poculum is considerably deformed (Fig. 408). Distribution (Fig. 381): Saint Vincent [Volcano Soufriére [NHMUK]; Sharp's Valley [NHMUK]) and Grenada (St. Georges [USNM]; Vendome Estate [NHMUK]; St. Andrew, Birch Grove [FSCA]). Number of specimens examined: 111



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TABLE 13. Measurements of Diapherodes gigantea gigantea (Gmélin, 1789) [mm] ♀ Saint Vincent [NHMW]

♀♀ Grenada

♂♂ Grenada

♂♂ Saint Vincent [NHMUK]


195.0

153.0–197.0

-

-

Body

181.0

140.0–183.0

92.0–116.3

116.5–126.0

Pronotum

8.4

5.8–8.2

4.3–4.8

4.1–5.0

Mesonotum

34.0

22.7–32.0

13.0–16.4

18.1–21.1

Metanotum

13.0

9.9–11.0

7.8–8.5

8.0–8.9

Median segment

13.6

11.8–13.5

8.7–9.5

10.0–11.0

Tegmina

15.3

11.6–14.8

14.6–17.0

15.8–18.0

Alae

8.5

5.7–6.3

50.0–59.0

62.0–69.5

Profemora

33.0

23.5–31.5

16.0–19.9

21.8–23.6

Mesofemora

30.0

19.6–26.7

14.2–17.0

18.1–20.0

Metafemora

43.0

27.0–40.0

18.9–23.0

27.0–28.0

Protibiae

34.5

20.4–28.6

15.9–19.5

21.6–21.7

Mesotibiae

28.5

12.8–25.4

12.6–15.1

19.3–21.5

Metatibiae

42.2

24.2–36.2

17.0–19.8

23.6–24.0

Antennae

> 85.0

59.0–98.0

54.0–68.0

> 66.0

Diapherodes gigantea saintluciae n. ssp. (Figs. 136–143, 367, 381) Diapherodes gigantea, Langlois, Lelong & Dorel, 2006: 42, figs. 19 a–g (♂, ♀, nymph), photos 5–6 (♀) & 27–29 (egg, REMphotos). Langlois & Lelong, 2010: 60 (in part—only records from Saint Lucia). HT, ♀: Saint Lucia, Mount Rochar 1100 ft., Nr. Dennery, 22.30 local time, 1994, on guava on side of track, AR + DP James [NHMUK]. PT, 2 ♀♀: Santa Lucia 89–99 [NHMUK]. PT, ♂: Saint Lucia, Prospect, 9.VI.1936, H.E. Box, Pres. by Com. Inst. Ent. B. M. 1952-302 [NHMUK]. PT, ♂: Saint Lucia, Castries, 9.XI.1934, H.E. Box, Pres. by Com. Inst. Ent. B. M. 1952-302 [NHMUK]. PT, ♂: La Perle Estate, Saint Lucia, W. Indies, J. Frew, B. M. 1935-270, Diapherodes sp. not in B. M. det. B. Uvarov 1935 [NHMUK]. PT, ♂: Saint Lucia: Union Agr. School, 25-V-1987, R.E. Woodruff [FSCA]. PT, 1 ♂, 1 ♀: Sainte-Lucie, Barre de l’Isle Trail, alt. 275–305 m, 13°55’34’’N 60°57’28’’E, 02.XI.2003 [MNHN, coll. ASPER; No’s SLU03-90 & 94]. PT, 3 ♀♀: Sainte-Lucie, Barre de l’Isle Trail, alt. 275–305 m, 13°55’34’’N 60°57’28’’E, 02.XI.2003 [coll. ASPER; No’s SLU03-91 to 93]. PT, ♀: Sainte-Lucie, Mount Beaujolais nord, alt. 38–274 m, 13°54’30’’N 60°55’45’’E, 22.X.2003 [coll. ASPER, No. SLU0318]. PT, 2 ♂♂: Sainte-Lucie, Banse 1 km nord, alt. 183–229 m, 13°47’09’’N 60°58’53’’E, 21.X.2003 [coll. ASPER, No’s SLU0398 to 99]. PT, ♂: Sainte-Lucie, Forestière, alt. 185–305 m, 13°57’29’’N 60°57’46’’E, 29.X.2003 [coll. ASPER, No. SLU03-074].

Diagnosis: Similar to the nominate form from Saint Vincent and Grenada but, apart from being geographically isolated, easily distinguished by the considerably smaller size, relatively shorter mesothorax and more numerous spiniform tubercles and spines of the mesonotum of both sexes. Females differ by: the more broadened mesothorax; longitudinal row of small spines near the lateral margins of the mesonotum (Figs. 136–137); more


HENNEMANN ET AL.

numerous granules and tubercles of the meso- and metasternum; lack of the distinct lamellate expansions of the mid and hind legs, as well as the almost smooth dorsal carinae of the meso- and metafemora and corresponding tibiae. ♂♂ differ from those of the nominate form by: the black mesothoracic armature; somewhat longer alae, which usually exceed the posterior margin of abdominal tergum VI (Figs. 138–139); broader posteromedian excavation of the anal segment (Fig. 143) and broader vomer, which has the terminal hook somewhat less acuminate and relatively shorter (Fig. 359). One of the ♂ paratypes (specimen from Prospect in NHMUK) has a rounded posterolateral lobe on abdominal tergum VII and the meso- and metafemora bear a sub-apical lobe dorsally, both of which features are also not known to occur in D. gigantea gigantea. The characterizations provided below emphasize on the distinctive features. Etymology: Neuter. The name “saintluciae” refers to the type-locality Saint Lucia, one of the Lesser Antillean islands, to which this new subspecies is endemic. Characterization: ♀ (Figs. 136–137). Of moderate size for the genus (body length including subgenital plate 118.5–132.0 mm), and fairly broad (body width at abdominal tergum III 11.0–14.0 mm). General colouration bright yellowish green to apple green, ventral body segments mid green with a slight whitish wash on abdominal sternites. All larger spines and spiniform tubercles of the thorax dull yellow with black tips. Meso- and metapleurae white ventrally with the longitudinal row of spines dark red to dull reddish brown. Antennae pale red, the scapus and pedicellus with a brownish hue. Eyes of moderate size, slightly elongate-oval; their length contained about 2.5x in that of cheek. Pronotum with a prominent median pair of blunt spines in the anterior half; posterior half armed with one or two pairs of blunt tubercles medially and a small, spiniform tubercle at each posterolateral angle. Mesothorax rather short and decidedly, gradually widened towards the posterior; about 1.4x longer than head and pronotum combined; mesonotum with posterior margin more than 2x wider than anterior margin (Figs. 136–137). Dorsal surface of mesonotum armed with two distinct pairs of spines close to anterior margin and a variable number of spines and spiniform tubercles in the remainder portion except the posterior 1/3; usually with two considerably enlarged spines medially. Along lateral margins with a longitudinal row of 6–12 moderately sized spines, which gradually decrease in size towards the posterior of segment (Figs. 136–137). Spines of the meso- and metapleurae acute and of moderate size, the largest spine less than 2x as long as its basal width. Meso- and metasternum unevenly tuberculose. Abdominal tergum VII ± deflexed with lateral margins gently rounded (Fig. 141). Posterior margin of anal segment broadly rounded, epiproct very small and fully hidden under anal segment (Fig. 141). Subgenital plate long and projecting over apex of abdomen by more than the combined length of tergites IX and X Figs. 140–141). Posterodorsal carina of profemora, as well as all carinae of the mid and hind legs finely but acutely granulose. Medioventral carina of meso- and metafemora faint and with 6–10 stout but distinct spines, which decrease in size towards the base of femur. Posteroventral carina with one, anteroventral carina with two moderately distinct, pointed sub-apical spines. ♂ (Fig. 138–139). Medium-sized for the genus (body length 83.0–89.0 mm). General colouration creamish mid brown, the ventral body segments except the prosternum whitish. Spines of the pro- and mesonotum black with olive bases, all smaller tubercles and granules of the thorax black. Antennae ochre. Anterior margin of tegmina and anterior margin in basal portion of alae pale yellow. If present, sub-apical dorsal lobes of the mesoand metafemora very dark brown. Vertex armed with two moderately decided ochre tubercles, back of vertex smooth. Pronotum with a well developed pair of blunt spines in front of the transverse median depression, and a pair of considerably smaller, spine-like tubercles in the posterior portion. Mesothorax about 2x longer than head and pronotum combined. Mesonotum dorsally armed with 8–12 distinct but blunt paired spines and usually a cluster of four paired spines close to anterior margin; posterior portion of dorsal surface irregularly covered with small, acute granules; a variable number of blunt tubercles present along lateral margins. Mesopleurae with a longitudinal row of 8–10 rather distinct tubercles, which gradually decrease in size towards the posterior. Alae reaching to or projecting over posterior margin of abdominal tergum VI. Anal region of alae pale transparent pink. Abdominal tergum VII either parallel-sided or with a rather distinct, rounded lobe posterolaterally. Anal segment with a broad but shallow posteromedian excavation (Fig. 143). Vomer roundly triangular, hardly longer than wide and with a fairly short and broad, acuminate apical hook (Fig. 367). Poculum with a distinct median incision at posterior margin (Fig. 367). Posterodorsal carina of meso- and metafemora sometimes with a ± prominent, roundly triangular lobe sub-apically; anterodorsal carina of meso- and metatibiae occasionally expanded and rounded subbasally and sub-apically. Spines on the medioventral carina of the meso- and metafemora fairly small.



81

FIGURES 136–139. Diapherodes gigantea saintluciae n.ssp.. 136. ♀ PT: Saint Lucia, Barre de l’Isle Trail [coll. ASPER]; 137. ♀ PT: Saint Lucia, Barre de l’Isle Trail [coll. ASPER]; 138. ♂ PT: Saint Lucia, Barre de l’Isle Trail [coll. ASPER]; 139. ♂ PT: Saint Lucia, Union Agricultural School [FSCA].

Variability: Both sexes show some variability in the size and number of the mesothoracic spines. The ♂ from Prospect in NHMUK is characteristic for having the lateral margins of abdominal tergum VII with a rounded posterior lobe, having a roundly triangular sub-apical lobe on the posterodorsal carina of the meso- and metafemora and having the anterodorsal carina of the meso- and metatibiae roundly to triangularly expanded sub-basally and sub-apically.


HENNEMANN ET AL.

FIGURES 140–143. Diapherodes gigantea saintluciae n.ssp.. 140. ♀ apex of abdomen (lateral view) [coll. ASPER]; 141. ♀ apex of abdomen (dorsal view) [coll. ASPER]; 142. ♂ apex of abdomen (lateral view) [coll. ASPER]; 143. ♂ apex of abdomen (dorsal view) [coll. ASPER].

Comments: Langlois, Lelong & Dorel (2006: 42) and Langlois & Lelong (2010: 69) reported D. giagntea saintluciae n. ssp. is not particularly common on Saint Lucia with merely nine specimens found at four of the thirteen prospected localities. However, these authors stated it was comparatively more abundant than other Diapherodes-species are on neighbouring islands. Diapherodes gigantea saintluciae n. ssp. occurs in various biotopes like montane and lowland hygrophilic forests as well as cultivated zones in xero-mesophilic environments and is usually found in vegetation 0.5–4 metres off the ground. In the wild mimosa (Mimosa camporum, Mimosaceae), guava (Psidium guajava, Myrtaceae) and mapou-baril (Sterculia caribea, Sterculiaceae) seem to represent natural food plants. Captive rearing in Europe has shown bay tree (Laurus nobilis, Lauraceae) and bramble (Rubus fruticosus, Rosaceae) to be accepted as alternative food plants by both nymphs and adults. Distribution (Fig. 381): Saint Lucia (Mount Rochar, nr. Dennery 1100 ft. [NHMUK]; Prospect [NHMUK]; Castries [NHMUK]; La Perle Estate [NHMUK]; Trail above Banse 183–229 m [ASPER]; Northern Mount Beaujolais 38–274 m [ASPER]; Forestière [ASPER]; Barre de l’Isle Trail 274–305 m [MNHN, ASPER] & Union Agricultural School [FSCA]). Endemic. Number of specimens examined: 16 TABLE 14. Measurements of Diapherodes gigantea saintluciae n. ssp. [mm] ♀, HT [NHMUK]

♂, PT [FSCA]

♀♀, PT* [MNHN & coll. ASPER]

♂♂, PT* [MNHN & coll. ASPER]


124.5

-

123.0–132.5

-

Body

119.0

82.0

-

86.0–89.0

Pronotum

5.5

3.0

5.5–6.0

3.0–3.5

Mesonotum

19.6

12.0

21.0–25.0

12.5–14.0

Metanotum

8.2

-

8.0–10.0**

13.0–14.0**

Median segment

10.0

-

Tegmina

9.8

11.2

10.5–13.0

10.5–12.0

Alae

5.0

46.5

-

48.0–50.0

Profemora

19.6

14.5

21.0–23.0

14.5–15.0

Mesofemora

18.3

12.9

20.0–22.5

13.5–14.0 ......continued on the next page



83

TABLE 14. (Continued) ♀, HT [NHMUK]

♂, PT [FSCA]

♀♀, PT* [MNHN & coll. ASPER]

♂♂, PT* [MNHN & coll. ASPER]

Metafemora

25.5

16.7

27.5–32.0

17.0–18.0

Protibiae

19.3

14.1

-

-

Mesotibiae

15.5

10.5

-

-

Metatibiae

23.5

16.3

-

-

Antennae

> 53.0

43.5

57.0–59.0

44.0–49.0

* according to Langlois & Lelong, 2010: 60 ** including median segment

Diapherodes martinicensis Lelong & Langlois, 2005 (Figs. 144–156, 349, 369, 381, 394–395) Diapherodes sp. Langlois, Lelong, Polidori & Dorel, 2000: 46, figs. 19a–g (♀, ♂, nymph) & photos 10–12 (egg). Diapherodes martinicensis Lelong & Langlois, 2005: 264, figs. 19–23 (♀, ♂, nymph), 40–42 (egg). HT, ♀: Martinique, trace Duclos, 10/XI/98, ref. No. MAR98-036 [MNHN]; AT, ♂: Martinique, Morne-Rouge, 15.X.1960, Diapherodes dominicae—Rehn et Hebard [coll. Pére Pinchon, Hôtel de Région Martinique]; PT, ♀: Martinique, Balata, 18.III.1963 [coll. Pére Pinchon, Hôtel de Région Martinique]; PT, ♀: Martinique [coll. H. Griffon, Martinique]. Further material [16 ♂♂, 18 ♀♀, 2 nymphs, eggs]: MARTINIQUE: 7 ♂♂, 6 ♀♀, 1 ♀ (n5), 1 ♀ (n4), eggs: ex Zucht F. Hennemann 2014–2015, Herkunft: Martinique, leg. C.B. Dupré 2011 [coll. FH, No’s 0750-1 to 15 & E]; 9 ♂♂, 12 ♀♀, eggs: ex Zucht. O. Conle, Herkunft: Martinique [coll. OC].

Diagnosis: This is the smallest species of the gigantea species-group and apart from the small size (body length of ♀♀ < 10 cm) well distinguished from all other representatives by: the short mesothorax, which is strongly gradually broadened towards the posterior, trapezoidal in dorsal aspect and swollen dorsally with a prominent central pair of spines in ♀♀ (Fig. 144); completely unarmed tibiae of both sexes; having only 37 antennomeres in ♀♀ (at least 50 in all other species); relatively shorter legs with the hind legs of ♀♀ not projecting over abdominal tergum VI, as well as the long alae of ♂♂ which at least reach to the posterior margin of abdominal tergum VII (Figs. 146–147). Females are also characterised by the bright red and black intersegmental membrane between abdominal sternites I and II (Fig. 154). Eggs are distinctive by their very strongly sculptured capsule surface and comparatively smaller micropylar plate, which covers less than half of the capsule length (Figs. 155–156). Description: ♀ (Figs. 144–145, 394): Very small for the genus (body length including subgenital plate 92.0–105.5 mm), with a conspicuously shortened and strongly broadened mesothorax and comparatively short, dorsally unarmed legs. General colouration plain bright apple-green, the ventral body surface and basal half of subgenital plate dull green, sometimes with a slight whitish hue. Lateral margins of meso- and metasternum white and a longitudinal white stripe along the ventral surface of the lateral margins of the abdominal tergites. Points of the two large spines on the vertex, as well as all larger tubercles of the pro- and mesonotum and the row of spines on the meso- and metapleurae dark pink; anterior half of mesopleurae dark pink as well. Bases of dorsal thoracal tubercles and spines yellow. Antennae dark pink except for the two basal segments which are blackish with pale outer margins, and a few pale annulations in the apical 1/3. Tegmina and alae of same colouration as body. Tarsi and subgenital plate becoming pale brown towards the apex. Eyes creamish mid brown. Head: Vertex strongly convex and armed with two very prominent, blunt spines; the dextral one larger (Fig. 148). A few minute granules are present posterior of these and a small pair of rounded granules on the frons. Eyes slightly elongate-oval, convex and of moderate size; their length contained about 2.6x in that of cheeks. Between the bases of the antennae with a fairly decided, slightly curved and transverse depression. Antennae reaching about half way along median segment; with 37 segments. Scapus dorsoventrally flattened, almost 3x longer than wide and gently narrowed towards the base. Pedicellus cylindrical and about 2/5 the length of scapus.


HENNEMANN ET AL.

FIGURES 144–147. Diapherodes martinicensis Lelong & Langlois, 2005. 144. ♀ HT: Martinique, trace Duclos (dorsal view) [MNHN]; 145. ♀: captive reared from Martinique [coll. FH, No. 0750-1]; 146. ♂: captive reared from Martinique [coll. FH, No. 0750-8]; 147. ♂: captive reared from Martinique [coll. FH, No. 0750-5].



85

FIGURES 148–156. Diapherodes martinicensis Lelong & Langlois, 2005. 148. ♀ head and pronotum [coll. FH, No. 0750-3]; 149. ♂ head and pronotum [coll. FH, No. 0750-5]; 150. ♀: apex of abdomen (lateral view) [coll. FH, No. 0750-3]; 151. ♀: apex of abdomen (dorsal view) [coll. FH, No. 0750-3]; 152. ♂: apex of abdomen (lateral view) [coll. FH, No. 0750-5]; 153. ♂: apex of abdomen (dorsal view) [coll. FH, No. 0750-5]; 154. Metasternum and abdominal sterna I–II of ♀ showing the colourful inter-segmental membrane [coll. FH, No. 0750-3]; 155. Egg (dorsal view) [coll. FH, No. 0750-E]; 156. Egg (lateral view) [coll. FH, No. 0750-E].

Thorax: Pronotum shorter and about as broad as the head; roughly 1.3x wider than long. In front of the very prominent transverse median depression armed with a pair of very large, blunt spines; posterior half set with a few minute granules (Fig. 148). Mesothorax short, only about 1.3x longer than head and pronotum combined, decidedly constricted anteriorly and strongly broadened towards the posterior; posterior margin almost 4x broader than anterior margin (Fig. 144). Mesonotum sparsely set with minute granules which slightly increase in size towards the anterior, otherwise dorsally armed with two pairs of prominent, blunt spines close to anterior margin and a further pair of similarly sized spines just before the middle (Fig. 148). Metanotum unarmed, about half the length


HENNEMANN ET AL.

of mesonotum and slightly wider than long. Spines of the meso- and metapleurae more acute than dorsal thoracal armature; mesopleurae with 12–13, metapleurae with 10–11 spines. Tegmina rather small and reaching about ¾ the way along metanotum (length 5.5–7.0 mm), posterior margin slightly angulate; not overlapping. Alae very small and projecting underneath tegmina by about 2.2–3.0 mm. Abdomen: Median segment slightly longer than metanotum and roughly trapezoidal with the anterior margin narrowed. Segments II–VI slightly increasing in length and gradually narrowing, II about 2x wider than long, VI 1.25x longer than wide. VII narrowest and about as long as VI, almost parallel-sided or with the posterolateral portions somewhat deflexed and forming a small, rounded lobe (Fig. 151). Tergites II–VII all with a faint, slightly curved longitudinal carina laterally. Intersegmental membrane between abdominal sternites I and II bright red and black (Fig. 154). Sternites II–VII with a blunt lateral longitudinal carina. Praeopercular organ formed by an elongate, keel-shaped swelling near posterior margin of sternum VII (Fig. 349). Tergum VIII slightly shorter and narrower than VII and widened towards the posterior; decidedly more convex than previous. IX less than half the length of VIII, slightly narrowed towards the posterior and very faintly keeled. Anal segment gently narrowed, posterior margin roundly angular and dorsally with a faint, longitudinal median keel. Epiproct small, broad and very slightly projecting underneath anal segment (Fig. 151). Subgenital plate long and considerably narrowed towards a slender and pointed tip; projecting over apex of abdomen by almost the combined length of tergites VIII–X (Figs. 150–151). Legs: All short, moderately broad and entirely unarmed, except for 4–5 minute spines on the two outer ventral carinae of the meso- and metafemora; these decrease in size towards the base. Profemora slightly longer than mesothorax and metatibiae slightly projecting over posterior margin of abdominal tergum V. Tarsi about half the length of corresponding tibia. Basitarsi indistinctly longer than second tarsomere. ♂ (Figs. 146–147, 395). Small (body length 64.0–81.5 mm) and moderately slender for the genus, with very long alae (43.5 mm) and a comparatively short, dorsally sparsely spinose mesothorax. General colouration of head and body pale yellowish brown; ventral body segments whitish. Posterior half of poculum pale cream. Tegmina and costal region of alae creamish mid brown, the tegmina with a broad, longitudinal white stripe along the anterior margin and the central hump marked with dull pink. Anal region of alae pale translucent orange. Femora pale green basally and gradually becoming mid brown towards the apex (with a slight dull pink hue ventrally); tibiae entirely pale green with only the apex slightly brownish. Tarsi pale brown but becoming decidedly darker towards the apex. Cephalad pair of spines and all larger spines and tubercles of the thorax tipped with black. Antennae pale brown with the basal portion slightly reddish and the two basal segments very dark brown; a few dark brown annulations close to the apex. Eyes dark reddish brown. Head: Generally as in ♀♀. Vertex strongly convex and smooth except for two prominent, blunt but slender cephalad spines (Fig. 149). Eyes large, very slightly oval and projecting hemispherically; their length contained less than 2x in that of cheeks. Antennae reaching about half way along abdominal tergum III, otherwise as in ♀♀. Thorax: Pronotum roughly quadrate and about as long and broad as head. Transverse median depression very prominent, almost straight and reaching lateral margins of segment. Anterior half with a pair of prominent blunt spines and posterior half with a pair of low tubercles (Fig. 149). Mesothorax short, just 1.5x longer than head and pronotum combined; mesonotum about 4x longer than wide. Dorsal surface with a few minute granules and six distinct, blunt paired spines; two pairs close to anterior margin (Fig. 149) and a further pair slightly before the mid of segment. Lateral margins with a longitudinal row a small greenish tubercles. Meso- and metapleurae each with a longitudinal row of minute granules. Meso- and metasternum sparingly granulose. Tegmina reaching to posterior margin of metanotum. Alae at least reaching to posterior margin of abdominal tergum VII (Figs. 152–153). Abdomen: Tergum VII with a distinct, rounded posterolateral lobe, which projects by about ¼ of the body width (Fig. 153). Tergites VIII–X very indistinctly broader than previous segments, VIII about ¾ the length of VII and IX 2/3 the length of VIII. Anal segment gently narrowed towards the apex with the posterior margin broadly rounded (Fig. 153). Cerci slender and narrowing towards a blunt apex. Vomer with a fairly small base and a very long, gradually narrowing and acutely pointed apical hook (Fig. 369). Poculum strongly convex and conical basally, just slightly projecting over posterior margin of tergum IX (Fig. Fig. 152); the posterior margin very broadly rounded. Legs: All of moderate length, profemora almost as long as pro- and mesonotum combined, metatibiae reaching about halfway along abdominal tergum VI. Posterodorsal carina of meso- and metafemora very slightly elevated and rounded sub-basally. Both outer ventral carinae of meso- and metafemora with 2–3 sub-apical spines; a few



87

more minute spines are present in the median section of the posteroventral carina of the mesofemora. Remaining carinae unarmed. Tarsi elongate, about half the length of corresponding tibia. Basitarsi about 1.5x longer than second tarsomere. Egg (Figs. 155–156): Rather small (capsule length 4.3–4.8 mm), capsule 1.7x longer than wide. Polar area decidedly impressed if seen in lateral aspect. Capsule surface irregularly covered with prominently raised structures, these crater-like and with several distinct circular pits dorsally. Micropylar plate small, its length contained about 2.5x in that of capsule. Central portion of micropylar plate with two irregularly raised ridges, which begin at the micropylar cup and converge towards the anterior, where they terminate about 0.4 mm off the anterior end of plate. Micropylar cup distinct. Operculum in the central portion with numerous irregular granules and tubercles; centre slightly lowered. General colour pale to dark brown. Outer margin of micropylar plate and micropylar cup mid brown. Measurements [mm]: Length 4.3–4.8, length (including operculum) 4.5–5.0 width 2.5–2.7, height 2.7–2.9, length of micropylar plate 1.7–1.8. Comments: Lelong & Langlois (2005) described Diapherodes martinicensis from three ♀♀ and one ♂ and provided REM-photographs of the eggs. According to the small number of known specimens, D. martinicensis appears to be rare. Lelong & Langlois (2005: 264) reported this species to be apparently uncommon and to feed on guava (Psidium guajava, Myrtaceae) and Coccoloba sp. (Polygonaceae) in the wild. Only one ♀ was found during extensive collections conducted by these authors in several localities throughout Martinique. Only two ♀♀ were collected by Christian Bouladou Dupré (Martinique) on several occasions during 2011. The specimens were kept alive in captivity in Martinique and accepted Acacia muricata (Fabaceae), Tapura latifolia (Dichapetalaceae), Pimenta racemosa (Myrtaceae), Ocotea coriacea (Lauraceae) and Avocado (Persea americana, Lauraceae), hence at least some of these are likely to represent native host plants. Eggs were sent to Bruno Kneubühler (Switzerland), who first successfully reared D. martinicensis in Europe. In captivity hawthorn (Pyracantha coccinea, Rosaceae), salal (Gaultheria shallon, Ericaceae), oak (Quercus robur, Fagaceae) and dog rose (Rosa canina, Rosaceae) are accepted as alternative food plants. Females produce an average of three eggs per day, which are simply flicked away by an abrupt movement of the abdomen. Eggs take some 4–5 months to hatch at temperatures of 20–25°C and nymphs reach maturity in 3–4 months. If disturbed, adult ♂♂ will readily flash their wings and are capable of short active flights. Females will either drop to the ground or pinch with their hind legs, to expose the bright red and black coloured intersegmental membrane between the abdominal sternites I and II (Fig. 154). Distribution (Fig. 381): Martinique (Trace Duclos [MNHN]; Balata [coll. H. Griffon] & Morne-Rouge [MNHN]). Endemic. Number of specimens examined: 40 TABLE 15. Measurements of Diapherodes martinicensis Lelong & Langlois, 2005 [mm] ♀♀, HT & PT*

♂, PT [coll. Pinchon]*

♀♀ [coll. FH]

♂♂ [coll. FH]

92.0–97.0

-

92.0–105.5

-

-

64.0

87.0–97.3

68.0–81.5

4.0

2.5

3.9–4.1

2.2–2.7

Mesonotum

12.5–13.0

9.5

13.0–14.0

8.9–10.6

Metanotum

6.0–8.0

12.0**

5.3–5.8

5.3–5.9

Median segment

6.5–8.5

-

7.0–7.5

6.0–7.0

7.0

11.5

5.0–6.2

9.0–11.0

-

43.5

2.2–3.0

40.5–50.0

Profemora

13.0–15.0

11.5

11.8–14.7

10.5–12.6

Mesofemora

12.0–14.5

11.0

10.6–12.5

9.4–11.0

Body (incl. sg. pl.) Body Pronotum

Tegmina Alae



HENNEMANN ET AL.

TABLE 15. (Continued) ♀♀, HT & PT*

♂, PT [coll. Pinchon]*

♀♀ [coll. FH]

♂♂ [coll. FH]

17.5–21.0

15.0

15.8–19.0

12.5–15.8

Protibiae

-

-

10.3–12.6

9.9–11.5

Mesotibiae

-

-

9.8–11.3

8.1–9.9

Metatibiae

-

-

14.6–16.0

12.0–14.0

Antennae

29.5–30.0

37.5

31.5–36.0

36.5–44.0

Metafemora

* according to Lelong & Langlois (2005: 266) ** including median segment

5.4.2. The jamaicensis species-group Characteristics: The three known members of the jamaicensis species-group are restricted to the two Greater Antillean islands of Jamaica and Puerto Rico (Fig. 375). They are of moderate size for the genus (body lengths: ♂♂ 60.0–94.0 mm, ♀♀ including subgenital plate 84.0–164.5 mm) with ♀♀ ranging from rather slender to moderately robust. Body surface of ♂♂ sub-glabrous to glabrous. Colouration of both sexes usually bright green (♀♀ more rarely brown or with brown markings). Pronotum without anterior spines; at best sparsely tuberculate. Mesonotum of ♂♂ entirely unarmed or with 2–6 paired anterior spines. Meso- and metasternum smooth (♂♂), mesosternum may have a few minute granules in ♀♀. Meso- and metapleurae of ♂♂ unarmed, of ♀♀ with a marginal row of granules or spiniform tubercles. Alae of ♀♀ ± as long as tegmina; anal region of both sexes plain pink to purple. Abdomen of ♀♀ broadened sub-basally with segment III broadest. Tergites IV–VII of ♀♀ ± decidedly multicarinate; VII ± strongly laterally expanded or with a triangular posterolateral lobe. Epiproct of ♀♀ large, triangular to shield-shaped and distinctly projecting over anal segment. All carinae of legs unarmed, except for spines on medioventral carina and sub-apical spines on the two outer ventral carinae of the meso- and metafemora. Dorsal carinae of meso- and metafemora may bear a ± tooth or lobe sub-apically. Eggs small to moderately sized (capsule length < 5.0 mm), micropylar plate small and usually < 1/3 the length of capsule. For a detailed comparison with the gigantea species-group see Table 10.

Diapherodes achalus (Rehn, 1904) n. comb. (Figs. 157–166, 350, 370, 382–383) Aplopus achalus Rehn, 1904: 68. HT, ♂: Adjuntas P.R. 12 Apr. 00, C.W. Richmond Collector, Cat. no. Aplopus achalus Rehn Type, Type No. 7344 U.S.N.M. [USNM, No. 7344]. Wetmore, 1916: 58. Wolcott, 1923: 23. Wolcott, 1936: 35. Wolcott, 1948: 50. Wolcott, 1951: 50. Diapherodes achalus, Van de Bussche et al., 1988: 422. Haplopus achalus, Kirby, 1904a: 363. Otte & Brock, 2005: 150. Diapherodes achalus, Moxey, 1972: 90 (in litt.). Diapherodes longiscapha Redtenbacher, 1908: 435. LT (by present designation), ♀: Porto-Rico, Antilles, Mr. H. de Saussure, Diapherodes longiscapha REDT. [MHNG]; PLT, ♀: 141, Portorico Coll. Krug, Diapherodes longiscapha Br. Brunner det, Type [MNHU] n. syn. Wolcott, 1923: 23. Wolcott, 1936: 35. Wolcott, 1948: 50. Wolcott, 1951: 50. Zompro & Brock, 2003: 16.



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Otte & Brock, 2005: 121. Diapherodes gigantea, Wolcott, 1923: 23. (Misidentification—not Mantis gigantea Gmélin, 1789) Wolcott, 1936: 35. Wolcott, 1948: 50. Wolcott, 1951: 50. Diapherodes krugii, Wolcott, 1923: 23. (Nomen nudum) Wolcott, 1936: 35. Further material [16 ♂♂, 15 ♀♀, 1 nymph, 1 egg]: PUERTO RICO: 1 ♂: Puerto Rico: Parador, Casa Grande, nr. Utuado, 16-III-1990, at light, R.E. & K.E. Wooruff [FSCA]; 1 ♀: Barranguitas, P.R., June 1959, coll. L.F. Martorell [USNM]; 1 ♀ (in Ryker mount): From Porto Rico Exp. Stat. […] Nov. 19.1913; Diapherodes longiscapha Redt. ♀ A.n.c. [USNM]; 1 ♀: Road S.G. to Mariacao, 18K6M side road, Beating from Tree, Feb. 13,'61, A.M. Nadler; P.R. Road from Maricao-Sabana Grande km 18.6, Beating tree, Feb. 13,'61, A.M. Nadler; Diapherodes longiscapha Redt. ♀, det. A.B. Gurney 1961 [USNM]; 1 ♂: P.R. ACC.NO. 69–50, Cayey, P.R., Sept. 26,1950; G.N. Wolcott & L.F. Martorell Collectors; USMNH; w0150; Diapherodes achalus (Rehn) det. CF Moxey 1972 [USNM]; 1 ♂: Utuado, P.R., Nov. 1948, Coll: E. Boneta; w0152; USMNH; Diapherodes achalus (Rehn) det. CF Moxey 1972 [USNM]; 1 ♂: Caguas, P.R., Jan. 8.1912, in bird stomach. Biol. Survey.; Aplopus achalus Rehn A.n.c [USNM]; 2 ♂♂: Barranquitas, P.R., Nov. 20 1948, Coll. S. Rodriguez [USNM]; 1 ♂: Mayaguez, P.R., 1/14/47, C. Feyre R. [USNM]; 1 ♂: Mayaguez, P.R., 1-19-1930, Coll. E. Rosa [USNM]; 1 ♂: Ponce, P.R., X-20 1947 [USNM]; 1 ♀: Juyuya PR, VIII-1454, H F Winters, PR-1278; Toro Negro Ar, Carribean Nat. Forest, Lot 45-19676 [ANSP]; 1 ♀ (valde defectum): Yauco, P.R., June 1945, Coll: [USNM]; 2 ♀♀: Rio Piedras, P.R., IV-2-1944, Coll: R. Casas [USNM]; 1 ♀: Barranquitas, P.R., Oct. 10 1948, Coll. S. Rodriguez [USNM]; 1 ♀: Maricao, P.R., Nov. 20 1990, Coll: Iv. Nazario [USNM]; 1 ♀: Aibonito, P.R., 8-9-33, R.G. Oakley; on leaf Inga vera; San José No. 4630; wO143 [ANSP]; 1 ♀: El Yunque Mountains, Luquillo, P.R., June 1935; cdl. L. F. Martore, Host: on weeds, Acc#: 29–35; wO144 [ANSP]; 1 ♀: Jayuya, PR, Aug. 23–45, H F Winters, PR No. 472; on Cinchona ledgeriana; Lot No. 56-1971; wO147 [ANSP]; 1 ♂: Jayuya, P.R., Aug. 23–45, H F Winters, PR No. 470; on Cinchona ledgeriana; wO151 [ANSP]; 1 ♂: Mayaguez, PR; wO149 [ANSP]; 1 ♀ (penultimate instar): Yauco, P.R., II.1934, Cdi J. Matini; wO146 [ANSP]; 1 ♀: Mayaguez, P. R,. 3. Oct. 1938; J. A. Ramos Collector [ANSP]; 1 ♂: Mayaguez, P. R., 21 Oct. 1937, Coll.: R. del Moral [ANSP]; 1 ♂: Aibonito, P. R., II-16-1935, Coll. J. Martinez [ANSP]; 1 ♀, 1 egg (ex ovipositor): Porto Rico, Mayaguez, 5.7.1906, C. Gaggo leg.; PHA 12, Zoologisches Museum Hamburg [ZMUH]; 1 ♀: Mayaguez, Porto Rico, C. Gaggo leg. 5.VII.1906, ded. 3.IX.1906; PHA 11, Zoologisches Museum Hamburg [ZMUH]; 1 ♂: Puerto Rico, Maricao, 13.10.2009, leg. Mel J. Rivera [coll. OC]. NO DATA: 1 ♂: F. Sein Collector; Aplopus jamaicensis (Drury) A.n.c. [USNM].

Diagnosis: Well characterized and distinguished from the two other species of the jamaicensis species-group by the elongate head, just gently rounded and almost unarmed vertex (Figs. 159–160), entirely unarmed mesothorax (Figs. 157–158) and having only two apical spines on the medioventral carina of the meso- and metafemora in both sexes. ♂♂ furthermore differ by the bold brown longitudinal median stripe on the tegmina (Fig. 158) and brown cheeks (Fig. 160). The characteristic eggs are unique within the entire genus by having a hollow polar area and a prominent conical swelling on the operculum (Figs. 165–166). Description: The colouration is described from several photos of live specimens taken at Maricao in Puerto Rico by Mel J. Rivera (Figs. 374–375). ♀ (Fig. 157, 382). Fairly small (body length including subgenital plate 92.0–116.0 mm) and rather massive member of the jamaicensis species-group with a moderately widened mesothorax and abdominal segment VII. Colour bright apple green, the meso- and metapleurae ochre with a white longitudinal stripe along lower margin. Ventral body surface mostly whitish. Tegmina and costal region of alae bright green with the anterior margin darker in colour. Anal region of alae dull pink to violet. Antennae dark ochre dorsally and blackish brown ventrally. Eyes pale ochre. Head: Elongate, ovate and 1.3x longer than wide, the vertex very gently convex and smooth except for a very faint pair of cephalad tubercles; the sinistral very faint (Fig. 159). Eyes rather small, circular and their length contained a little more than 3x in that of cheeks. Antennae broken in all specimens at hand but at least reaching to metanotum. Scapus dorsoventrally compressed, constricted basally with the outer lateral margin gently rounded, 2.3x longer than wide. Pedicellus oval in cross-section about 2/3 the length of scapus. Thorax: Pronotum considerably shorter and narrower than head, hardly longer than wide and gently narrowing towards the posterior. Transverse median depression deep but very short and curved. Surface unarmed except for a pair of small tubercles just in front and behind the median sulcus (Fig. 159). Mesothorax about 1.6x longer than


HENNEMANN ET AL.

head and pronotum combined, constricted in anterior portion and then gradually widened towards the posterior; posterior margin more than 3x wider than anterior margin. Mesonotum very slightly tectiform longitudinally and widened towards the middle, then roughly parallel-sided; surface smooth except for a few scattered small tubercles, in particular in the anterior portion. Mesopleurae with a marginal row of spiniform tubercles or short spines; metapleurae armed with a row of about eight short spines. Pro-, meso- and metasternum smooth; the mesosternum occasionally with a small pair of granules posteriorly. Tegmina oval, reaching to posterior margin of metanotum and with the central protuberance moderate. Alae roughly equal in length to tegmina and reaching about half way along median segment.

FIGURES 157–160. Diapherodes achalus (Rehn, 1904). 157. ♀: Puerto Rico, Mayaguez [ANSP]; 158. ♂: Puerto Rico, Parador Casa Grande [FSCA]; 159. ♀ head and pronotum: Puerto Rico, Mayaguez [ANSP]; 160. ♂ head and pronotum Puerto Rico, Parador Casa Grande [FSCA].

Abdomen: Median segment slightly longer than wide and very gently widening towards the posterior; smooth. Abdomen strongly swollen sub-basally with segment II widening, IV narrowing and III widest. II–IV transverse and III almost 1.5 wider than long. V–VIII longer than wide, VII moderately expanded and with the lateral margins rounded (Fig. 162). Tergites II–IV smooth, V–VII with four roughly parallel, longitudinal carinae which are most decided on VII (Fig. 162). Sternites II–VII smooth except for a fine carina along lateral margins. Praeopercular organ formed by a brown tubercle close to posterior margin of sternum VII (Fig. 350). Tergites VIII–X narrower than previous segments and roughly of uniform width. VIII about ¾ the length of VII and very slightly constricted medially, about 1.2x longer than wide; IX indistinctly longer than wide. Anal segment about equal in length to IX, with a slight longitudinal median carina in basal portion and flattened towards the posterior; posterior margin



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rounded and with a very small and shallow median indentation (Fig. 162). Epiproct small and semi-circular with a keel dorsally (Fig. 162). Cerci very small, round in cross-section and tapered towards a pointed apex. Subgenital plate elongate, lanceolate, longitudinally keeled and variable in length; projecting over apex of abdomen by more than the combined length of tergites IX–X (Figs. 161–162). Legs: Profemora slightly shorter, mesofemora about ¾ the length of mesothorax and metafemora reaching about half way along abdominal segment IV. Medioventral carina of all femora with two rather small spines in apical portion. Posteroventral carina of meso- and metafemora with one, the anteroventral carina with two distinct, black-tipped sub-apical spines. Basitarsi no longer than following two tarsomeres combined. ♂ (Figs. 158, 383). Small (body length 60.0–74.0 mm) and fairly robust for the jamaicensis-group with a very characteristic colouration, body surface glabrous, with very long alae (36.8–42.0 mm). General colour bright green, washed with brown, the lateral surfaces of the head, entire pronotum, mesonotum, mesopleurae, posterior portion of metapleurae and bases of all femora ochraceous. Abdominal tergites V–X green laterally, yellow dorsally and V–IX with a mid to dark brown longitudinal median stripe (Fig. 321). Ventral surface of abdomen with a whitish wash. Head bright green with the cheeks ochraceous brown and two bold, washed yellow markings between the eyes (Fig. 160). Anterior margin of tegmina and basal portion of the same region of the alae white, tegmina with a further broad brown longitudinal stripe, the innermost portion green with darker green veins and basal portion yellow (Fig. 158); costal region of alae plain green with dark green longitudinal veins, the anal region very pale and translucent pink. Eyes mid to dark brown. Antennae greyish ochre. Ventral sub-apical spines of the meso- and metafemora black. Head: Ovate and only about 1.2x wider than long, otherwise generally as in ♀♀; cephalad tubercles very faint (Fig. 160). Eyes of moderate size and projecting hemispherically, their length contained less than 2x in that of cheeks. Antennae projecting over posterior margin of abdominal segment II and moderately robust; with 36–37 antennomeres. Scapus oval in cross-section, roundly rectangular with the inner margin concave and about 1.3x longer than wide. Pedicellus cylindrical and about 2/3 the length of scapus. Thorax: Pronotum slightly shorter and narrower than the head, with the anterior portion slightly broader than posterior portion; median transverse depression distinct, gently curved and almost reaching lateral margin of segment. Surface smooth or with a very few minute granules (Fig. 160). Mesothorax 1.8x longer than head and pronotum combined; mesonotum at best with a few very obsolete granules anteriorly. Mesopleurae smooth, metapleurae with a few small granules. Meso- and metasternum smooth. Tegmina oval and with a rather longitudinal and acute central protuberance, roughly reaching to posterior margin of metanotum. Alae ± reaching to posterior margin of abdominal tergum VII (Fig. 158). Abdomen: Median segment gently narrowed towards the posterior. Segments II–VI slightly decreasing in length and width, II 2.7x and VI only about 2x longer than wide. elongate, cylindrical, parallel-sided and gradually decreasing in length. II 3.5x, VII 2.5x longer than wide. VII about equal in length to VI and very slightly widened posteriorly. Sternites II–VII smooth. Tergum VIII about 3/5 the length of VII, widened towards the posterior and slightly tectate. IX slightly shorter than VIII and very slightly narrowed towards the posterior. Anal segment about as long as IX with two longitudinal carinae dorsally, slightly constricted towards the posterior and the posterior margin sub-truncate with a shallow median emargination (Fig. 164). Vomer roughly triangular with a moderately long terminal hook (Fig. 370). Cerci about 2/3 the length of anal segment, rather slender, cylindrical and very slightly in-curving with the apex blunt. Poculum moderately convex, cymbiform and longitudinally carinate, the posterior margin rounded and with a narrow median emargination; very slightly projecting over posterior margin of tergum IX (Fig. 163). Legs: Shape and armature generally as in ♀♀. Egg (Figs. 165–166): Medium sized (capsule length 5.0 mm), capsule about 2x longer than wide and slightly higher than wide. Capsule surface srtrongly and irregularly sculptured with several raised swelling and an irregular swollen keel indicated, that surrounds the micropylar plate and posteriorly runs towards lateral surfaces of capsule. Another distinctly raised, blunt keel below micropylar plate, which almost reaches to the polar area. The polar area hollow, impressed and surrounded by an irregularly toothed rim. Anterior margin with an irregularly crown-like rim of tooth like and anteriorly directed swellings. Micropylar plate small, heart-shaped and its length contained about 3.5x in that of capsule. Central portion impressed but otherwise strongly swollen. Operculum sub-circular with a large, irregularly sculptured conical swelling in centre. General colour pale to mid ochraceous to reddish brown.


HENNEMANN ET AL.

FIGURES 161–166. Diapherodes achalus (Rehn, 1904). 161. ♀ apex of abdomen of synonym HT of D. longiscapha (lateral view) [MHNG]; 162. ♀ apex of abdomen of HT of synonym D. longiscapha (dorsal view) [MHNG]; 163. ♂ apex of ♂ abdomen (lateral view) [FSCA]; 164. ♂ apex of abdomen (dorsal view) [FSCA]; 165. Egg: Puerto Rico, Mayaguez (dorsal view) [ZMUH]; 166. Egg: Puerto Rico, Mayaguez (lateral view) [ZMUH].

Measurements [mm]: Length 5.0, length (including operculum) 6.2 width 2.5, heigth 2.9, length of micropylar plate 1.3. Comments: Moxey (1972: 10, in litt.) stated D. achalus to have been taken on Guarea ramiflora (Meliaceae) and Inga vera (Mimosaceae) which might both be part of its natural diet. Body lengths cited by Moxey (1972: 92, in litt.) are 61.0–74.0 mm for ♂♂ and 92.0–114.0 mm including the subgenital plate for ♀♀. Distribution: Puerto Rico: Barranquitas [USNM]; Mayaquez [ANSP, ZMUH]; Maricao Forest [ANSP, coll. OC]; Yauco [ANSP]; between Maricao and Sabana Grande km 18.6 [USNM]; Cayey [USNM]; Adjuntas [ANSP, USNM]; Utuado [USNM]; Jayuya [ANSP]; Aibonito [ANSP]; Ponce [USNM]; Rio Piedras [USNM]; Caguas [USNM]; Naguabo, El Yunque Quadrangle, Naguabo [Van den Bussche et al., 1988: 423] & Luquillo Experimental Forest, El Verde [ANSP]. Endemic. Number of specimens examined: 35 TABL 16. Measurements of Diapherodes achalus (Rehn, 1904) n. comb. [mm] ♂, HT achalus [USNM]

♂ [FSCA]

♀, LT longiscapha [MHNG]

♀, PLT longiscapha [MNHU]

-

-

116.0

> 104.0

Body

61.5

60.0

101.5

97.4

Pronotum

2.8

2.6

4.2

5.7

Mesonotum

9.0

8.7

17.9

16.2





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TABL 16. (Continued) ♂, HT achalus [USNM]

♂ [FSCA]

♀, LT longiscapha [MHNG]

♀, PLT longiscapha [MNHU]

Metanotum

4.9

9.5*

3.0

7.5

Median segment

5.1

-

9.8

6.0

Tegmina

10.0

8.8

8.2

8.8

Alae

39.0

36.8

9.0

8.9

Profemora

11.8

10.0

16.2

14.8

Mesofemora

11.0

8.9

14.1

13.4

Metafemora

14.8

12.2

14.8

19.0

Protibiae

10.0

9.1

14.0

12.9

Mesotibiae

9.3

7.4

12.5

10.7

Metatibiae

10.7

10.2

18.4

16.7

Antennae

ca. 47.0

> 27.0

30.0

> 35.0

* metanotum + median segment

Diapherodes jamaicensis (Drury, 1773) n. comb. (Figs. 167–182, 351, 371, 375, 388–391) Mantis jamaicensis Drury, 1773: 88, pl. 49: 1 (♂). HT, ♂: Jamaica [not traced—believed lost]. Fabricius, 1787: 227. Olivier, 1792: 634. Fabricius, 1793: 15. Aplopus jamaicensis, Rehn, 1903: 132. Brock, 1998b: 33. Otte & Brock, 2003: 302. Cyphocrana jamaicensis, St. Fargeau & Audinet-Serville, 1825: 445. Diapherodes jamaicensis, Moxey, 1972: 102 (in litt.). Haplopus jamaicensis, Westwood, 1859: 86. Kirby, 1904a: 363. Redtenbacher, 1908: 430. Otte & Brock, 2005: 151. Phasma jamaicensis, Fabricius, 1798: 188. Phasma jamaicense, St. Fargeau & Audinet-Serville, 1825: 101 Phasma (Platycrana) jamaicensis, Westwood, 1837: 99, pl. 44: 1 (♂). Platycrana jamaicensis, Gray, 1835: 38. Mantis bispinosa Fabricius, 1775: 274. LT, ♂: “America. Mus. Dom. Hunter” [specimen with wings open] [HMUG]; PLT, 2 ♂♂: same data as LT [HMUG]. [Synonymised by Redtenbacher, 1908: 430] Aplopus bispinosus Brock, 1998c: 33. [Lectotype designation] Gmélin, 1789: 2054. Olivier, 1792: 633. Mantis 2 spinosa, Fabricius, 1787: 227. Fabricius, 1793: 15. Haplopus bispinosus, Westwood, 1859: 87. Kirby, 1904a: 430. Otte & Brock, 2003: 303. Phasma bispinosa, Fabricius, 1798: 188. Latreille, 1806: 87. Phasma bispinosum, Audinet-Serville, 1831: 58. Gray, 1835: 24. Haplopus christopheri Westwood, 1859: 84, pl. 33: 4, 4a (♀). LT (by present designation), ♀: St. Christopheri, W. Indies, E. Shepard; Diapherodes christopheri Westwood, p. 84, pl. 33 f4; Type—Westwood, Haplopus christopheri, Cat Phasm.


HENNEMANN ET AL.

1859, p. 84, pl. 33 f. 4; Type Orth: 614 1/2, Diapherodes christopheri Westw., Hope Dept. Oxford [OXUM, No. 614]; PLT, ♀: St. Christopher, 1858, W. Indies, E. Shepard; Diapherodes christopheri Westwood, p. 84, pl. 33 f4; Cat Phasm. 1859, p. 84, pl. 33 f. 4; Type Orth: 614 2/2, Diapherodes christopheri Westw., Hope Dept. Oxford [OXUM, No. 614] n. syn. Diapherodes christopheri, Kirby, 1904a: 462. Redtenbacher, 1908: 434. Moxey, 1972: 94 (in litt.). Otte & Brock, 2005: 120. Diapherodes glabricollis Gray, 1835: 33. HT, ♀: Type; Haplopus glabricollis G.R. Gray,—?, glabricollis Gray 108, Diapherodes glabricollis Gray, HT; BMNH(E) #844944 [NHMUK]. n. syn. Brock et al., (in press) Haplopus glabricollis, Westwood, 1859: 88. Kirby, 1904a: 364. Redtenbacher, 1908: 430. Otte & Brock, 2005: 151. Diapherodes glabricollis, Burmeister, 1838: 575. Haplopus murinus Redtenbacher, 1908: 429. LT, ♀ (by present designation): Coll. Br. v. W., Jamaika, Burr; Det. Br. v. W., Haplopus murinus; 25.068; Stony Hill, Jamaica, July 29/93, Hwm; Lectotypus, Haplopus murinus Redt., P.D. Brock [NHMW, No. 834]; PLT, ♀: Cuming, Jan., Jamaica, Burr; det. Br. v. W. Haplopus murinus, Paralectotypus, Haplopus murinus Redt., P.D. Brock [NHMW, No. 834]. n. syn. Aplopus murinus, Brock, 1998a: 44. Haplopus murinus, Otte & Brock, 2005: 152. Diapherodes murinus, Moxey, 1972: 103 (in litt.). Diapherodes pulverulentus Gray, 1835: 34. HT, ♀: Diapherodes pulverulentus Gray; pulverulentus Gray III Haplopus pulverulentus G. R. Gray;—? Diapherodes pulverulentus Gray HT; BMNH(E) #844949 [NHMUK]. n. syn. Brock et al., (in press) Haplopus pulverulentus, Kirby, 1904a: 364. Redtenbacher, 1908: 431. Otte & Brock, 2005: 152. Further material [70 ♂♂, 47 ♀♀, 7 nymphs, eggs]: JAMAICA: 1 ♀ [brown form]: Kingston, Jamaica, C.R. Orcutt.; USMNH; w0214; Diapherodes jamaicensis (Drury) det. C.F. Moxey 1972 [USNM]; 1 ♀ [brown form]: Jam., Quick Step. nr. Aberdee, VIII.8.'41, W.G. Lynn; USDMNH; w0211; Diapherodes jamaicensis (Drury) det. C.F. Moxey 1972 [USNM]; 1 ♀: Jamaica, Recd. 1928, C.R. Orcutt; USMNH; w0210; Diapherodes jamaicensis (Drury) det. C.F. Moxey 1972 [USNM]; 1 ♂: Fellowship Rio Grande, July 27,1962, Farr, O. & R. Flint, Jamaica, Portland Parish ; USMNH; Diapherodes jamaicensis (Drury) det. C.F. Moxey 1972 [USNM]; 1 ♂: Bath, Jamaica, July 1902; Diapherodes jamaicensis (Drury) det. C.F. Moxey 1972 [USNM]; 3 ♂♂, 1 ♀: Coll. Br. v. W., Jamaica; det. Br. v. W., Haplopus jamaicensis [NHMW, No. 833]; 1 ♀: Coll. Br. v. W., Jamaica; det. Br. v. W., Haplopus jamaicensis; Haplopus jamaicensis [18]93 [NHMW, No. 833]; 1 ♂, 1 ♀: Cumming, Jan. 847; Mus. Caes. Vind., Jamaika, Cumming leg.; det. Redtenb. Haplopus jamaicensis [NHMW, No. 833]; 1 ♂: Coll. Br. v. W., Jamaica; det. Br. v. W., Haplopus jamaicensis; 22.767; Haplopus jamaicensis Drury, Jamaica [NHMW, No. 833]; 1 ♂: Jamaica; 45 110 [NHMUK]; 1 ♂: Jamaica; 46 84; Haplopus jamaicensis (Drury), Jamaica [NHMUK]; 1 ♂: Jamaica; 46 84 [NHMUK]; 1 ♂, 1 ♀: Jamaica; 61 63 [NHMUK]; 1 ♂: Jamaica, Mandeville, L. J. Bertram, B. M. 1923-574 [NHMUK]; 1 nymph (n1): Jamaica 96-220 [NHMUK]; 1 ♀: Jamaica, Dr. Leon, Brit. Mus. 1922-162, Haplopus murinus Redt., Det. B. Uvarov ♀ [NHMUK]; 1 ♂: 54 76; Brasil [NHMUK]; 1 ♀: 34J, Jamaica, Haplopus an foem. Jamaicensis ?, Jamaica [NHMUK]; 1 ♀: Jamaica: Blaudeville, Da Careret College, 26.V.1969, K. Stanton, B. M. 1969-506, Diapherodes jamaicensis (Drury) det. Moxey 1972 [NHMUK]; 1 ♂: Jamaica: Portland, Parrish, Millbank, 4-VIII-1985, C.B. & H.V. Weems Jr., G.B. Edwards [FSCA]; 1 ♀: M. Delfosse, Jamaique, ♀ Elevage 02/04/2005 [MNHN]; 1 ♀: M. Delfosse, Jamaique, ♀ Elevage 10/04/ 2005 [MNHN]; 1 ♂: Haplopus jamaicensis T. Jourdan ♂ Elevage 06/2010 [MNHN]; 5 ♂♂, 6 ♀♀, 3 ♀♀ (nymphs), 1 ♂ (nymph), eggs: ex Zucht: F. Hennemann, urspr. Jamaika, 2000–2002 [coll. FH, No’s 0428-1 to 11, 14 to 16, E & MP]; Right tegmen and ala of ♂: ex Zucht: F. Hennemann, urspr. Jamaika, 2000–2002 [coll. FH, No’s 0428-12 and 13]; 5 ♀♀, 7 ♂♂, 2 ♀♀ (penultimate instar), 1 ♀ (n5), eggs: ex Zucht F. Hennemann 2008/09, Herkunft: Jamaica, Blue Mts., leg. T. & P. James (PSG No. 214) [coll. FH, No’s 0428-17 to 30 & 33]; 2 ♀♀, 6 ♂♂: ex Zucht F. Hennemann 2009, Herkunft: Jamaica, Blue Mts., leg. T. & P. James (PSG No. 214) [coll. FH, No’s 0428-31, 32 & 34-39]; 2 ♀♀, 2 ♂♂: Jamaica, F. Klages; Coll. W. Holland; A.N.S.P., Ex Carn. Mus., Bruner Cln. [ANSP]; 1 ♂: Jamaica; wO222 [ANSP]; 1 ♂: Jasmaica; wO217 [ANSP]; 30 ♂♂, 18 ♀♀, eggs: ex Zucht O. Conle, Herkunft: Jamaika [coll. OC]. NO / ERRONEOUS DATA: 2 ♀♀: Africa, J. E. Teede, B. W. 1928-311 [NHMUK]; 1 ♂: bispinosum Fab., 106 [NHMUK]; 1 ♂: Brazil, 54, 76; Haplopus bispinosus (Fabr.) Brazil [NHMUK]; 1 ♂: Brazil, 54, 70 [NHMUK].



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FIGURES 167–170. Diapherodes jamaicensis (Drury, 1773). 167. ♀: captive reared from Jamaica (typical green specimen with a short subgenital plate) [coll. FH, No. 0428-20]; 168. ♀: captive reared from Jamaica (green specimen with a brownish wash and a very short subgenital plate) [coll. FH, No. 0428-19]; 169. ♀: captive reared from Jamaica (typical green specimen with a long subgenital plate and strongly developed leg-armature) [coll. FH, No. 0428-17]; 170. ♀, PLT of synonym Haplopus murinus Redtenbacher, 1908: Jamaica (brown colour-form) [NHMW, No. 834].

Diagnosis: This species differs from the other two representatives of the jamaicensis species-group by the slender body of ♀♀ in particular, which makes it resemble members of the genus Haplopus Burmeister, 1838, and short alae of ♂♂, which merely reach to abdominal segment VI (VII in the other two species). From the Puerto Rican D. achalus (Rehn, 1904) ♀♀ furthermore differ by the spines and tubercles of the mesonotum and prominent cephalad horns, while they are at once distinguished from the second Jamaican species D. laevicollis Redtenbacher, 1908 by the just gently widened mesothorax, shorter alae, which merely reach half way along the median segment, and not broadly rounded abdominal tergum VII. Males differ from those of D. achalus by the plain green tegmina and anterior spines of the mesonotum and from D. laevicollis by the dorsally brown scapus and pedicellus, more sender body and much shorter terminal hook of the vomer (Fig. 371). Description: ♀ (Figs. 167–170, 390–391). Medium-sized to large (body length including subgenital plate 104.0–164.5 mm) and moderately slender for the genus with a rather slender and almost parallel-sided abdomen and just slightly widened mesothorax. Colouration very variable and occurring in a green and brown colour-form, which are here described separately i) Green form (Figs. 167, 169, 390): Colour bright apple green, the ventral


HENNEMANN ET AL.

body surface dull green. Lateral margins of mesonotum ochre, meso- and metapleurae ochre with a white longitudinal stripe along lower margin; membranes along lateral margins of the abdomen whitish. Tegmina and costal region of alae bright green, the anterior margin of tegmina brown and interiorly bordered by an obscure yellow longitudinal stripe; basal portion of alae with a bold brown marking. Anal region of alae plain pink (Figs. 167–168). Antennae pale brown dorsally, black ventrally and annulated in apical 1/3. Eyes reddish brown. Tips of the anterior mesonotal spines black. ii) Brown form (Fig. 170): General colour greyish mid brown and often all over furnished with pale grey or whitish mottling and speckles; occasionally washed with some green. Tegmina and costal region of alae mid to purplish dark brown with the venations of a paler colour; anal region of alae dark red to purple with all veins marked by dark brown. Culture stock in Europe occasionally also produces intermediate forms, which are mostly green with the body to a variable degree furnished with well-defined bold chestnut-brown markings or with a brownish wash (Figs. 168, 391). Head: Indistinctly longer than wide, strongly globose with cheeks parallel-sided. Vertex convex, covered with a variable number of differently sized tubercles and in centre armed with two ± prominent, pointed or blunt, conical horns; the dextral one carinate and the sinistral one considerably smaller (Figs. 173–174). Near posterior margin usually with a transverse row of four spiniform tubercles, the exterior ones sometimes enlarged. Often two further small granules are present on the cheeks. Eyes of moderate size, sub-circular and their length contained about 2x in that of cheeks. Antennae slightly projecting over posterior margin of metanotum; consisting of about 45 antennomeres. Scapus dorsoventrally compressed, roundly rectangular and about 1.5x longer than wide. Pedicellus cylindrical, sub-spherical and less than half the length of scapus. Third antennomere slightly shorter than pedicellus. Thorax: Pronotum slightly shorter and narrower than head, 1.3x longer than wide, gently constricted medially and with the anterior margin slightly widened. Transverse median depression distinct, slightly curved and almost reaching lateral margins of segment. Surface close to posterior margin with a pair of minute spiniform tubercles and in centre often with four small tubercles, which roughly form a square (Figs. 173–174). Mesothorax about 1.6x longer than head and pronotum combined, constricted anteriorly and gently widened towards the posterior. Mesonotum with a very faint longitudinal median carina and covered with a variable number of tubercles, which become more distinct towards the lateral margins and less numerous towards the posterior of segment; anteriorly with 1–3 pairs of ± prominent, forward directed spines (Figs. 173–174). Mesopleurae with a marginal row of 8–14 spiniform tubercles; metapleurae smooth or with a few small tubercles. Meso- and metasternum sparsely granulose. Tegmina oval, scale-like, with a very shallow central hump and very slightly projecting over posterior margin of metanotum. Alae slightly shorter than tegmina and reaching about half way along median segment. Abdomen: Median segment longer than metanotum, 1.3x longer than wide and very gently widening towards the posterior; smooth. Segments II–VII almost parallel-sided and of uniform length, about 1.3x longer than wide; III and IV slightly broader than remaining. Tergites II and III occasionally with two small spine-like projections at posterior margin, II–VI otherwise smooth except for a few scattered granules. VI with two and VII with four fine longitudinal carinae. Lateral margins of VII slightly expanded posteriorly and forming a narrow, triangular lobe or tooth. Sternites II–VII smooth except for two very faint and short sub-parallel carinae close to posterior margin. Praeopercular organ formed by an elongate brown median hump close to posterior margin of sternum VII (Fig. 351). Tergites VIII–X roughly equal in width and considerably narrower than previous segments. VIII about 2/3 the length of VII and slightly constricted medially, IX quadrate and about 2/3 the length of VIII. Anal segment about as long as IX, with a slight longitudinal median carina, the posterior margin rounded and with a very small median indentation (Fig. 178). Epiproct slightly projecting over anal segment, shield shaped and roughly semi-circular with a keel dorsally (Fig. 178). Cerci very small, round in cross-section and conical with the apex pointed. Subgenital plate elongate and variable in length, lanceolate, longitudinally keeled and narrowed towards a rather acute apex; projecting over apex of abdomen by more than the length of anal segment (Figs. 177–178). Legs: All rather short and stocky. Profemora and mesofemora shorter than mesothorax, metafemora slightly projecting over posterior margin of abdominal segment III. Medioventral carina of profemora with 2–3 spines in apical portion, mesofemora with 4–6 and metafemora with 5–8 spines. Dorsal carinae of meso- and metafemora each with a ± prominent triangular lobe sub-apically (less distinct or sometimes lacking on metafemora), the posteroventral carina with one and the anteroventral carina with two pointed sub-apical spines. Anterodorsal carina of meso- and metatibiae occasionally with a ± distinct rounded to roundly triangular sub-basal and sub-apical expansion. Medioventral carina raised and rounded sub-basally and armed with a few small teeth in apical portion. Basitarsi about as long as following two tarsomeres combined. REVISION OF HAPLOPODINI


97

FIGURES 171–176. Diapherodes jamaicensis (Drury, 1773). 171. ♂: captive reared from Jamaica [coll. FH, No. 0428-26]; 172. ♂: captive reared from Jamaica [coll. FH, No. 0428-35]; 173. ♀ head and pronotum: captive reared from Jamaica (specimen with large and irregularly tuberculose horns) [coll. FH, No. 0428-31]; 174. ♀ head and pronotum: captive reared from Jamaica (specimen with moderately sized, conical horns) [coll. FH, No. 0428-32]; 175. ♂ head and pronotum: Jamaica: Portland, Parrish (specimen two large anterior mesonotal spines) [FSCA]; 176. ♂ head and pronotum: captive reared from Jamaica (specimen with a cluster of six anterior mesonotal spines) [coll. FH, No. 0428-37].


HENNEMANN ET AL.

FIGURES 177–182. Diapherodes jamaicensis (Drury, 1773). 147. ♀ apex of abdomen (lateral view); 148. ♀ apex of abdomen (dorsal view); 149. ♂ apex of abdomen (lateral view); 150. ♂ apex of abdomen (dorsal view); 151. Egg: captive reared from Jamaica (dorsal view) [coll. FH, No. 0428-E]; 152. Egg: captive reared from Jamaica (lateral view) [coll. FH, No. 0428-E].

♂ (Figs. 171–172, 388–389). Of moderate size (body length 63.0–94.0 mm) and rather slender for the jamaicensis species-group with well developed alae (32.0–53.5 mm); body surface sub-glabrous. General colouration bright green, the mesonotum with a slight yellowish wash. Tegmina green with a bold longitudinal white stripe along anterior margin. Costal region of alae slightly translucent green with a white longitudinal stripe running along the basal 1/3 of the anterior margin. Anal region translucent pink to orange (Fig. 171). Eyes reddish brown. Antennae reddish mid brown with the three or four basal antennomeres black ventrally. Pedicellus usually darker brown than all other antennomeres. Anterior spines of the mesonotum often black. Bases of all femora and tibiae ochre. Tarsi greenish brown. Head: Similar in shape to ♀♀ but vertex much less convex and smooth except for two rounded unequal cephalad humps and two blunt tubercles close to the posterior margin; sometimes also with a few minute granules (Figs. 175–176). Eyes very prominent, sub-circular and projecting hemispherically; their length contained hardly more than once in length of cheeks. Antennae projecting over posterior margin of abdominal segment II and moderately robust; consisting of 57–60 antennomeres. Scapus oval in cross-section, roundly rectangular and about 1.3x longer than wide. Pedicellus cylindrical and about 2/3 the length of scapus. Thorax: Pronotum somewhat shorter and narrower than the head, slightly trapezoidal and gradually narrowed towards the posterior; the posterior margin gently rounded. Median transverse depression distinct, gently curved and almost reaching lateral margin of segment. Surface smooth (Figs. 175–176). Mesothorax about 2x longer than head and pronotum combined, mesonotum mostly with 1–4 pairs of prominent black, forward directed anterior spines (Figs. 175–176); rarely lacking. Posterior 2/3 of mesonotum and mesosternum sometimes with a few minute granules. Meso- and metapleurae as well as metasternum smooth. Tegmina elongate, oval, reaching 1/3 along median segment and with a blunt central protuberance, which is slightly displaced towards the anterior. Alae reaching to abdominal tergum VI. Abdomen: Median segment gently narrowed towards the posterior. Segments II–VII parallel-sided and gradually decreasing in length; II 3.5x, VII 2.5x longer than wide. Sternites II–VII smooth. Tergum VIII ¾ the length of VII, widened towards the posterior and slightly tectate. IX ¾ the length of VIII, about as long as wide and very slightly narrowed towards the posterior. Anal segment about as long as IX with a slight longitudinal median carina, the posterior portion narrowed and rounded on both sides of the triangular median emargination (Fig. 180). Vomer with the base rather broad and roundly triangular, the terminal hook slightly up-curving and almost as long as basal portion (Fig. 371). Cerci almost as long as anal segment, slender, narrowed towards a blunt apex and very



99

slightly in-curving (Fig. 371). Poculum convex, cymbiform, longitudinally carinate and with the posterior margin distinctly indented and bi-lobate medially (Fig. 179). Legs: Generally as in ♀♀ but relatively longer, more slender and lacking the dorsal armature of the meso- and metafemora and tibiae. Profemora roughly equal, mesofemora slightly shorter than mesothorax, metafemora reaching about half way along abdominal segment IV. Basitarsi a little longer than following two tarsomeres combined. Nymphs: Newly hatched nymphs are bright yellowish green with brown antennae. Later instars and ♀ nymphs in particular tend to be various shades of brown rather than green like the adults. From 3rd instar on the lateral margins of tergum VII are conspicuously elevated and form a rounded lobe. The sub-apical dorsal teeth of the meso- and metafemora and dorsal elevations of the tibiae of ♀ nymphs are more prominent than in adults. ♂ nymphs may have small granules on the mesonotum. ♀ nymphs are very similar in general appearance to those of certain Haplopus-species. Variability: While ♂♂ merely show variability in the size and number of anterior spines of the mesonotum (Figs. 175–176), ♀♀ exhibit considerable variability in several aspects. Features generally showing variation are the number and size of the anterior spines of the mesonotum and armature of the head (Figs. 173–174), length of the subgenital plate and degree of the anterior armature of the mid and hind legs. Despite these morphological features ♀♀ occur in two distinct colour-forms described in detail above, which has caused the description of several synonyms. Specimens of the more common green form (Figs. 167, 169, 390) are averaging smaller than specimens of the more rarely encountered brown form (Fig. 170). Culture stock in Europe occasionally also produces green intermediate forms which are to a variable degree prettily furnished with rather geometrical chestnut-brown markings and possess a brownish hue all over the body (Fig. 168, 391). Egg (Figs. 181–182): The following description is based on a large number of eggs in the first author's collection (coll. FH) laid by captive reared specimens. Rather small, capsule 1.8x longer than wide and with dorsal surface more convex than ventral surface; polararea decidedly impressed if seen in lateral aspect. Capsule surface irregularly covered with raised ridge-like structures, tubercles and granules; a rather blunt longitudinal keel running from the micropylar plate to the polararea. Micropylar plate small, roughly heart-shaped and its length contained about 2.6x in that of capsule. Central portion of micropylar plate granulose, micropylar cup distinct and placed in posteromedian gap of plate. Operculum with a rim of irregularly raised tubercles; centre slightly lowered. General colour pale to mid brown, the outer margin of the micropylar plate and micropylar cup dark brown. Measurements [mm]: Length 3.5–3.8, length (including operculum) 3.7–4.0, width 2.0–2.2, height 2.1–2.3, length of micropylar plate 1.3–1.4. Comments: Drury's illustration of the ♂ HT does not show the anterior spines on the mesonotum mostly present and a very variable feature in D. jamaicensis. There are several old specimens in the collection NHMUK but none of these could be confirmed as Drury's holotype (personal communication with J. Marshall and P.D. Brock). Hence, the designation of a neotype may become necessary, but this deserves further evaluation and research in other collections, which are likely to contain specimens examined by Drury. Diapherodes glabricollis Gray, 1835 was described from the ♀ and is the opposite sex of Drury's species (n. syn.). Examination of the holotype of Diapherodes christopheri Westwood, 1859 in OXUM leaves no doubt it is conspecific with D. jamaicensis (Drury) (n. syn.). The type-locality “St. Christopher” [= St. Kitts] is however very doubtful and most certainly erroneous. Former authors distinguished D. christopheri from D. jamaicensis by the smooth metapleurae, which however is a variable feature in ♀♀ of jamaicensis. Both, D. pulverulentus Gray, 1835 and H. murinus Redtenbacher, 1908 are ♀♀ of the brown form of D. jamaicensis, hence also synonyms (n. syn.). D. jamaicensis is being successfully reared in captivity in Europe from stock imported by Tony and Pat James (England) in the late 1990's and has subsequently been included on the Phasmid Study Group culture-list as culture No. 214 “Haplopus jamaicensis”. It readily accepts oak (Quercus robur & Q. petraea, Fagaceae), eucalyptus (Eucalytus gunnii, Myrtaceae), bramble (Rubus fruticosus, Rosaceae), raspberry (Rubus idaeus, Rosaceae), rose (Rosa spp., Rosaceae), salal (Gaultheria shallon, Ericaceae) and guava (Psidium guajava, Myrtaceae) as alternative food plants and does well in moderately humid but well ventilated conditions. If adults are disturbed, both ♀♀ and ♂♂ will spread their pink hindwings. Males are very quick moving and capable of active flight. Mating usually takes place at night. Distribution: Jamaica: Batia [ANSP]; Bath [USNM]; Liguanea Plain [MCZC; Moxey, 1972: 104, in litt.];


HENNEMANN ET AL.

Quick Step nr. Aberdeen [USNM]; St. James, Montego Bay [NHMJ; Moxey, 1972: 104, in litt.]; Snug Harbour [ANSP]; Fountain Hill [NHMJ; Moxey, 1972: 104, in litt.]; Trelawny, Westwood [NHMJ; Moxey, 1972: 104, in litt.]; Manchester, Mandeville [NHMJ; Moxey, 1972: 104, in litt.]; Christiana [AMNH; Moxey, 1972: 104, in litt.]; St. Mary, Castleton Gardens [MCZC, Moxey, 1972: 104, in litt.]; St. Catherine, Boston Town Postal Agency [NHMJ; Moxey, 1972: 104, in litt.]; St. Andrew; Hermitage Reservoir [NHMJ; Moxey, 1972: 104, in litt.]; Blaudeville [NHMUK]; Cinchona [AMNH; Moxey, 1972: 104, in litt.]; Kingston [USNM]; Portland, Millbank [FSCA] & Portland Parish, Fellowship Rio Grande [USNM, NHMJ, Moxey, 1972: 104, in litt.]. Endemic. Number of specimens examined: 134 TABLE 17. Measurements of the green form of Diapherodes jamaicensis (Drury, 1773) n. comb. [mm] ♀, HT glabricollis [NHMUK]

♀ [NHMW]

♀♀ captive reared [coll. FH]

♂♂ captive reared [coll. FH]

♂ [NHMW]


139.0

137.5

104.0–120.0

-

-

Body

124.0

126.0

100.5–117.5

63.0–75.0

76.5

Pronotum

5.6

5.2

4.6–4.8

2.5–2.8

3.2

Mesonotum

23.8

23.0

17.8–20.5

9.8–11.9

13.8

Metanotum

7.9

8.1

6.1–7.0

4.8–5.2

-

Median segment

8.0

8.3

6.2–7.1

6.6–7.0

-

Tegmina

10.9

9.6

7.1–8.1

7.0–9.1

7.4

Alae

10.0

8.9

6.5–7.0

32.0–39.0

38.0

Profemora

19.8

18.9

14.2–16.0

10.4–12.0

12.3

Mesofemora

17.5

17.3

11.6–13.1

9.4–10.3

-

Metafemora

24.6

24.2

16.0–18.0

12.0–13.8

14.5

Protibiae

19.2

18.2

12.8–14.8

10.0–11.9

11.5

Mesotibiae

18.2

14.1

10.2–11.0

7.9–8.6

-

Metatibiae

24.0

20.2

14.5–25.5

10.3–12.1

12.3

Antennae

-

> 34.0

34.0–43.0

37.0–45.0

> 29.5

TABLE 18. Measurements of the brown form of Diapherodes jamaicensis (Drury, 1773) n. comb. [mm] ♀ HT pulverulentus [NHMUK]

♀ LT murinus [NHMW]

♀ PLT murinus [NHMW]

♀♀ [NHMUK]

♂ [FSCA]


120.5

164.5

132.5

109.5–152.5

-

Body

109.0

152.0

121.0

102.5–131.0

94.0

Pronotum

4.8

5.4

5.1

4.7–5.7

3.9

Mesonotum

20.1

27.0

22.8

17.9–26.3

15.3

Metanotum

5.5

8.0

7.2

7.0–7.4

14.0**

Median segment

7.2

9.4

8.1

6.2–9.9

-

Tegmina

7.8

11.2

8.0

7.3–10.3

11.2

Alae

6.1

13.0

> 9.0

6.8–9.0

53.5

Profemora

16.5

20.5

16.3

15.1–18.3

15.7

Mesofemora

13.4

17.2

14.4

13.2–16.3

14.9

Metafemora

19.6

24.9

19.3

18.3–21.8

19.5




101

TABLE 18. (Continued) ♀ HT pulverulentus [NHMUK]

♀ LT murinus [NHMW]

♀ PLT murinus [NHMW]

♀♀ [NHMUK]

♂ [FSCA]

Protibiae

14.0

18.9

14.9

14.5–16.7

14.8

Mesotibiae

10.8

15.0

12.1

11.6–13.2

12.3

Metatibiae

15.7

21.8

16.9

16.0–27.6

16.2

Antennae

-

> 34.0

> 33.0

> 40.0

ca. 50.0

** metanotum + median segment

Diapherodes laevicollis (Redtenbacher, 1908) (Figs. 183–189, 372) Diapherodes laevicollis Redtenbacher, 1908: 434. HT, ♀: Coll. Br. v. W., Jamaica, Staudinger; det. Br. v. W. Diapherodes laevicollis; 18.887 [NHMW, No. 839]. Moxey, 1972: 106 (in litt.). Brock, 1998a: 38. Otte & Brock, 2005: 121. Aplopus (?) laevicollis, Rehn & Hebard, 1938: 52. Further material [1 ♂]: JAMAICA: 1 ♂: Jamaica: Saint Andrew, Parrish, Hard war Gap, 26–27-VII-1985, C.B. & H.V. Weems, G.B. Edwards [FSCA].

Diagnosis: Differing from the other two species in the jamaicensis species-group by the strongly laterally expanded abdominal tergum VII (Fig. 186) and strongly swollen, transverse abdominal segments II–IV of ♀♀, as well as the black scapus and pedicellus of ♂♂. It is closely related to D. jamaicensis (Drury, 1773) but in addition to these characteristic features distinguished by: the broader body (♀♀ in particular); more robust legs and longer alae of both sexes; more blunt cephalad humps of ♀♀ as well as the broader abdominal tergum VIII (Fig. 188) and much longer terminal hook of the vomer of ♂♂ (Fig. 372). Description: ♀ (Figs. 183–184). Medium-sized (body length including subgenital plate 84.0–110.0 mm) and very massive member of the jamaicensis species-group with a distinctly widened mesothorax and abdominal segment VII; tegmina (7.0–10.0 mm) and alae (7.0–11.0 mm) rather well developed. Colour bright apple green, the meso- and metapleurae ochre with a white longitudinal stripe along lower margin. Meso- and metasternum dull green with a brownish wash, ventral surface of abdomen mostly whitish. Tegmina and costal region of alae bright green with the anterior margin broadly brown and interiorly bordered by a rather narrow, weakly defined yellow stripe. Anal region of alae plain pink. Antennae ochre dorsally and black ventrally. Eyes pale ochre. Tips of the anterior mesonotal spines dark brown. Head: Indistinctly longer than wide, strongly globose, the cheeks ± parallel-sided. Vertex convex and covered with several differently sized tubercles; in centre with two prominent, conical humps, the dextral carinate and the sinistral one considerably smaller. Posterior of head with a transverse row of four distinct tubercles. Eyes of moderate size, sub-circular and their length contained just a little more than 2x in that of cheeks. Antennae broken in the holotype. Thorax: Pronotum about as long but narrower than head, 1.5x longer than wide and gently narrowing towards the posterior. Transverse median depression distinct, curved and almost reaching lateral margins of segment. Surface unarmed except for a pair of very low tubercles in anterior portion and a pair of short spines just before posterior margin. Mesothorax about 1.5x longer than head and pronotum combined, constricted anteriorly and strongly gradually widened towards the posterior. Mesonotum with a very faint longitudinal median carina and widened towards the posterior with posterior margin almost 3x wider than anterior margin (Fig. 184); surface sparsely granulose, with a distinct anterior pair of spines and a longitudinal row of small tubercles along lateral margins. Mesopleurae with a marginal row of 10–15 spiniform tubercles or short spines; metapleurae merely with


HENNEMANN ET AL.

a few granules. Pro-, meso- and metasternum smooth. Tegmina broadly ovate, indistinctly longer than wide, reaching about 1/3 the way along median segment and with the central protuberance very shallow. Alae slightly longer than tegmina and reaching to posterior margin of median segment.

FIGURES 183–189. Diapherodes laevicollis (Redtenbacher, 1908). 183. ♀ HT: Jamaica (dorsolateral view) [NHMW, No. 839]; 184. ♀ HT: Jamaica (dorsal view) [NHMW, No. 839]; 185. ♂: Jamaica, Saint Andrew, Parrish [FSCA]; 186. ♀ HT: Apex of abdomen (dorsal view—apex of subgenital plate reconstructed based on ♀ in NHMJ) [NHMW, No. 839]; 187. ♀ HT: Apex of abdomen (lateral view—apex of subgenital plate reconstructed based on ♀ in NHMJ) [NHMW, No. 839]; 188. ♂ apex of abdomen (dorsal view) [FSCA]; 189. ♂ apex of abdomen (dorsal view) [FSCA].



103

Abdomen: Median segment longer than metanotum, about 1.3x longer than wide and very gently widening towards the posterior; smooth. Abdomen strongly swollen sub-basally with segment II widening towards the posterior, III widest, IV–V gradually narrowing and VI almost parallel-sided. II–V wider than long, II trapezoidal and longer than III–V; these equal in length with III 2x wider than long. VI hardly wider than VIII–X and about 1.4x wider than long. Tergum VII greatly laterally deflexed with the lateral margins rounded and almost 2x wider than VI (Fig. 186). Tergites II–IV with two irregular converging carinae, V–VII with four roughly parallel, longitudinal carinae which are most decided on VII (Fig. 186). Sternites II–VII smooth except for a fine carina along lateral margins. Praeopercular organ formed by an elongate brown tubercle close to posterior margin of sternum VII. Tergites VIII–X narrowest segments and roughly of uniform width. VIII about ¾ the length of VII and very slightly constricted medially, about 1.3x longer than wide; IX quadrate. Anal segment a little longer than IX, with a slight longitudinal median carina and flattened towards the posterior; posterior margin rounded and entire (Fig. 186). Epiproct small and semi-circular with a keel dorsally (Fig. 186). Cerci very small, round in crosssection and tapered towards a pointed apex. Subgenital plate elongate, lanceolate, keeled longitudinally and projecting over apex of abdomen by more than the length of anal segment (Figs. 186–187). Legs: Profemora and mesofemora shorter than mesothorax, metatafemora reaching to posterior margin of abdominal segment IV. Medioventral carina of profemora with three small spines, of meso- and metafemora with five rather strong and back-curving spines. Posteroventral carina of meso- and metafemora with one, the anteroventral carina with two sub-apical spines. Tips of all femoral spines black. Dorsal carinae of meso- and metafemora each with a low triangular sub-apical elevation. Anterodorsal carina of protibiae slightly lamellate and undulate, in meso- and metatibiae with a shallow rounded elevation sub-basally. Basitarsi about as long as following two tarsomeres combined. Dorsal carina of probasitarsus very gently rounded. ♂ (Fig. 185). Fairly small (body length 69.0 mm) and robust for the jamaicensis species-group, body surface sub-glabrous, with very long alae (42.0 mm) that reach as far back as the posterior margin of abdominal tergum VIII. General colouration bright green, the dorsal surface of the anal segment and lateral margins of the pronotum yellow. Head yellow with a broad green postocular streak. Anterior margin of tegmina and the basal portion of the same region of the alae white, otherwise pale green with the veins mid green. Anal region of alae pale translucent pink. Eyes mid brown. Antennae ochraceous and becoming slightly darker towards the apex, scapus and pedicellus black. Anterior spines of the mesonotum black, cerci and tarsi pale creamish brown. Head: Globose, hardly wider than long, otherwise generally as in ♀♀ but cephalad tubercles less distinct. Eyes prominent and projecting hemispherically, their length contained hardly 1.5x in that of cheeks. Antennae projecting over posterior margin of abdominal segment II and moderately robust; with about 53 antennomeres. Scapus oval in cross-section, about 1.2x longer than wide and with the lateral margins rounded. Pedicellus cylindrical and about 2/ 3 the length of scapus, III shorter than pedicellus. Thorax: Pronotum about equal in length but slightly narrower than head, roundly rectangular and about 1.6x longer than wide; median transverse depression distinct, very gently curved and almost reaching lateral margin of segment. Surface smooth except for a very few minute granules; a slightly more distinct pair of granules before transverse median depression and another near posterior margin. Mesothorax 1.8x longer than head and pronotum combined. Mesonotum in front with a pair of distinct, forward-pointing spines and behind with two longitudinal rows of four spiniform tubercles which decrease in size towards the posterior; a row of small granules is present along lateral margins. Mesopleurae with a marginal row of about 10 small tubercles; metapleurae with a few small granules. Mesosternum with a few scattered granules in anterior portion, metasternum smooth. Tegmina oval and with a longitudinal central protuberance and reaching more than half way along median segment. Alae ± reaching to posterior margin of abdominal tergum VIII (Fig. 185). Abdomen: Segments II–VII roughly of uniform width and slightly decreasing in length, all rectangular; II 2.5x and VII only about 2.2x longer than wide. Sternites II–VII smooth. Tergum VIII about ¾ the length of VII and with the lateral margins distinctly expanded and rounded (Fig. 188). IX a little shorter than VIII and very slightly narrowing towards the posterior. Anal segment about as long as IX, roughly parallel-sided, the posterior margin bilobate with a distinct median emargination (Fig. 188). Vomer with the basal portion fairly small and the terminal hook very long, slender, slightly up-curving and distinctly longer than basal portion (Fig. 372). Cerci almost as long as anal segment, slender, cylindrical and gently in-curving with the apex blunt. Poculum moderately convex, cymbiform and longitudinally carinate; roughly reaching to posterior margin of tergum IX (Fig. 189).


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Legs: Shape and armature generally as in ♀♀, but dorsal carinae of meso- and metafemora entirely smooth and tibiae not elevated sub-basally. Comments: Redtenbacher (1908: 434) described Diapherodes laevicollis based on a unique ♀ in NHMW. The fact that only four specimens have been recorded so far suggests this distinctive species is apparently rare and considerably less abundant than the sympatric D. jamaicensis (Drury, 1773). Based on the record of a museum specimen, Moxey (1972: 10) stated H. laevicollis to be found on the endemic Eugenia alpina (Myrtaceae) which might thus be a natural food-plant in Jamaica. Moxey (1972: 106, in litt.) recorded body lengths of 84.0–110.0 mm based on two ♀♀ in NHMJ. Eggs unknown. Distribution: Jamaica: Portland, Summit of Blue Mountain Peak [NHMJ; Moxey, 1972: 106, in litt.] & Saint Andrew [FSCA]). Endemic. Number of specimens examined: 2 TABLE 19. Measurements of Diapherodes laevicollis Redtenbacher, 1908 [mm] ♀, HT [NHMW]

♂ [FSCA]

109.0*

-

Body

99.0

69.0

Pronotum

5.3

3.1

Mesonotum

17.5

11.4

Metanotum

14.0**

9.9**

-

-

Tegmina

10.0

9.7

Alae

11.0

42.0

Profemora

16.1

11.2

Mesofemora

13.8

10.6

Metafemora

19.8

14.9

Protibiae

15.8

10.8

Mesotibiae

11.9

9.3

Metatibiae

18.2

12.2

Antennae

-

37.0


Median segment

* according to Redtenbacher (1908: 435) ** metanotum + median segment

5.5. Genus Haplopus Burmeister, 1838 Type-species: Cyphocrana micropterus St. Fargeau & Audinet-Serville, 1828: 445, by monotypy. Haplopus Burmeister, 1838: 560, 576. [Emendation and replacement name for Aplopus Gray, 1835] De Haan, 1842: 107, 127. Westwood, 1859: 85 (in part). Saussure, 1871–1872: 192. Stål, 1875: 31. Bolivar, 1888: 140. Kirby, 1904a: 363 (in part). Redtenbacher, 1908: 429 (in part). Zompro, 2005a: 30. Otte & Brock, 2005: 150 (in part). Aplopus Gray 1835: 34. [Preoccupied by Aplopus Dejean, 1821 (Coleoptera: Curculionidae)] Rehn, 1904: 63 (in part).



105

Caudell, 1905: 83. Rehn, 1909: 200. Karny, 1923: 240. Rehn & Hebard, 1938: 52. Adams & Adams, 1982: 263. Brock, 1998c: 33. Cyphocrana, St. Fargeau & Audinet-Serville, 1828: 445. Diapherodes Gray, 1835: 33 (in part). Moxey, 1972: 85 (in litt.; in part). [As a synonym of Aplopus Gray, 1835 in error] Phasma, Stoll, 1813: 61 & 1788: pl. 21: 77 (♀). [Not: Haplopus, Shelford, 1908: 355, 365. → Pterinoxylus Audinet-Serville, 1838]

Description: ♂♂, ♀♀. Medium-sized to large (body length ♀♀ (including subgenital plate) 115.5–172.0 mm, ♂♂ 82.0–103.0 mm), slender and elongate Haplopodini, ♀♀ brachypterous, ♂♂ brachypterous or with well developed alae. Body ± cylindrical in cross-section. Colouration of ♀♀ various shades of brown, straw and grey, rarely greenish. ♂♂ rather colourful insects and sometimes multi-colorous but mostly green or greenish brown; body surface glabrous. Head indistinctly longer than wide and globose, vertex ± convex and distinctly bi-spinose or bicornute; the dextral spine larger than the sinistral. Antennae filiform in ♀♀ and considerably thickened and robust in ♂♂; in ♀♀ almost as long as head and complete thorax combined, in ♂♂ reaching at least to abdominal segment III. Pronotum about as long as head and with a ± distinct pair of anterior tubercles or spines; otherwise ± tuberculose. Mesothorax elongate and parallel-sided, at least 2x longer than head and pronotum combined. Mesonotum in ♀♀ rather irregularly and to a variable degree armed with granules or spiniform tubercles, in ♂♂ armed with a variable number of distinct spines. Mesopleurae smooth in ♂♂ and with a longitudinal row of tubercles in ♀♀. Meso- and metasternum irregularly set with a variable number of granules or tubercles; metasternum may be smooth in ♂♂. Tegmina broadly oval and with a moderately distinct central hump in ♂♂; ± reaching posterior margin of metanotum. Alae of ♀♀ ± as long as tegmina; of ♂♂ variable, either slightly shorter than tegmina or reaching as far as abdominal segment VI. Anal region reticulate with distinct brown to black radial and transverse veins, translucent pink or red in ♂♂. Abdomen considerably longer than head and thorax combined. Median segment longer than metanotum, smooth. Segments II–VII distinctly longer than wide. Tergites unarmed in ♂♂, II–V often with a ± distinct pair of posterior spines in ♀♀. VII parallel-sided, with lateral margins gently rounded posteriorly, or with a small posterolateral lobe or tooth. VIII and IX narrower (♀♀) or slightly wider (♂♂) than previous segments. Sternites II–VII smooth. Praeopercular organ of ♀♀ formed by a ± distinct, elongate and longitudinal wart-like median tubercle close to posterior margin of sternum VII (Figs. 288–289). Anal segment with a longitudinal median carina and the posterior margin with a ± distinct median notch; tapered in ♀♀. Epiproct very small in ♂♂, triangular or shield-shaped and ± projecting beyond anal segment in ♀♀. Vomer of ♂♂ well developed, broadened basally and with an elongate, ± papillate terminal hook (Figs. 304–310). Cerci straight, small and tapered in ♀♀, obtuse, about as long as anal segment and cylindrical in ♂♂. Poculum of ♂♂ convex, cup-like, often longitudinally carinate and with a blunt hump or spine basally. Subgenital plate of ♀♀ long, ± lanceolate with the apex narrow and ± pointed; projecting over apex of abdomen by at least the combined length of tergites IX and X. Legs of moderate length, pro- and mesofemora shorter than mesothorax in ♀♀, about as long or a little longer in ♂♂. Hind legs ± reaching (♂♂) or clearly not reaching (♀♀) to the apex of abdomen. Two outer ventral carinae of meso- and metafemora each with 1–3 sub-apical spines, the medioventral carina of these femora with a longitudinal row of 4–7 ± distinct spines. Dorsal carinae of all femora unarmed, those of the meso- and metafemora often forming a shallow to acutely triangular sub-apical tooth or lobe. Tibiae unarmed except for a few small teeth in the apical portion of the medioventral carina; anterodorsal carina in ♀♀ sometimes gently elevated sub-basally and sub-apically. All basitarsi slender and rather elongate, longer than following two tarsomeres combined. Eggs: Medium-sized (capsule length 3.1–4.4 mm), capsule ovoid, round in cross-section and > 1.5x longer than wide, polar area rounded. Capsule surface minutely granulose or rugulose. Micropylar plate elongately heartshaped, the anterior end narrowed and the widened posterior end with a prominent median notch; > 2/3 as long as capsule. No median line (Fig. 40). Operculum with a prominent, raised, squamiform, hat or knob-like capitulum. Colouration of capsule variable, often prettily mottled in various shades of brown, straw, cream, grey and orange. Differentiation: Very close to Parhaplopus n. gen. but differing by: the convex and distinctly bi-cornute vertex and anterior pair of tubercles or spines on the pronotum of both sexes; slender mesothorax and slender probasitarsus of ♀♀, as well as the reticulate anal region of the alae; larger and conspicuously thickened cerci and


HENNEMANN ET AL.

more convex poculum with a ± distint central spine of ♂♂. The eggs clearly differ from those of Parhaplopus n. gen. by having the capsule surface much less sculptured and possessing a ± distinct conical, hat or knob-like capitulum. For a more detailed differentiation see Table 29. Comments: Burmeister (1838: 576) established Haplopus as an amendment to Aplopus Gray, 1835 based on the original Greek pronounciation. This was however an unjustified emendation (ICZN, 1999: Article 32.5) and hence Haplopus is a junior objective synonym of Aplopus Gray (ICZN, 1999: Article 33.2.3). Aplopus Gray, 1835 however is preoccupied and a junior homonym of Aplopus Dejean, 1821 (Coleoptera: Curculionidae) and therefore replaced by Haplopus Burmeister, 1838 (Zompro, 2005a: 30), which is an available name (ICZN, 1999: Article 19.1). In the composition it was used by most former authors (e.g. Kirby, 1904a; Redtenbacher, 1908 or Moxey, 1972 in litt.) Haplopus Burmeister, 1838 was not a natural grouping, which had already been suggested by Rehn & Hebard (1938: 52). This is obviously seen in two clearly distinct types of eggs and several obvious distinctive features of the insects. Eggs of typical Haplopus exhibit a more or less prominently conical, knob or hat-like capitulum and have a rather faintly coriaceous capsule surface, whereas those of the second group lack a central capitulum and have the capsule surface prominently sculptured with raised ridges and humps. Species belonging to the second group are here transferred to the “jamaicensis species-group” of Diapherodes Gray, 1835. One species formerly attributed to Haplopus, i.e. Haplopus dubius (Gray, 1835), is definitely misplaced and here removed from the genus. The exact systematic position however remains uncertain (→ 5.9). Distribution (Fig. 376): Bahamas, Florida Keys, Dry Tortugas, Greater Antilles (Hispaniola and Puerto Rico), Gonaive, Mona Island east of Hispaniola, Navassa Island west of Hispaniola, Virgin Islands (St. Croix, St. Thomas, St. John, Anegada, Guana, Little Thatch, Moskito and Tortola), Cayman Islands and Swan Islands. Although no definite records are available so far, the genus is most certainly also represented on the Caicos Islands north of Hispaniola (Fig. 312). Species included: 1. Haplopus bicuspidatus de Haan, 1842: 128. [Distribution: Hispaniola] 2. Haplopus brachypterus n. sp. [Distribution: Hispaniola] 3. Haplopus intermedius n. sp. [Distribution: Hispaniola] 4. Haplopus micropterus (St. Fargeau & Audinet-Serville, 1828: 445) [Cyphocrana]. = Phasma angulata Stoll, 1813: 61, pl. 21: 77 (♀). = Haplopus bituberculatum de Haan, 1842: 128. n. syn. = Haplopus cythereus Westwood, 1859: 86, pl. 18: 5, 5a & b (♂). n. syn. = Haplopus ligiolus Redtenbacher, 1908: 432. n. syn. = Haplopus ligia Westwood, 1859: 89, pl. 1: 1, 1a, 1b (♂) & 2, 2a (♀). n. syn. = Haplopus obtusus Redtenbacher, 1908: 431. n. syn. = Diapherodes spinipes Gray, 1835: 34. n. syn. [Distribution: Hispaniola, Mona Id., Puerto Rico, St. Croix & Virgin Islands] 5. Haplopus scabricollis (Gray, 1835: 34) [Diapherodes]. = Aplopus mayeri Caudell, 1905: 83. n. syn. = Aplopus similis Caudell, 1904: 65. n. syn. [Distribution: Bahamas, S-Florida, Florida Keys, Dry Tortugas, Navassa Id., Cayman Ids. & Swan Ids.] 6. Haplopus sobrinus n. sp. [Distribution: Cuba] 7. Haplopus woodruffi n. sp. [Distribution: Cayman Islands (Cayman Brac)]



107

Keys to the species of Haplopus ♀♀ 1. 2. -

3. 4. 5. 6. -

Vertex convex; alae as long as or longer than tegmina and covering > 2/3 of median segment. . . . . . . . . . . . . . . . . . . . . . . . . . 4 Vertex gently rounded; alae at best equal in length to tegmina and covering at best ½ of median segment . . . . . . . . . . . . . . . . 2 Spines of the vertex blunt or obsolete; anal region of alae not red . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 Spines of the vertex acute (Fig. 268); alae much shorter than tegmina and covering only 1/3 of median segment, anal region red; epiproct distinct, shield-shaped (Fig. 273); Bahamas, Florida Keys, Dry Tortugas, Navassa Island, Cayman Islands & Swan Islands . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .scabricollis Spines of the vertex obsolete (Fig. 277); tegmina distinctly shorter than alae and covering only 1/3 of median segment (Fig. 278), anal region reticulate; epiproct acutely triangular and keeled dorsally; Cuba. . . . . . . . . . . . . . . . . . . . . . . .sobrinus n. sp. Spines of the vertex blunt (Fig. 281); alae subequal in length to tegmina and covering about ½ of median segment, anal region pale transparent pink; epiproct very small (Fig. 283); Cayman Brac . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . woodruffi n. sp. Abdominal tergum VII parallel-sided; epiproct small, triangular; only Hispaniola . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5 Abdominal tergum VII ± broadened / expanded, or with a lobe posterolaterally; epiproct distinct. . . . . . . . . . . . . . . . . . . . . . . 6 Epiproct very small; anal segment with a shallow concave excavation posteromedially (Fig. 208); profurcasternum tuberculate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . brachypterus n. sp. Epiproct acutely triangular; anal segment with a distinct, triangular posteromedian incision (Fig. 197); profurcasternum smooth . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . bicuspidatus Epiproct roundly triangular to semi-circular, ± shield-shaped (Figs. 241–242); mesonotum < 2.3x longer than head and pronotum combined; alae covering > ¾ of median segment; antennae red ventrally . . . . . . . . . . . . . . . . . . . . . . . . . . . . . micropterus Supraanal plate acutely triangular (Fig. 215); mesonotum 2.4x longer than head and pronotum combined; alae covering < ¾ of median segment; antennae brown . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . intermedius n. sp.

♂♂ 1. 2. 3. -

4. -

5. -

Brachypterous, alae not reaching to apex of median segment . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 Alae well-developed, exceeding abdominal tergum IV . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5 Large (body > 80 mm); horns of the head prominent and acute; three terminal abdominal segment scarcely broader than previous; tergum VII with a posterolateral tooth or lobe . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 Small (body < 80 mm); horns of the head very small and blunt (Fig. 282); three terminal abdominal segments strongly swollen and club-like, distinctly broader than previous (Fig. 285); tergum VII slender; Cayman Brac . . . . . . . . . . . . . . woodruffi n. sp. Alae scarcely shorter than tegmina and covering roughly ½ of median segment, anal region reticulate; no white markings on pronotum and median segment; . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4 Alae considerably shorter than tegmina and covering only 1/3 of metanotum, anal region red; lateral margins of pronotum white; median segment with a longitudinal white marking; Bahamas; Florida Keys; Dry Tortugas, Navassa Island, Cayman Islands & Swan Islands . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .scabricollis Abdominal tergum VII slightly expanded posteriorly (Fig. 210); dorsal carinae of meso- and metafemora smooth; vomer with base large and terminal hook only about 1.2x longer than basal portion (Fig. 350) . . . . . . . . . . . . . . . . . . . . brachypterus n. sp. Abdominal tergum VII with a distinct triangular posterolateral lobe (Fig. 217); dorsal carinae of meso- and metafemora with a low triangular tooth sub-apically; vomer with base fairly small and the terminal hook papillate and almost 2x longer than basal portion (Fig. 351) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . intermedius n. sp. Head with cheeks white (Figs. 231–233); alae reaching to abdominal segment VI; anterior of pronotum with a distinct pair of spines (Figs. 231–233) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . micropterus Head uniformly green (sometimes with a faint yellow marking on frons, Fig. 194); alae at best reaching half way along abdominal segment V; anterior of pronotum merely with a pair of small tubercles (Fig. 194) . . . . . . . . . . . . . . . . . . . . . . . bicuspidatus

Haplopus bicuspidatus de Haan, 1842 (Figs. 190–201, 338, 357, 380) Haplopus bicuspidatus de Haan, 1842: 128. HT, ♀: ?; RMNH Leiden, TYPE, Haplopus bicuspidatum de Haan, 1842 [RMNH]. Westwood, 1859: 87. Kirby, 1904a: 364. Redtenbacher, 1908: 433. [Suggesting possible synonymy with H. ligius Westwood, 1859] Otte & Brock, 2005: 150. Aplopus bicuspidatus, Bragg, 1996: 109. Bragg, 2001: 706. Aplopus sp., Adams & Adams, 1982: 263. Gange, 1990: 18. [Culture report] Kirby, 1904a: 364 (in part). Redtenbacher, 1908: 432 (in part). Moxey, 1972: 114 (in litt.; in part).


HENNEMANN ET AL.

Further material [2 ♂♂, 9 ♀♀, eggs]: HISPANIOLA (DOMINICAN REPUBLIC): 1 ♀: Dominican Republic, Proyecto Biologico «La Huronera» La Victoria, Distrito Nacional, IX.1992, Kelvin A. Guerrero, NHMUK(E) 2005-98 [NHMUK]; 1 ♀: Santo Domingo, Dominican Republic 1925; G. Russo coll; ANSP; USNM; w0139; Diapherodes spinipes Gray det. C.F. Moxey 1972 [USNM]; 1 ♀: Santo Domingo, Dominican Republic 1925; G. Russo coll; USNM; w0140; Aplopus ligius Westw. ♀ Topotype, A.n.c.; Diapherodes spinipes Gray det. C.F. Moxey 1972 [USNM]; 1 ♂, 2 ♀♀: ex Zucht: F. Hennemann, urspr.: Dominikanische Republik, 1993, PSG No. 48 [coll. FH, No. 03421 to 3]; 1 ♂, 4 ♀♀, eggs: ex Zucht: F. Hennemann, urspr.: Dominikanische Republik, VI.1995, PSG No. 48 [coll. FH, No. 0342-4 to 8 & ED].

Diagnosis: Very similar and presumably the sister-taxon of H. brachypterus n. sp.. Females differ by: the larger size of both sexes; slight posterolateral lobe of abdominal tergum VII; alae which are longer than the tegmina (shorter in brachypterus); deeper posteromedian incision of the anal segment and larger and more acutely triangular epiproct (Fig. 197). Males differ from those of H. brachypterus n. sp. by: the fully developed alae (37.0–41.0 mm); almost entirely green head (Fig. 194); obsolete anterior tubercles of the pronotum (Fig. 194); more decidedly excavated anal segment (Fig. 199) and shorter terminal hook of the vomer (Fig. 349). From H. micropterus (St. Fargeau & Audinet-Serville, 1838) it is easily distinguished by: the more elongate and slender body and less distinct anterior pair of tubercles on the pronotum of both sexes; smaller and acutely triangular supraanal plate of ♀♀ (Fig. 197), as well as the entirely green head (Fig 194); ochraceous antennae (orange to red in micropterus) and shorter alae of ♂♂, which merely reach to the anterior margin of abdominal segment V (reaching to tergum VI in micropterus). The eggs clearly differ from those of H. micropterus by the more elongate and more decidedly sculptured capsule and roundly convex to irregularly conical operculum (Figs. 200–201). Description: ♀ (Figs. 190–191). Of moderate size (body length including the subgenital plate 134.0–153.5 mm) but slender for the genus with a rather weakly developed thoracic armature. Colour ranging from drab over pale to dark brown, sometimes with a reddish or greenish wash. Abdomen to a variable degree furnished with paler and darker mottling and speckles. Head with the cheeks irregularly whitish and an elongate black marking running from the upper portion of the eyes towards the occipital horns; occasionally a further pair of black spots on the frons between the bases of the antennae. Antennae pale to mid brown. Spines of the thorax either brown or very dull green. Tegmina and costal region of alae mid to dark brown, rarely with a greenish wash; the latter blackish basally. Anal region of alae transparent with all major longitudinal and transverse veins broadly marked by dark brown bands. Head: About 1.2x longer than wide, globose. Vertex rounded and with a pair of prominent but variably sized cephalad spines; these usually with a pointed tip (Fig. 193). Eyes circular and contained almost 3x in length of cheeks. Antennae ± reaching half way along median segment and consisting of 54–56 segments. Scapus almost 2x longer than wide and slightly narrowed towards the base. Pedicellus about half the length of scapus and distinctly shorter than III. Thorax: Pronotum a little longer but slightly narrower than head, about 1.5x longer than wide, roughly rectangular but the lateral margins with a deep ± semi-circular median emargination. Transverse median sulcus deep, strongly curved but not reaching lateral margins of segment. Dorsal surface with a pair of low spiniform tubercles in the anterior portion, otherwise unarmed (Fig. 193). Prosternum and profurcasternum smooth. Mesothorax 2.5–2.6x longer than head and pronotum combined. Mesonotum rather narrow anteriorly and very slightly widened in median portion which is about as wide as posterior margin; surface with a variable number of spiniform tubercles or low spines; a rather defined marginal row of evenly sized tubercles is present laterally. Meso- and metapleurae with an irregularly longitudinal marginal row of spiniform tubercles. Meso- and metasternum sparsely granulose. Metanotum about ¼ the length of mesonotum, unarmed. Tegmina oval, coriaceous, with the venation very distinct, dense and irregularly disposed, and slightly projecting over posterior margin of metanotum; median protuberance very shallow. Alae longer than tegmina and reaching 2/3 to ¾ the way along median segment.



109

FIGURES 190–192. Haplopus bicuspidatus de Haan, 1842. 160. HT ♀: locality unknown [RMNH]; 191. ♀: captive reared from Dominican Republic [coll. FH, No. 0342-2]; 192. ♂: captive reared from Dominican Republic [coll. FH, No. 0342-1].

Abdomen: Median segment 2x longer than wide and slightly narrowed medially. All segments unarmed. Segments II–V of roughly equal length, VI a little shorter than V, VII just a little more than ¾ the length of V; on average 2x longer than wide. VII with lateral margins very slightly expanded posteriorly (Fig. 196). Praeopercular organ formed by a rough V-shaped swelling and two slight semicircular expansions at posterior margin of sternum VII (Fig. 338). VIII slightly shorter than VII, gently narrowed medially and almost 2.5x longer than wide. IX rectangular, slightly longer than wide and roughly half the length of VIII. Anal segment with a slight longitudinal carina which becomes gradually more decided towards the posterior, narrowed in posterior half and with a small triangular posteromedian indentation. Epiproct very small and acutely triangular (Fig. 197). Cerci very small, conical with a fairly acute tip and slightly laterally compressed; hardly projecting over posterior margin of anal segment. Subgenital plate very long, lanceolate, longitudinally carinate and with a ± acute apex; extending greatly over apex of abdomen (Figs. 195–196).


HENNEMANN ET AL.

FIGURES 193–201. Haplopus bicuspidatus de Haan, 1842. 193. ♀ head and pronotum: captive reared from Dominican Republic [coll. FH, No. 0342-5]; 194. ♂ head and pronotum: captive reared from Dominican Republic [coll. FH, No. 0342-8]; 195. ♀ apex of abdomen (lateral view); 196. ♀ apex of abdomen (dorsal view); 197. ♀ anal segment and epiproct (dorsal view); 198. ♂ apex of abdomen (lateral view); 199. ♂ apex of abdomen (dorsal view); 200. Egg:captive reared from Dominican Republic (dorsal view) [coll. FH, No. 0342-E]; 201. Egg captive reared from Dominican Republic (lateral view) [coll. FH, No. 0342-E].

Legs: Profemora ¾ the length of mesothorax, mesofemora reaching about ¾ the way along abdominal segment II and metafemora almost reaching posterior margin of abdominal segment IV. Profemora occasionally with 1–3 minute spines in the apical half of the medioventral carina. Anteroventral carina of meso- and metafemora with two, posteroventral carina with only one sub-apical spine; medioventral carina armed with four distinct spines. Basitarsi about as long as following three tarsomeres combined. ♂ (Fig. 192). Of moderate size (body length 97.0–98.0 mm) and rather slender for the genus with well developed alae (length 37.0–41.0 mm). Colouration fairly complex with head and great parts of mesometapleurae, ventral body surface and legs green. Head plain green with the frons slightly yellowish (Fig. 194). Dorsal surface of thorax creamish mid brown, and drab to pale brown in abdomen. Spines of thorax with black tips. Meso- and metapleurae each with a pale brown longitudinal band along lower margin. Lateral margins of abdominal tergites VIII and IX broadly white. Tegmina and costal region of alae pale to mid brown, the latter with



111

a few irregularly disposed pale markings along anterior margin and black basally. Basal portion of posterior margin of tegmina broadly white. Anal region of alae pink with all major veins brown. Antennae drab to ochraceous. Tarsi pale brown. Head: Generally as in ♀♀ but with the cephald horns more slender and acute (Fig, 194). Eyes more prominent, projecting hemispherically and their length contained less than 2x in that of cheeks. Antennae very robust and ± reaching posterior margin of abdominal segment III; with > 50 segments (broken in all specimens at hand). Thorax: Pronotum slightly longer but narrower than head, general shape as in ♀♀; surface smooth except for a faint pair of low tubercles in the anterior portion (Fig. 194). Mesothorax about 2.3x longer than head and pronotum combined. Mesonotum with 4–8 small but pointed, rather irregularly disposed spines in the anterior 2/3. Mesosternum and metasternum with a few irregularly disposed granules to low spiniform tubercles. Tegmina oval and slightly projecting over posterior margin of metanotum, central protuberance rather distinct and roundly conical. Alae slightly projecting over posterior margin of abdominal segment IV. Abdomen: Segments II–IV of equal length and about 3.5x longer than wide. V–VII gradually decreasing in length with VII no more than 2.3x longer than wide. All tergites and sternites smooth. VII ± expanded posteriorly (Fig. 199). VIII shorter than VII and gently widening towards the posterior; IX ¾ the length of VIII and constricted medially. Anal segment with a faint longitudinal median carina which becomes gradually more decided towards the apex. Posterior portion strongly narrowed and laterally emarginated, the posterior margin with a distinct concave emargination medially (Fig. 199), slightly swollen and on ventral surface armed with several small, black incurving denticles. Epiproct small and with a posteromedian notch. Vomer with a large, roundly triangular base and a moderately long, up-curving terminal hook, which is about equal in length to basal portion (Fig. 357). Cerci large, obtuse, about equal in length to anal segment and slightly laterally compressed basally. Poculum moderately convex, cup-like and with a prominent, conical basal hump (Fig. 198); the posterior portion carinate longitudinally (Fig. 357). Legs: Pro- and mesofemora a little longer than mesothorax, metafemora projecting over posterior margin of abdominal segment IV. All legs less carinate than in ♀♀ but armature generally alike. Tarsi relatively more elongate and basitarsi a little longer than following three tarsomeres combined. Egg (Figs. 200–201): Rather large for the genus, capsule elongate ovoid and 1.8x longer than wide; dorsal surface more convex than ventral surface. Capsule surface strongly coriaceous and irregularly granulose. Micropylar plate almost 3/5 the length of capsule and generally shaped like a bold inverted “Y”; the anterior half narrow and roughly parallel-sided, the posterior portion strongly diverging. Posterior margin with a wide and deep triangular emargination. Micropylar cup small and wart-like. Operculum ± round in cross-section, irregularly conically raised and strongly tuberculate. Colouration of capsule variable, ranging from pale drab over ochre to grey and to a variable degree furnished with weakly defined, dark brown to black markings. Capitulum pale drab to grey. Measurements [mm]: Length 3.9–4.4, length (including operculum) 4.1–4.7, width 2.2–2.3, heigth 2.4–2.7, length of micropylar plate 2.0–2.1. Variability: This species shows much less intraspecific variability than e.g. H. micropterus (St. Fargeau & Audinet-Serville, 1838) or H. scabricollis (Gray, 1835). In addition to the size and colouration the only features that underlie some variation in ♀♀ are the length of the subgenital plate, number and size of the mesothoracic tubercles and size of the cephalad horns. No considerable variation is seen in the two ♂♂ at hand, except for one having eight and the other one only four spines on the mesonotum. One specimen only has two or three spines on the medioventral carina of the mesofemora. Comments: This species was originally described from a single ♀ in RMNH and just briefly recognized by subsequent authors (e.g. Westwood, 1859 and Redtenbacher, 1908). Examination of the holotype and comparison with captive reared specimens originating in the Dominican Republic has shown these to be conspecific. The captive reared specimens include the so far unknown ♂♂ and eggs. Since only one additional wild-caught specimen is recorded here and none are represented in the extensive material from the Dominican Republic at hand H. bicuspidatus appears to be apparently rare. Culture stock of H. bicuspidatus collected in the Dominican Republic was imported to Europe in the early 1980's (Adams & Adams, 1982) and included on the Phasmid Study Group culture-list as culture No. 48 “Aplopus sp.” (later corrected to Haplopus cytherea). In captivity it readily accepts rose (Rosa spp., Rosaceae), bramble (Rubus fruticosus, Rosaceae), raspberry (Rubus idaeus, Rosaceae), oak (Quercus robur & Q. petraea, Fagaceae),


HENNEMANN ET AL.

guava (Psidium guajava, Myrtaceae), eucalyptus (Eucalyptus gunnii, Myrtaceae) and hawthorn (Pyracantha spp., Rosaceae) as alternative food plants. Distribution (Fig. 380): Hispaniola (Dominican Republic: Distrito Nacional) [NHMUK, USNM]. Number of specimens examined: 12 TABLE 20. Measurements of Haplopus bicuspidatus de Haan, 1842 [mm] ♀, HT [RMNH]

♀♀ [coll. FH)]

♂♂ [coll. FH]

♀ [MNHU]

♂ [MNHU]


140.5

134.0–153.5

-

-

-

Body

125.0

122.5–131.5

97.0–98.0

134.5

105.0

Pronotum

5.1

5.1–5.3

3.7–3.9

5.3

4.0

Mesonotum

30.4

24.7–27.6

19.8–19.9

30.0

20.7

Metanotum

6.9

7.0–7.5

6.8

7.0

-

Median segment

9.2

8.5–9.1

7.9

8.9

-

Tegmina

10.2

8.2–10.2

10.0–11.2

10.0

10.8

Alae

11.3

9.6–12.9

37.0–41.0

12.0

44.5

Profemora

21.8

19.5–23.0

21.8–22.2

23.2

23.5

Mesofemora

19.9

17.2–19.8

21.4–25.2

19.8

21.0

Metafemora

26.3

22.8–25.1

24.0–25.8

25.5

27.0

Protibiae

22.8

19.8–23.1

22.2–23.0

23.9

24.5

Mesotibiae

20.2

17.8–20.0

19.1–19.6

20.0

21.3

Metatibiae

27.8

23.2–27.5

25.0

26.5

26.5

Antennae

-

46.0–50.0

> 53.0

> 45.5

73.0

* a captive reared ♀ in coll. FH is unusually small, measuring only 116.0 mm (including the subgenital plate) and hence excluded from the table above.

Haplopus brachypterus n. sp. (Figs. 193–210, 339, 358, 380) HT, ♂: Dominican Republic, RD-248 Entrance to Sabana Iglesia, Santiago Prov., 372 m, 19°20.196’N 70°45.483’W,29.IV.2004, DPerez, BHierro, R. Bastardo. (d) [USNM]. PT, 1 ♀, 1 ♂: Dominican Republic, RD-248 Entrance to Sabana Iglesia, Santiago Prov., 372 m, 19°20.196’N 70°45.483’W, 29.IV.2004, DPerez, BHierro, R. Bastardo. (d) [USNM]. PT, 2 ♂♂: Dominican Republic, Janico, Santiago Prov., 30.IX.1996, on Pictetia spinifolia (Tabacuelo), D. Perez, S. Navarro [USNM]. PT, 1 ♀, 1 ♂ (penultimate instar), 1 ♀ (nymph n4): Dominican Republic, RD-247 Rd. Inoa—El Caimito, nr. San Josè de las Matas, Santiago Prov., 552 m, 19°22.225’N 71°00.661’W, 28.IV.2004, D. Perez, BHierro, RBastardo (d/n) [USNM]. PT, 1 ♀: Dominikaner Republik Haiti; Santiago de los Caballeros, P. Thumb leg., Eing. Nr. 92/37; PHA 7, Zoologisches Museum Hamburg [ZMUH]. PT, 1 ♀: Dominikaner Republik Haiti; Santiago de los Caballeros, P. Thumb leg., Eing. Nr. 92/37; PHA 8, Zoologisches Museum Hamburg [ZMUH]. PT, 1 ♀: Dominikaner Republik Haiti; Santiago de los Caballeros, P. Thumb leg., Eing. Nr. 92/37; PHA 9, Zoologisches Museum Hamburg [ZMUH]. PT, 1 ♀: Dominikaner Republik Haiti; Santiago de los Caballeros, P. Thumb leg., Eing. Nr. 92/37; PHA 10, Zoologisches Museum Hamburg [ZMUH]. PT, 3 ♂♂: Dominikaner-Republik Haiti, Oktober 1936, Santiago de los Caballeros, P. Thumb leg., Eing. Nr. 92, 1937 [ZMUH—in alcohol].



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FIGURES 202–210. Haplopus brachypterus n. sp.. 202. ♀ PT: Dominican Republic, Santiago Province [USNM]; 203. ♂ HT: Dominican Republic, Santiago Province [USNM]; 204. Head and pronotum of ♀ PT [USNM]; 205. Head and pronotum of ♂ HT [USNM]; 206. Tegmina and alae of ♂ PT [USNM]; 207. Apex of abdomen of ♀ PT (lateral view) [USNM]; 208. Anal segment and epiproct of ♀ PT (dorsal view) [USNM]; 209. Apex of abdomen of ♂ HT (lateral view) [USNM]; 210. Apex of abdomen of ♂ HT (dorsal view) [USNM].


HENNEMANN ET AL.

Diagnosis: Very similar to H. bicuspidatus de Haan, 1842 and H. intermedius n. sp.. From the first it differs by: the slightly smaller dimensions of both sexes; presence of posterior tubercles on the pronotum; tuberculose profurcasternum; parallel-sided abdominal tergum VII; more shallow posteromedian excavation of the anal segment and smaller, transverse epiproct of ♀♀ (Fig. 208), as well as the conspicuously shortened alae (5.6–6.0 mm); distinct yellow markings on the frons and posterior portion of the cheeks (Fig. 205); more prominent and spiniform anterior spines of the pronotum (Fig. 205); less decidedly excavated anal segment (Fig. 208) and longer papillate terminal hook of the vomer of ♂♂ (Fig. 350). From H. intermedius n. sp., with which ♂♂ share the shortened alae, it can be distinguished by: the conspicuously less distinct posterolateral lobe of abdominal tergum VII and lack of a sub-apical tooth on the dorsal carinae of the meso- and metafemora of both sexes. ♀♀ also differ by: the slightly smaller dimensions; considerably smaller cephalad horns (Fig. 204); relatively longer mesothorax; absence of distinct paired posterior spines on abdominal tergites II–IV; absence of a sub-basal and sub-apical elevation of the anterodorsal carina of the meso- and metatibiae, and smaller epiproct (Fig. 208). Males have the anal region of the alae smaller and less well developed than in H. intermedius n. sp. (Fig. 206) and the cerci round in cross-section (compressed basally and carinate dorsally in intermedius). The short alae of ♂♂ also resemble H. scabricollis (Gray, 1835) from the Bahamas, Florida Keys, Navassa Island and Cayman Islands. They however clearly differ from ♂♂ of H.scabricollis by: the more slender body; lack of longitudinal white stripes along the lateral margins of the pronotum, and white median markings on the median segment and abdominal tergites VIII–X; convex and prominently bi-cornute vertex (Fig. 205); more decidedly excavated anal segment (Fig. 210) and more distinct central spine of the poculum (Fig. 209). Females are well distinguished from H. scabricollis by: the globose and prominently bi-cornute vertex (Fig. 204); longer alae; less decidedly tuberculose pronotum and smaller epiproct (Fig. 208). Etymology: The name “brachypterus” refers to the conspicuously shortened alae of both sexes. Description: ♀ (Fig. 202). Of moderate size (body length including the subgenital plate 131.0–139.0 mm) and slender for the genus; body surface slightly glabrous. Colour creamish mid to dark brown with abdomen to a variable degree furnished with paler and darker mottling and speckles. Head with the cheeks irregularly whitish and a black marking above the eyes; two further small black spots on the frons between the bases of the antennae. Antennae dark ochraceous with a very slight reddish wash ventrally. Spines of the thorax dark reddish brown. Anterior portion of metapleurae whitish. Tegmina and costal region of alae dark brown to black with the veinature sepia; the latter blackish basally. Anal region of alae transparent and all major longitudinal and transverse veins broadly marked with dark brown. Head: About 1.2x longer than wide, globose with the cheeks rather convex. Vertex rounded and with a pair of moderately sized and apically pointed cephalad spines (Fig. 204). Eyes circular and contained about 3x in length of cheeks. Antennae ± reaching half way along median segment and consisting of about 64 segments. Scapus 2.2x longer than wide and gently expanded pre-apically. Pedicellus less than half the length of scapus and about equal to III. Thorax: Pronotum a little longer but distinctly narrower than head, about 1.4x longer than wide, roughly rectangular but the lateral margins with a wide ± semi-circular emargination pre-medially. Transverse median sulcus deep, strongly curved but not reaching lateral margins of segment. Dorsal surface with a pair of low spines in the anterior portion and two pairs of small tubercles in the posterior portion (the posterior pair may be lacking, Fig. 204). Prosternum smooth. Profurcasternum with three pairs of tubercles; a distinct pair marking the anterolateral angles of the segment, a much smaller pair at the posterior angles and a further even smaller pair posterior of distinct anterior tubercles. Mesothorax about 2.8x longer than head and pronotum combined. Mesonotum fairly narrow anteriorly but otherwise almost parallel-sided; surface with a variable number of spiniform tubercles or low spines; a rather defined marginal row of evenly sized tubercles is present laterally. Meso- and metapleurae with an irregular longitudinal marginal row of short but acute spines. Mesosternum sparsely set with some very low, irregularly paired spines, metasternum usually with two pairs of tubercles. Metanotum about ¼ length of mesonotum, unarmed. Tegmina oval, coriaceous, with the venation very distinct, dense and irregularly disposed, and slightly projecting over posterior margin of metanotum; median protuberance very shallow and slightly displaced towards the apex of tegmen. Alae very indistinctly longer than tegmina and reaching about ¾ the length along median segment. Abdomen: Median segment 2x longer than wide and very gently narrowed medially. All segments unarmed, except for a pair of very shallow tubercles near posterior margin of tergum II. Segments II–IV very slightly



115

increasing in length, V as long as IV; the latter two segments about 2x longer than wide. VI a little shorter and narrower than V, almost 2.5x longer than wide. Tergum VII slightly shorter than VI and parallel-sided, the posterolateral angles very slightly rounded; dorsal surface with two short converging carinae in posterior half. Praeopercular organ formed by an acute, longitudinal ridge in posterior portion of sternum VII (Fig. 339). Tergum VIII slightly shorter than VII, gently narrowed medially and almost 2.5x longer than wide. IX rectangular and about 2/3 the length of VIII. Anal segment with a very faint longitudinal median carina, narrowed in posterior half and with a shallow posteromedian emargination. Epiproct very small, roughly triangular and the projecting part wider than long (Fig. 208). Cerci very small, conical with a rather acute apex and slightly compressed laterally; hardly projecting over posterior margin of anal segment. Subgenital plate very long, lanceolate, longitudinally carinate and with a ± acute apex; extending greatly over apex of abdomen (Fig. 207). Legs: Profemora about 2/3 the length of mesothorax, mesofemora reaching about 1/3 the way along abdominal segment II and metafemora reaching about half way along abdominal segment IV. Profemora occasionally with 1–3 very minute spines in the apical half of the medioventral carina. Anteroventral carina of meso- and metafemora with two, posteroventral carina with one sub-apical spine; medioventral carina armed with four distinct spines. Basitarsi about as long as following three tarsomeres combined. ♂ (Fig. 203). Of moderate size (body length 83.2–87.5 mm) and rather slender for the genus with conspicuously shortened alae (length 5.4–5.9 mm). Colouration rather complex with great parts of head, mesometapleurae, ventral body surface and legs bright green. Head green with the frons yellow and a yellow postocular stripe along cheeks (Fig. 205). Dorsal surface of thorax creamish mid brown, of abdomen drab to pale brown. Pronotum yellowish laterally. Dorsal spines of pro- and mesothorax dark reddish brown with black points. Mesoand metapleurae with a washed longitudinal yellow marking in posterior portion and each with a dull yellow longitudinal band along lower margin. Lateral margins of abdominal tergites VIII and IX broadly white. Tegmina and costal region of alae mid to dark greyish brown; anterior margin and basal portion of posterior margin of tegmina, as well as anterior margin of alae broadly white; base of alae blackish. Anal region of alae pink with all major veins brown (Fig. 206). Antennae drab to ochraceous. Tarsi pale to mid reddish brown. Head: Generally as in ♀♀ but with the cephalad horns more prominent, slender and acutely pointed (Fig. 205). Eyes more prominent, projecting hemispherically and their length contained only about 2x in that of cheeks. Antennae moderately robust; broken in all specimens at hand. Thorax: Pronotum slightly longer but narrower than head, general shape as in ♀♀; surface smooth except for a moderate pair of blunt spines or spiniform tubercles in the anterior portion (Fig. 205). Mesothorax about 2.5x longer than head and pronotum combined. Mesonotum with 7–8 pointed, irregularly dispersed but paired spines in the anterior 2/3. Mesosternum and metasternum with several irregularly dispersed ± spiniform tubercles. Tegmina oval and ± reaching posterior margin of metanotum, central protuberance very shallow. Alae small, slightly shorter than alae and reaching only about 2/3 the way along median segment (Fig. 206). Abdomen: Segment II a little shorter than III–IV and equal in length to V, about 2.7x longer than wide. III and IV longest segments and almost 3.5x longer than wide. V–VII gradually decreasing in length with VII no more than 2.3x longer than wide. All tergites and sternites smooth. VII slightly expanded posteriorly (Fig. 210). VIII shorter than VII and gently widening towards the posterior; IX about ¾ the length of VIII and narrowed towards the posterior. Anal segment with a faint longitudinal median carina which becomes gradually more decided towards the base. Posterior portion narrowed and laterally emarginated, the posterior margin with a shallow median emargination (Fig. 210), slightly swollen and on ventral surface armed with several small, black in-curving denticles. Epiproct very small (Fig. 210). Vomer with a rather small, roughly semi-circular base and a very long, papillate, up-curving terminal hook which is >1.5x longer than the basal portion (Fig. 358). Cerci large, obtuse and about equal in length to anal segment, very slightly laterally compressed basally. Poculum moderately convex, cuplike and with a prominent, acute and spiniform basal hump (Fig. 209); posterior portion carinate longitudinally (Fig. 358). Legs: Pro- and mesofemora very slightly shorter than mesothorax, metafemora projecting a little over posterior margin of abdominal segment IV. All legs less carinate than in ♀♀ but armature generally alike. Tarsi relatively more elongate and basitarsi a little longer than following three tarsomeres combined. Nymphs: The half-grown ♀ and penultimate instar ♂ nymph at hand are greyish dark brown with fine pale grey mottling and both lack the sub-apical dorsal lobe on the meso- and metafemora, which distinguish them from


HENNEMANN ET AL.

nymphs of the very similar H. intermedius n. sp.. The ♀ nymph has abdominal tergum VII with a distinct, rounded posterolateral lobe, hence has this much better developed than adult specimens. Comments: This new species appears to be not uncommon in the northwestern Dominican Republic and is known to feed on “Palo de Tabaco” (Pictetia spinifolia, Fabaceae) at Janico. Eggs unknown. Distribution (Fig. 380): Hispaniola, NW-Dominican Republic (Santiago Province) [USNM, ZMUH]. Number of specimens examined: 15 TABLE 21. Measurements of Haplopus brachypterus n. sp. [mm] ♂, HT [USNM]

♂♂, PT

♀♀, PT

-

-

131.0–139.0

Body

87.5

83.2–86.5

116.5–123.5

Pronotum

3.8

3.5–3.7

5.0–5.1

Mesonotum

19.3

18.2–18.9

25.8–27.0

Metanotum

5.0

4.9–5.1

6.6.6.7

Median segment

6.7

6.3–6.5

7.8–8.2

Tegmina

6.1

5.8–6.5

8.5.–8.8

Alae

6.0

5.4–5.9

7.7–8.0

Profemora

19.0

17.9–18.4

18.8–20.0

Mesofemora

17.2

15.4–16.8

16.6–18.4

Metafemora

21.0

19.8–21.4

21.0–24.5

Protibiae

18.9

17.6–18.8

18.9–21.5

Mesotibiae

16.5

14.6–15.7

17.6–19.0

Metatibiae

21.8

20.2–21.6

22.9–26.3

Antennae

> 60.0

> 58.0

49.0–53.0


Haplopus intermedius n. sp. (Figs. 211–219, 359, 380) HT, ♂: Dominican Republic, RD-158, ~ 2 km N Maizal, Valverde Prov., dry forest, 19°39.819’N 70°58.422’W, 23.VII.2003, D. Perez, R. Bastardo, B. Hierro. (day) [USNM]. PT, ♀: Dominican Republic, RD-088 4 km E Montecristi, Montecristi Prov., 19°49.657’N 71°37254’W, 10.XII.2002, D. Perez, R. Bastardo (night) [USNM]. PT, ♂ (nymph n5): Dominican Republic, RD-088 4 km E Montecristi, Montecristi Prov., 19°49.657’N 71°37254’W, 10.XII.2002, D. Perez, R. Bastardo (night) [USNM]. PT, ♂ (nymph n4): Dominican Republic, RD-028 5.II.02, 3 km S. Montecristi, near road, Montecristi Prov., near sea level.Day. RB, BH, DP [USNM].

Diagnosis: Very similar to H. bicuspidatus de Haan, 1842 and H. brachypterus n. sp. but well distinguished from both species by the triangular sub-apical tooth on the dorsal carinae of the meso- and metafemora of both sexes, as well as the sub-basal and sub-apical elevation of the anterodorsal carina of the meso- and metatibiae of ♀♀ and basally compressed and dorsally carinate cerci of ♂♂. From the first species ♀♀ also differ by: the presence of posterior tubercles on the pronotum and tubercles on the profurcasternum; slightly shorter mesothorax; considerably larger epiproct (Fig. 215) and distinct paired posterior spines on abdominal tergites II–IV (Fig. 218). Males readily differ from those of H. bicuspidatus by the shortened alae (length 5.6 mm) and the posterolateral tooth or lobe of abdominal tergum VII (Fig. 217). From H. brachypterus n. sp. this species can be distinguished by: the distinct posterolateral lobe of abdominal tergum VII and sub-apical tooth on the dorsal carinae of the meso- and metafemora of both sexes. Females furthermore differ by: the slightly larger dimensions; larger horns of the vertex; REVISION OF HAPLOPODINI


117

relatively shorter mesothorax; presence of a sub-basal and sub-apical elevation of the anterodorsal carina of the meso- and metatibiae, and larger epiproct (Fig. 215). Males have the anal region of the alae larger and better developed than those of H. brachypterus n. sp.. The short alae of ♂♂ also resemble H. scabricollis (Gray, 1835) from the Bahamas, Florida Keys, Navassa Island and the Cayman Islands. They however clearly differ from ♂♂ of H. scabricollis by: the more slender body; lack of longitudinal white stripes along the lateral margins of the pronotum and white median markings on the median segment and abdominal tergites VIII–X; convex and prominently bi-cornute vertex; more decidedly excavated anal segment (Fig. 217) and more acute central spine of the poculum (Fig. 216). ♀♀ are well distinguished by: the globose and prominently bi-cornute vertex, longer alae and typical armature of the mid and hind legs. Etymology: The specific name “intermedius” (lat. = inbetween) is meant to refer to the close relation to the genus Cephaloplopus n. gen. in several morphological aspects. Description: ♀ (Fig. 211). Rather large (body length including the subgenital plate 152.0 mm) and slender for the genus; body surface slightly glabrous. Metathorax and great parts of abdomen discoloured to very dark brown during the process of dehydration in the unique ♀ paratype. Colour creamish mid brown and all over furnished with pale cream to whitish mottling and speckles. Head with lateral surfaces whitish and two small black markings above the eyes; vertex with irregular bown markings. Antennae mid grey dorsally, brown ventrally. Eyes dark reddish brown. Spines of the thorax dark sepia with the points black. Tegmina and costal region of alae dark brown with the venations of a slightly paler colour; the latter black basally. Anal region of alae transparent drab and all major longitudinal and transverse veins broadly marked with dark brown (Fig. 219). Legs and subgenital almost entirely creamish white with faint grey mottling. Head: About 1.2x longer than wide, globose with the cheeks fairly convex. Vertex rounded and armed with a pair of large apically pointed cephalad horns. Eyes circular and contained almost 3.5x in length of cheeks. Antennae broken in the only specimen available. Scapus 2.2x longer than wide with the lateral margins gently rounded. Pedicellus less than half the length of scapus and a little shorter than antennomere III. Both antennae broken in the only specimen available. Thorax: Pronotum longer but considerably narrower than head, almost 2x longer than wide, roughly rectangular and with the lateral margins gently emarginated medially. Transverse median sulcus deep, almost straight and not reaching lateral margins of segment. Dorsal surface with a pair of moderately sized but pointed spines in the anterior portion; in posterior portion with two pairs of smaller spines. Prosternum smooth. Profurcasternum with three pairs of small tubercles, one pair marking the anterolateral angles of the segment, a much smaller pair at the posterior angles and a further even smaller pair positioned posterior of the distinct anterior tubercles. Mesothorax about 2.7x longer than head and pronotum combined. Mesonotum almost parallel-sided, the surface with a good number of irregularly disposed spines of variable size; along lateral margins with a marginal row of about eight small but pointed spines. Meso- and metapleurae armed with an irregular longitudinal marginal row of moderately sized spines. Mesosternum with four pairs of low spines, metasternum with two pairs of blunt spines. Metanotum about ¼ the length of mesonotum, unarmed. Tegmina oval, coriaceous, with the venation very distinct, dense and irregularly disposed; slightly projecting over posterior margin of metanotum. The median protuberance very shallow and conspicuously displaced towards the apex of tegmen. Alae a little longer than tegmina and reaching about 2/3 the way along median segment. Abdomen: Median segment 2x longer than wide and very gently narrowed medially. All segments unarmed, except for a pair of compressed posteromedian spines on tergites II–IV; this pair of spines most prominent on II (Fig. 218). Segments II–IV very slightly increasing in length, V as long as IV; the latter two segments about 1.8x longer than wide. VI a little shorter and narrower than V, about 2x longer than wide. Tergum VII slightly shorter than VI and parallel-sided, with the posterolateral angles protuded into a rather narrow, blunt triangular lobe (Fig. 214). VIII slightly shorter than VII, gently narrowed medially and roughly 2.5x longer than wide. IX rectangular and slightly more than half the length of VIII. Anal segment with a very faint longitudinal median carina, narrowed in posterior half and with a deep triangular posteromedian emargination. Epiproct prominent, acutely triangular and projecting considerably over apex of anal segment (Fig. 215). Cerci small, conical with a rather acute tip and slightly compressed laterally; somewhat projecting over posterior margin of anal segment. Subgenital plate very long, lanceolate, carinate longitudinally and with a fairly acute apex; extending over apex of abdomen by more than the combined length of tergites VIII–X (Figs. 213–214).


HENNEMANN ET AL.

FIGURES 211–219. Haplopus intermedius n. sp.. 211. ♀ PT: Dominican Republic, Montecristi Province, nr. Montecristi [USNM]; 212. ♂ HT: Dominican Republic, Valverde Province, nr. Maizal [USNM]; 213. Apex of abdomen of ♀ PT (lateral view) [USNM]; 214. Apex of abdomen of ♀ PT (dorsal view) [USNM]; 215. Anal segment and epiproct of ♀ PT (dorsal view) [USNM]; 216. Apex of abdomen of ♂ HT (lateral view) [USNM]; 217. Apex of abdomen of ♂ HT (dorsal view) [USNM]; 218. Abdominal tergites II and II of ♀ PT showing the paired spiniform posterior projections (dorsal view) [USNM]; 219. Right ala of ♀ PT [USNM].

Legs: Profemora about 3/5 the length of mesothorax, mesofemora slightly projecting over median segment and metafemora reaching about 2/3 the way along abdominal segment IV. Profemora occasionally with three very minute spines in the apical half of the medioventral carina, otherwise unarmed. Anteroventral carina of meso- and metafemora with two, posteroventral carina with one sub-apical spine; medioventral carina armed with four prominent and strong spines. Both dorsal carinae with a triangular lobe-like tooth sub-apically. Anterodorsal carina



119

of meso- and metatibiae with roundly triangular elevation sub-basally and sub-apically. Basitarsi about as long as following three tarsomeres combined. ♂ (Fig. 212): Of moderate size (body length 82.5 mm) and rather slender for the genus with conspicuously shortened alae (length 5.6 mm). Great parts of head, mesometapleurae, ventral body surface and legs mid green. Head green with a faint yellow marking on the frons and a fine yellow postocular stripe. Dorsal surface of thorax and abdomen creamish mid brown. Dorsal spines of pro- and mesothorax mid brown with black points. Meso- and metapleurae with a very faint longitudinal yellowish marking in posterior portion and each with a dull orange longitudinal stripe along lower margin. Lateral margins of abdominal tergites VIII and IX broadly white. Tegmina and costal region of alae greyish mid brown and becoming darker towards the base; anterior margin and basal portion of posterior margin of tegmina, as well as anterior margin of alae broadly white; base of alae black. Anal region of alae pink with all major veins brown. Antennae greyish mid brown. Tarsi mid brown. Head: Generally as in ♀♀ but eyes more prominent, projecting hemispherically and their length contained only about 2x in that of cheeks. Antennae moderately robust and reaching to abdominal segment VI; with 61 segments. Thorax: Pronotum slightly longer but narrower than head, general shape as in ♀♀ but relatively shorter and anterior margin slightly broader than posterior margin; surface smooth except for a moderate pair of spines in the anterior portion. Mesothorax about 2.7x longer than head and pronotum combined. Mesonotum in the HT with four paired spines at anterior margin and five further pointed spines in the median portion. Mesosternum with seven and metasternum with two pairs of spiniform tubercles. Tegmina oval and very slightly projecting over posterior margin of metanotum, central protuberance very shallow. Alae small, a little shorter than alae and hardly reaching 2/3 the way along median segment; anal region well developed. Abdomen: Segments II a little shorter than III–IV and equal in length to V, about 3.2x longer than wide. III and IV longest segments and almost 4x longer than wide. V–VII gradually decreasing in length with VII no more than 2.2x longer than wide. All tergites and sternites smooth. VII with a distinct, roundly triangular posterolateral tooth (Fig. 187). VIII slightly shorter than VII and very gently widening towards the posterior; IX about ¾ the length of VIII and narrowed towards the posterior. Anal segment with a very indistinct longitudinal median carina in basal portion and posterior margin with a distinct triangular median emargination (Fig. 217), slightly swollen and on ventral surface armed with several small, black in-curving denticles. Epiproct very small and roughly triangular (Fig. 217). Vomer with a broad, roughly semi-circular base and a very long, papillate, up-curving terminal hook (Fig. 359). Cerci large and about equal in length to anal segment; distinctly laterally compressed in basal portion and slightly longitudinally carinate dorsally (Figs. 187 & 306). Poculum moderately convex, cup-like and with a conical basal hump (Fig. 216); posterior portion with a distinct longitudinal median keel. Legs: Pro- and mesofemora about 2/3 the length of mesothorax, metafemora projecting a little over posterior margin of abdominal segment IV. All legs less carinate than in ♀♀ but armature generally alike, except for the dorsal sub-apical teeth of the meso- and metafemora less distinct and dorsal elevations of the meso- and metatibiae lacking. Tarsi relatively more elongate and basitarsi a little longer than following three tarsomeres combined. Nymphs: The two half-grown ♂ nymphs at hand are of a greyish brown colour with pale mottling and already show the characteristic features that distinguish this new species from H. brachypterus n. sp., having the sub-apical tooth on the dorsal carinae of the meso- and metafemora and the posterolateral lobe of abdominal tergum VII well developed. Comments: This pretty new species emphasizes the close relation between Haplopus Burmeister, 1838 and Cephaloplopus n. gen. by ♂♂ having basally compressed and slightly carinate cerci (Figs. 217, 351), a character otherwise unusual for Haplopus. Eggs unknown. Distribution (Fig. 380): Hispaniola, NW-Dominican Republic (Valverde Prov. & Montecristi Prov.) [USNM]. Number of specimens examined: 4 TABLE 22. Measurements of Haplopus intermedius n. sp. [mm]


♂, HT [USNM]

♀, PT [USNM]

-

152.0

82.5

133.0 ......continued on the next page


HENNEMANN ET AL.

TABLE 22. (Continued) ♂, HT [USNM]

♀, PT [USNM]

Pronotum

3.4

6.3

Mesonotum

17.7

30.0

Metanotum

4.8

7.1

Median segment

6.3

9.2

Tegmina

5.9

10.0

Alae

5.6

9.8

Profemora

18.1

22.3

Mesofemora

16.2

19.5

Metafemora

20.5

25.3

Protibiae

17.8

21.0

Mesotibiae

15.4

18.9

Metatibiae

21.4

25.1

Antennae

ca. 63.0

> 34.0

Haplopus micropterus (St. Fargeau & Audinet-Serville, 1828) (Figs. 220–256, 340, 360, 380, 399–405) Cyphocrana microptera St. Fargeau & Audinet-Serville, 1828: 445. HT, ♀: St. Thomas [RMNH]. [Replacement name for Phasma angulata Stoll, 1813; not Mantis angulata Fabricius, 1793] Aplopus micropterus, Gray, 1835: 34. Rehn, 1903: 136. Wolcott, 1923: 23. Wolcott, 1936: 35. Wolcott, 1951: 50. Diapherodes micropterus, Moxey, 1972: 107 (in litt.; in part). Haplopus micropterus, Westwood, 1859: 87. Kirby, 1904a: 363. Redtenbacher, 1908: 431. Otte & Brock, 2005: 152. Phasma (Haplopus) micropterum, de Haan, 1842: 128. Phasma angulata Palisot de Beauvois, 1805: 166, pl. 14: 4 (♀). HT, ♀: St. Domingo [presumed lost]. [Junior homonym of Mantis angulata Fabricius, 1793] Phasma angulata Stoll, 1813: 61, pl. 21: 77 (♀). HT, ♀: St. Thomas [RMNH]. [Junior homonym of Mantis angulata Fabricius, 1793] Haplopus angulatus, Burmeister, 1838: 577. Haplopus bituberculatum de Haan, 1842: 128. HT, ♀: Bituberculatum de Haan, ?; RMNH Leiden; TYPE, Haplopus bituberculatum de Haan 1842; RMNH Holotype [RMNH]. n. syn. Kirby, 1904a: 364. Redtenbacher, 1908: 433. Aplopus bituberculatus, Bragg, 1996: 109. Bragg, 2001: 706. Haplopus bituberculatus, Otte & Brock, 2005: 151. Haplopus cytherea Westwood, 1859: 86, pl. 18: 5, 5a & b (♂). LT, ♂ (by present designation): St. Dom.; E Coll (1870-73) W. W. Saunders, Purchased and pres. ’73 by Mrs. F. W. Hope; Type ♂– Westwood, Haplopus cythereus Westw., Cat. Phasm. 1859, p. 86, pl. 18 f. 5; Type Orth: 617, Haplopus cythereus Westw., Hope Dept. Oxford; Haplopus cythereus Westwood, Lectotype, det. Brock, 1996 [OXUM, No. 617]; PLT, ♂: St. Dom.; Cytherea W.; Type ♂—Westwood, Haplopus cythereus Westw., Cat. Phasm. 1859, p. 86, pl. 18 f. 5; Type Orth: 617, Haplopus cythereus Westw., Hope Dept. Oxford; Haplopus cythereus Westwood; Paralectotype, det. Brock, 1996 [OXUM, No. 617]. n. syn.



121

Kirby, 1904a: 363. Moxey, 1972: 114 (in litt.). [Listed as a synonym of Diapherodes spinipes Gray, 1835] Haplopus cythereus, Redtenbacher, 1908: 432. Otte & Brock, 2005: 151. Aplopus cytherea, Rehn, 1904: 63. [Description of ♂] Haplopus ligia Westwood, 1859: 89, pl. 11: 1, 1a, 1b (♂) & 2, 2a (♀). LT, ♂ (by present designation): St. Dom. 55.1, Haplopus ligia Westw. pl. 11 fig. 1, Westw. ♂, Haplopus ligia Westwood, ST [NHMUK]; PLT, ♂: St. Dom. 55.1 [NHMUK]; PLT, ♀: St. Dom. 55.1, Haplopus Ligia Westw. (St. Domingo) [NHMUK]; PLT, ♀: St. Dom. 55.1, Haplopus ligia Westw. pl. 11 fig. 2, Westw. ♀, Haplopus ligia Westwood, ST [NHMUK]. n. syn. Brock et al., (in press) Kirby, 1904a: 364. Redtenbacher, 1908: 432. Moxey, 1972: 114 (in litt.). Otte & Brock, 2005: 151. Haplopus ligiolus Redtenbacher, 1908: 432. LT, ♂ (by present designation): 104; Insel Mona, zw. Haiti u. Porto Rico, W. Bock leg. ded. 12.IX.1892; PHA 22, Zoologisches Museum Hamburg [ZMUH]; PLT, ♀: 105; Ins. Mona b. Hayti, C. Bock leg. ded. 16.V.184; cf. Stoll Taf. 21 fig. 17 Spectre à ailes petiter; PHA 26, Zoologisches Museum Hamburg [ZMUH]; PLT, ♀: 106; 10; Ins. Mona b. Hayti, C. Bock leg. ded 16.V.1894; PHA 27, Zoologisches Museum Hamburg [ZMUH]; PLT, ♀: “Westindien” [NHMB, No. VI.D.18]. n. syn. Werner, 1929: 7, fig. A (♂). Moxey, 1972: 107 (in litt.). [Listed as a synonym of H. micropterus (St. Fargeau & Audinet-Serville, 1825)] Hennemann & Conle, 1999: 11. Otte & Brock, 2005: 151. Haplopus obtusus Redtenbacher, 1908: 431, pl. 19: 5 (♀). HT, ♀: Coll. Br. v. W., ex. coll. Sommer, Sta. Kruz, det. Br. v. W., Haplopus obtusus, 7646, 7646, P. angulata Fabr., P. gigas Drury, Sta. Crux [NHMW, No. 835]. n. syn. Moxey, 1972: 107 (in litt.). [As a synonym of H. micropterus (St. Fargeau & Audinet-Serville, 1825)] Aplopus obtusus, Brock, 1998a: 47. Haplopus obtusus, Otte & Brock, 2005: 152. Diapherodes spinipes Gray, 1835: 34. HT, ♀: no data [not traced—presumed lost]. [Replacement name for Phasma angulata Palisot de Beauvois, 1805: 166; not Fabricius, 1793, not Stoll, 1813] n. syn. Diapherodes spinipes, Moxey, 1972: 114 (in litt.). Haplopus spinipes, Westwood, 1859: 87. Kirby, 1904a: 363. Redtenbacher, 1908: 431. Otte & Brock, 2005: 152. Aplopus sp., Wolcott, 1941: 40. Ramos, 1946: 9. Wolcott, 1948: 50. Wolcott, 1951: 50. [Not: Phasma angulata Fabricius, 1793: 13, = Diapherodes angulata angulata (Fabricius, 1793)] [Not: Phasma angulata Fabricius, 1793: 13, = Diapherodes angulata angulata (Fabricius, 1793)] Further material [86 ♂♂, 111 ♀♀, 51 nymphs, eggs]: VIRGIN ISLANDS (ST. CROIX): 1 ♀: Annaly St. Croix, Virgin Is., on grass in field, Oct. 15,59, C.-W. Miskimen, 60 4291; USMNH; w0195; Diapherodes micropterus (St. F. & Serv.) det. C.F. Moxey 1972 [USNM]. VIRGIN ISLANDS (ST. THOMAS): 1 ♀: 806; micropterus Gray, Westw.*, Cyphocr. microptera Serv., angulata, St. Thomas, Moritz [MNHU]; 1 ♀: St. Thomas, W.-Ind., C. Calwood leg., ded. 28.XII.1895; PHA 25, Zoologisches Museum Hamburg [ZMUH]. VIRGIN ISLANDS (ST. JOHN): 1 ♀, 1 ♀ (penultimate instar): St. John [MNHU]. PUERTO RICO: 1 ♂: Guánica, P.R., 2.Oct.1937, W. García; USMNH; w0199 [USNM]; 1 ♀: Rincon, P.R., Nov. 1938, Coll. S.R. Sada; USMNH; w0196; Diapherodes micropterus (St. F. & Serv.) det. C.F. Moxey 1972 [USNM]; 2 ♀♀, 5 ♂♂, eggs: ex Zucht F. Hennemann, Herkunft: Puerto Rico, VI.1995 [coll. FH, No’s 0243-20 to 26 & E3]. MONA ISLAND: 1 ♂: Mona Island, Aug 10.1938, Coll. A. Palmer; USMNH; w0198; Diapherodes micropterus (St. F. & Serv.) det. C.F. Moxey 1972 [USNM]. HISPANIOLA (DOMINICAN REPUBLIC): 2 ♂♂: Dominican Republic, Isla Beata, 27.VII.1977, ? Eugenio Marcano, NHMUK(E) 2005-98, ex. Museo Nacional de Historia Natural Sto. Domingo [NHMUK]; 1 ♂: Dominican Republic, Cerretera Romana—Bayahibe, 19.IX.1994, Kelvin


HENNEMANN ET AL.

A. Guerrero, NHMUK(E) 2005-98 [NHMUK]; 1 ♂: Dominican Republic, Punta Villa Mella, Distrito Nacional, 3.X.1993, Kelvin A. Guerrero, NHMUK(E) 2005-98 [NHMUK]; 3 ♂♂, 2 ♀♀, 4 eggs: Dominican Republic, RD-052 Pueblo Nuevo, Baní Peravia Prov., 27.VII.2002, 97 m (400 ft.), 18°17.757’N 70°19.601’W, D. Perez, R. Bastardo [USNM]; 3 ♂♂, 1 ♀, 1 ♂ (nymph n4): Dominican Republic, RD-062 ~ 5 km W Las Américas, Santo Domingo, 18°28.158’N 69°43.601’W, 19.XI.2002, D. Perez, R. Bastardo, B. Hierro [USNM]; 2 ♂♂, 3 ♀♀, 2 ♀♀ (penultimate instar), 4 ♀♀ (nymph n4), 2 ♀♀ (nymphs n3), 1 ♀ (nymph n2), 1 ♂ (penultimate inatar): Dominican Republic, RD-060 Monte Rio, ~ 70 m, dry scrub, 18°22.972’N 70°43.036’W, 17.XI.2002, D. Perez, R. Bastardo, B. Hierro. (night) [USNM]; 1 ♀: Dominican Republic, RD-091, Sierra Prieta, Santo Domingo Prov., 133 m, 18°39.010’N 69°58.409’W, 18.III.2003, D. Perez, R. Bastardo, B. Hierro (night) [USNM]; 2 ♂♂, 2 ♀♀, 1 egg: Dominican Republic, RD-034 Boca de la Cañada, ~ 15 km S Oviedo, Pedernales Prov., 4.VII.2002, 17°54.901’n 71°30.067’W, B. Hierro, R. Bastardo, D. Perez [USNM]; 1 ♀, 1 ♀ (nymph n3), 7 eggs: Dominican Republic, RD-153 La Poza de Agua Nueva, El Curro, Sierra Martín Garcia, Azua Prov., 18°18.324’N 70°57.176’W, ~ 800 m, 15–16.VII.2003, D. Perez, R. Bastardo, B. Hierro. (day/night) [USNM]; 1 ♀, 3 eggs: Dominican Republic, RD-146 ~ 2 km S Pinosdel Edén, Sierra de Neiba, Independencia Prov., 18°34.758’n 71°45.804’w, 8.VII.2003, D. Perez, R. Bastardo, B. Hierro. (night) [USNM]; 1 ♀ (penultimate instar): Dominican Republic, RD-079 Isla Beata, trail to Cueva del Tubo, PNJ, 17°36.844’N 71°31.379’W, 3.XII.2002, Dperez, Bhierro, Rbastardo. (day/night). [USNM]; 1 ♀ (nymph n5): Dominican Republic, San Cristóbal Prov., Sabana de Hatillo, on Malpighia gossipifolia (cerazas), 13.XII.1994, D. E. Perez [USNM]; 1 ♂ (penultimate instar): Dominican Republic, RD-069 Km 54 Rd. Azua-Barahona, Barahona Prov., 70 m, 18°21.871’N 71°09.080’W, 24.XI,2002, D. Perez, B. Hierro, H. Andújar. (night). [USNM]; 1 ♂ (nymph n5): Dominican Republic, RD-156 La Furnia, Berreras, Azua Prov., 18°19.289’N 70°54.755’W, 18.VII.2003, D. Perez, Rbastardo, Bhierro (n) [USNM]; 1 ♀ (n4): Dominican Republic, RD-063 Sierra Prieta, Santo Domingo Prov., 18°38.315’N 69°58.302’W, 20.XI.2002, D. Perez, R. Bastardo, B. Hierro [USNM]; 1 ♀ (n4): Dominican Republic, RD025 1–2.II.02 km 10 on Rd. to Los Anones, Ocoa Prov., 1070 m, 347–605 mE 2052–511 mN. RB, BH, DP [USNM]; 1 ♀ (n3): Dominican Republic RD-210 Mirador del Hoyo de Pelempito, P. N. Sierra de Bahoruco, Pedernales Prov., 1.250 m, 18°05.396’N 71°30.663’W, 5.IV.2004, D. Perez, R. Bastardo, B. Hierro (d/n) [USNM]; 1 ♂: El Choco, Puerto Plata 2-IV2000 en Bidens cynapiifolia R. H. Bastardo [USNM]; 4 ♀♀: Dominican Republic, 200 m N Puente Coronel Barahona Prov., 22.ix.2000 D. Perez, R. Bastardo, B. Hierro [USNM]; 1 ♀: Dominican Republic, La Altagracia Prov., Bayahibe, 2001, D. E. Perez [USNM]; 1 ♀: Dominican Republic, RD-219 Sierra Prieta, Villa Mella, Santo Domingo Prov., 142 m, 18º38.925’N 69º58.303’W, 12.iv.2004, D. Perez, B. Hierro, R. Bastardo. (n) [USNM]; 2 ♂♂ (nymphs): Dominican Republic, RD-153 La Poza de Agua Nueva, El Curro, Sierra Martín García, Azua Prov., 18°18.324’N 70°57.176’W, ~800 m, 15–16.vii.2003, D. Perez, R. Bastardo, B. Hierro. (day/night) [USNM]; 1 ♂: Dominican Republic, Azua Prov., Rd. Monte Rio—Puerto Viejo, 18°23.570’N 70°42.758’W, 120 m, 15.viii.2006, D. Perez, R. Bastardo, B. Hierro, S. Medrano (night) [USNM]; 1 ♂, 1 ♀ (nymphs): Dominican Republic, RD-272 Caseta 1, P N Sierra de Bahoruco, Independencia prov., 18º16.038’N 71º32.691’W, 1,239 m, 14.vii.2004, D. Perez (d) [USNM]; 1 ♂: Dominican Republic, RD-276 Boca de Yuma, P N del Este, La Altagracia prov., 18º19.554’N 68º48.503’W, near sea level, 19–20.vii.2004, D. Perez (d/n) [USNM]; 1 ♂: Dominican Republic, RD-092 Blanco (near hydroelectric), Bonao, Monseñor Nouel Prov., 18°52.946’N 70°30.337’W, 19.iii.2003, D. Perez, R. Bastardo, B. Hierro. (night) [USNM]; 1 ♂: Dominican Republic, RD-103 Rd. to Majagual, Sierra de Neiba, Bahoruco Prov., 669 m, 18°32.340’N 71°18.118’W, 25.iii.2003, D. Perez, R. Bastardo, B. Hierro. (night) [USNM]; 1 ♀ (nymph): Dominican Republic, RD-069 km 54 Rd. Azua—Barahona, Barahona Prov., 70 m, 18°21.871’N 71°09.080’W, 24.xi.2002, D. Perez, B. Hierro, H. Andujar. (night) [USNM]; 3 ♂♂: Dominican Republic, RD-079 Isla Beata, on way to Cueva del Tubo, Parque Nacional Jaragua, nr. sea level, 17°36.844’N 71°31.379’W, 3.xii.2002, D. Perez, B. Hierro, R. Bastardo. (night) [USNM]; 1 ♂ (nymph): Dominican Republic, RD-182 Loma Quita Espuela, Firme de loma, S. F. de Macorís Prov., 19º21.101’N 70º08.930’W, 715 m, 3–4.xii.2003, D. Perez, R. Bastardo, A. Marmolejos (day/night) [USNM]; 1 ♀: Dominican Republic RD-171 El Maizal, near Presa de Valdesia, Peravia Prov., 18°22.942’N 70°16.859’W, 97 m, 31.vii.2003, D. Perez, R. Bastardo, B. Hierro. (day) [USNM]; 2 ♀♀: Dominican Republi,c RD-212 ̴150 m N bridge on road Cabo Rojo—Aceitillar, Pedernales prov., 16 m, 17º58.530’N 71º39.034’W, 7.iv.2004, D. Perez, B. Hierro, R. Bastardo. (d/n) [USNM]; 2 ♂♂, 4 ♂♂ (nymphs), 1 ♀ (nymph): Dominican Republic, La Altagracia Prov., Bayahibe, 26/ix/1999 (night) D. Perez, R. Bastardo, L. Ramos. [USNM]; 1 ♂ (nymph): Dominican Republic, Jánico, Santiago Prov., 30 ix 1996, on Pictetia spinifolia (Tabacuelo) D. Perez, S. Navarro [USNM]; 1 ♂ (nymph): Dominican Republic, RD-182 Loma Quita Espuela, Firme de loma, S. F. de Macorís Prov., 19º21.101’N 70º08.930’W, 715 m, 3–4.xii.2003, D. Perez, R. Bastardo, A. Marmolejos (day/night) [USNM]; 2 ♂♂, 1 ♀: RD: La Altagracia Higüey Punta Cana 20–28-vii-2000 R. Bastardo [USNM]; 1 ♂: Dominican Republic, RD-005 15.i.02 Las Yayitas, Azua Prov., 240 m, 316–230 mE 2046–941 mN. D. Otte, D. Perez, R. Bastardo, S. Medrano [USNM]; 1 ♂: Dominican Republic, RD-056 Pueblo Nuevo, Baní, Peravia Prov., dry scrub, 18°17.672’N 70°19.922’W, 16.xi.2002, D. Perez, R. Bastardo, B. Hierro. (night) [USNM]; 1 ♀: Dominican Republic, RD-027 2.ii.02 Km 7 on Rd. to Presa Jiguey, Ocoa Prov., 860 m, 345–778 mE 2053–018 mN. Night collection. D. Perez, R. Bastardo, B. Hierro [USNM]; 1 ♀: Dominican Republic, RD-156 La Furnia, Barreras, Azua Prov., 18°19.289’N 70°54.755’W, 18.vii.2003, D. Perez, R. Bastardo, B. Hierro. (night) [USNM]; 1 ♀ (nmyph): Dominican Republic, RD-062 ~5 km W of Las Americas Airport, nr. LADOM, Santo Domingo Prov., 18°28.158’N 69°43.601’W, 19.xi.2002, D. Perez, R. Bastardo, B. Hierro. (night) [USNM]; 3 ♀♀: Dominican Republic, RD-052 Pueblo Nuevo, Baní, Peravia Prov., 27.vii.2002, 97 m (400 ft.), 18º17.757’N 70º19.601’W, D. Perez, R. Bastardo [USNM]; 1 ♂: Dominican Republic, RD-016 19.i.02 Km 26 Rd. Cabo Rojo- Aceitillar, Pedernales Prov., 680 m, 222–642 mE 2004–651 mN, transition forest. D. Otte, D. Perez, R. Bastardo, S.



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Medrano, B. Hierro [USNM]; 1 ♂ (nymph): Dominican Republic, RD-041 Sierra Prieta, Villa Mella, Santo Domingo Prov., 18º38.939’N 69º58.373’W, 8.vii.2002, 500 ft., 8.vii.2002. D. Perez, B. Hierro, R. Bastardo, S. Medrano. [USNM]; 1 ♂: Dominican Republic, RD-019 22.i.02 Talanquera, San Pedro de Macoris Prov., disturbed scrub forest, about 20 m above sea level, 18º26.35’N 69º24.06’W, D. Otte, D. Perez [USNM]; 1 ♂: Dominican Republic, RD-060 Monte Rio, ̴4 km W of beach, ~70m, dry scrub, 18°22.972’N 70°43.036’W, 17.xi.2002, D. Perez, R. Bastardo, B. Hierro. (night) [USNM]; 2 ♀♀: Dominican Republic, RD-034 Boca de la Cañada, ~15 km S Oviedo, Pedernales Prov., 4.vii.2002, 17º54.901’N 71º30.067’W, B. Hierro, R. Bastardo, D. Perez. [USNM]; 4 ♂♂, 3 ♀♀: Dominican Republic, RD-199 Boca de Yuma, P. N. Del Este, La Altagracia Prov., 20 m, 18º21.875’N 68º37.081’W, 16–17.xii.2003, D. Perez, R. Bastardo (day/night). [USNM]; 2 ♂♂, 3 ♀♀: Dominican Republic, RD-055 ~2 km N Bayahibe, La Altagracia Prov., 31.vii.2002, 18º23.423’N 68º50.453’W, D. Perez, R. Bastardo, B. Hierro [USNM]; 1 ♂: Dominican Republic, RD-067 Laguna Grande, Las Lagunas, Padre Las Casas, Azua Prov., 18°48.088’N 70°52.842’W, 23.xi.2002, 1,033 m, D. Perez, B. Hierro, H. Andujar. (day) [USNM]; 4 ♂♂ (nymphs), 4 ♀♀ (nymphs): Dominican Republic, Pedernales, Lado Fundacipe, Pedernales Prov., 18°01.8’N 71°44.7’W, 19–20 iii 1999, D. E. Perez, S. Navarro [USNM]; 1 ♂, 1 ♀, 1 ♂ (nymph): Dominican Republic, Paraiso, Barahona Prov. 17°59.1’N 71°10’W 21 iii 1999 D. Perez, S. Navarro [USNM]; 1 ♂: Dominican Republic, Punta Cana Beach Resort, Reserva Ecologica Punta Cana, La Altagracia Prov., 23/xii/2008 D. E. Perez [USNM]; 1 ♂, 5 ♀♀, 1 ♂ (nymph), 1 ♀ (nymph): Dominican Republic, San Cristobal Prov., Sabana de Hatillo, on Malpighia gossipifolia (cerezas), 25.ix.1996, D. E. Perez [USNM]; 2 ♂♂, 1 ♀, 1 ♀ (nymph): Dominican Republic, RD-195 ~6 km S of Cabral, dry forest, Barahona Prov., 18º12.252’N 71º14.401’W, 247 m, 13.xii.2004, D. Perez, R. Bastardo, B. Hierro (night). [USNM]; 1 ♂, 5 ♀♀: Dominican Republic, RD-196 ~4 km S of Cabral, Barahona Prov., 18º13.835’N 71º14.251’W, 105 m, 14.xii.2003, D. Perez, R. Bastardo, B. Hierro (night). [USNM]; 7 ♂♂, 5 ♀♀, 1 ♀ (nymph): Dominican Republic, RD-191 Around Caseta No. 1, Parque Nacional Sierra de Bahoruco, 1,239 m, Independencia Prov., 18º16.038’N 71º32.691’W, 11–12.xii.2003, D. Perez, R. Bastardo, B. Hierro. (day/night) [USNM]; 1 ♂: Dominican Republic, Prov. Barahona, 1 km from Entrada to Vincente Noble, R.E. Woodruff, coll. 30-IV-78 [FSCA]; 1 ♀: Dominican Republic, WI Baoruco Mts., 1250 m, 11. Sept. 1973, Coll. #3, TJ Wakler, JC Schuster [FSCA]; 1 ♀: Dominican Republic, Pedernales Prov., N of Cabo Rojo, 18.IX.1973, JC Schuster [FSCA]; 2 ♂♂: Dominican Republic, Pr. Pedernales, Km 12 N. Cabo Rojo, 21-vi-1999 R. E. Woodruff, Prosopis at night [FSCA]; 1 ♂, 1 ♂ (nymph): Dominican Republic, 15 km N Cabo Rojo, 21.vi.1999 R. E. Woodruff & R. M. Baranowski [FSCA]; 1 ♂: Dominican Republic, Isla Saona, 6.vii.2002 C. Nuñez [MNHNSD]; 1 ♀: Dominican Republic, RD-063 Sierra Prieta, Santo Domingo Prov., 18°38.315’N 69°58.302’W, 20.XI.2002, D. Perez, R. Bastardo, B. Hierro [MNHNSD]; 3 ♀♀, 2 ♂♂: Dominican Republic, RD-065 Monte Rio, ~4.5 km W of beach, Azua Prov., dry scrub, 18°23.728’N 70°42.726’W, 22.xi.2002, D. Perez, B. Hierro, H. Andujar. (night) [MNHNSD]; 1 ♂, 1 ♀: Dominican Republic, RD-078 El Cajuil, Oviedo, Pedernales Prov., 52 m, 17°48.783’N 71°21.538’W, 2.xii.2002, D. Perez, B. Hierro, R. Bastardo. (night) [MNHNSD]; 1 ♀, 1 ♀ (nymph): Dominican Republic, RD-083 Juan Dolio, San Pedro de Macorís Prov., sea level, nr.18°26.35’N 69°24.06’W, 6.xii.2002, D. Perez, R. Bastardo. (night) [MNHNSD]; 1 ♂, 1 ♀ (nymph): Dominican Republic, RD-084 ~10 km S of Ocoa, nr. main road, Ocoa Prov., 468 m, 18°28.350’N 70°29.670’W, 7.xii.2002, D. Perez, R. Bastardo. (night) [MNHNSD]; 1 ♀: Rep. Dom., Prov. Santo Domingo, Distrito Nacional, San Juan Bosco #63, 16.VIII.2005, R. Gonzáles y M. Díaz [MNHNSD]; 1 ♀: Rep. Dom., Prov. Santo Domingo Este, Boca Chica, La Malena, 20.X.1979, Marcano y Domínguez [MNHNSD]; 1 ♀: Rep. Dom., Prov. San Cristóbal, Villa Altagrica, La Cumbre, 14.VI.2001, C. Núñez [MNHNSD]; 1 ♀ (penultimate instar): Rep. Dom., Prov. San Cristóbal, Los Desamparados, 24.VII.1981, H. Domínguez [MNHNSD]; 1 ♂: Rep. Dom., Prov Pedernales, Cabo Rojo, 10.I.1998, M. Hernández [MNHNSD]; 1 ♂: Rep. Dom., Prov. Santo Domingo Norte, Mirador Norte, 7.X.2001, H. Takizawa [MNHNSD]; 1 ♂: Rep. Dom., Prov. Santo Domingo Norte, Mirador Norte, 12.VIII.2001, H. Takizawa [MNHNSD]; 1 ♂ (penultimate instar): Rep. Dom., Prov. San Cristobal, Haina, 23.VI.1981, H. Domínguez [MNHNSD]; 1 ♂: Rep. Dom., Prov. Azua, El Número, 4.VII.2002, H. Takizawa [MNHNSD]; 1 ♂ (penultimate instar): Rep. Dom., Isla Saona, de Catuano a Damaney, 6.VII.2002, H. Takizawa [MNHNSD]; 1 ♂: G.V, P53, 23.VII.1992 [MNHNSD]; 1 ♀: República Dominicana Prov. San Cristóbal, 11.II.1977 E. Marcano; #21060 [IIBZ]; 1 ♀ (apex of abdomen missing): República Dominicana Prov. San Cristóbal, Cambita, 8.VI.1976, E. Marcano; #20069 [IIBZ]; 1 ♀ (apex of abdomen missing): República Dominicana Prov. San Cristóbal, Campus Loyola, 30.X.1975, E. Marcano; #19189 [IIBZ]; 1 ♀: República Dominicana Santo Domingo, Capital, 11.VI.1973 E. Marcano; #13677[IIBZ]; 1 ♀: República Dominicana Prov. San Cristóbal, Campus Loyola, 30.X.1975, E. Marcano; #19190 [IIBZ]; 1 ♀: República Dominicana Prov. San Cristóbal, Politécnico Loyola, VII.1989, E. Marcano [IIBZ]; 1 ♀: República Dominicana Prov. San Cristóbal, Cambita, 8.VI.1976, E. Marcano; #20070 [IIBZ]; 1 ♀: República Dominicana Prov. Santo Domingo, 8.XII.1964, E. Marcano; #909 [IIBZ]; 1 ♀: República Dominicana Prov. San Cristóbal, cuidad, 19.V.1976, E. Marcano; #20065 [IIBZ]; 1 ♀: República Dominicana Prov. Barahona, 1.VI.1967, E. Marcano & I. Fernández; #4212 [IIBZ]; 1 ♀: República Dominicana Prov. San Cristóbal, Campus Loyola, 30.X.1975, E. Marcano; #19187 [IIBZ]; 1 ♀: República Dominicana Santo Domingo, Distrito Nacional, Engombe, 19.VI.1979 [IIBZ]; 1 ♀: República Dominicana Prov. San Cristóbal, ciudad, 10.IV.1975, E. Marcano; #17985 [IIBZ]; 1 ♀: República Dominicana Prov. San Cristóbal, ciudad, 19.V.1976, E. Marcano; #20064 [IIBZ]; 1 ♂: República Dominicana Prov. San Cristóbal, Campus Loyola, 23.X.1975, E. Marcano,; #19038 [IIBZ]; 1 ♀: República Dominicana Prov. San Cristóbal, Campus Loyola, 30.X.1975, E. Marcano; #19190 [IIBZ]; 1 ♀. República Dominicana Prov. San Cristóbal, Campus Loyola, 5.III.1976, E. Marcano; #19687 [IIBZ]; 1 ♀: República Dominicana Prov. San Cristóbal, 21.I.1975 E. Marcano ;#17497 [IIBZ]; 1 ♀: República Dominicana Prov. Barahona, 10.VI.1967, E. Marcano; #4196 [IIBZ]; 1 ♀: República Dominicana Prov. San Cristóbal, cuidad, 19.V.1976, E. Marcano; #20063 [IIBZ]; 1 ♀:


HENNEMANN ET AL.

República Dominicana Prov. Santo Domingo, 9.II.1969, E. Marcano,;#4854 [IIBZ]; 1 ♀: República Dominicana Prov. San Cristóbal, Campus Loyola, 30.X.1975, E. Marcano; #19188 [IIBZ]; 1 ♀: República Dominicana Prov. San Cristóbal, cuidad, 19.V.1976, E. Marcano; #20062 [IIBZ]; 1 ♀ (apex of abdomen missing): República Dominicana Prov. La Vega, Constanza, La Palma, 1.VIII.1968, E.Marcano & S. Raud; #4589 [IIBZ]; 1 ♂: República Dominicana, Santo Domingo, Distrito Nacional, octubre del 1970 [IIBZ]; 1 ♀: República Dominicana Prov. San Cristóbal, Campus Loyola, 6.I.1976, E. Marcano; #19549 [IIBZ]; 1 ♀ (apex of abdomen missing): República Dominicana Prov. San Cristóbal, Cambita, 8.VI.1976, E. Marcano; #20070 [IIBZ]; 1 ♀: República Dominicana Prov. San Cristóbal, Campus Loyola, 30.X.1975, E. Marcano; #19190 [IIBZ]; 1 ♀: Dominican Republic, Pedernales 30 km N Cabo Rojo 1070 m, 18-07N W71–39W 27 September 1991 R. Davidson, C. Young, S. Thompson, J. Rawlins Reservoir, pine woods [CMNH]; 1 ♂: Republique Dominicaire, route Oviedo à Pedernales km 17, Jaragua National Park, 11-V-2004, J. Barlout réc. [MNHN]; 1 ♂: République Dominicaine, route Oviedo à Pedernales km17, Jasagua National Park, 11.V.2004, J. Barbut réc [MNHN]; 8 ♂♂, 4 ♀♀, eggs: ex Zucht: F. Hennemann, VI.1995, Herkunft: Dominikanische Rep., PSG No. 61 [coll. FH, No’s 0243-1 to 12 & E1]; 1 ♂, 6 ♀♀: ex Zucht: F. Hennemann, 2001–2002, Herkunft: Dominikanische Rep., PSG No. 61 [coll. FH, No’s 0243-13 to 19 & E2]. HISPANIOLA (HAITI): 1 ♂: Furcy, Haiti, I.21.1945, Anthony Curtis!, wO134; Diapherodes spinipes (Gray) det. C. F. Moxey [ANSP]; 1 ♂: Furcy, Haiti, I.21.1945, Anthony Curtis!, wO136; Diapherodes spinipes (Gray) det. C. F. Moxey [ANSP]; 1 ♀: Furcy, Haiti, Feb. 5.1943, A. Curtis; wO138; Diapherodes spinipes (Gray) det. C. F. Moxey [ANSP]; 1 ♀: Furcy, Haiti, Feb. 5.1943, A. Curtiss; wO139; Diapherodes spinipes (Gray) det. C. F. Moxey [ANSP]; 1 ♂: Haiti, Dept. De l’Ouest, Port au Prince, 5.IX.1973, JC Schuster [FSCA]; 1 ♀: Coll. Br. v. W., Port au Prince, Stevens; det. Br. v. W. Haplopus ligia Westw.; 72., 6479, 6479 [NHMW]; 1 ♀: Port au Prince, Haiti, Dr. Fritz Rauch leg., ded. 31.XII.1900; PHA 24, Zoologisches Museum Hamburg [ZMUH]; 1 ♀: Port au Prince, Haiti, Dr. Rauch leg., W. Pohlmann durch Gust. Keitel jr., ded. 29.V.1901; PHA 23, Zoologisches Museum Hamburg [ZMUH]. NO DATA: 1 ♀: Aplopus ligius Westw. A.n.c. [USNM]; 1 ♂: W. Ind.; Aplopus cythereus Westw. ♂ det. Rehn [USNM].

Diagnosis: This widely distributed and very variable species is readily distinguished from all other species in the genus by the large roundly triangular, shield-shaped or almost semi-circular epiproct of ♀♀ (Figs. 241–242), as well as the distinctly white cheeks (Figs. 231–233), orange to red antennae, short terminal hook of the vomer (Fig. 352) and long alae of ♂♂, which clearly project over posterior margin of abdominal segment V (Figs. 224–227). Description: The description of the colouration is mostly based on colour photographs of various live wild and captive reared specimens. ♀ (Figs. 220–223, 386–387). Very variable in size (body length including the subgenital plate 115.5–172.0 mm) and several morphological features like the armature of the head and thorax, abdominal tergites, shape of the epiproct, and length of the subgenital plate and alae (→ comments on variability below). Colour ranging from dull green, over straw, grey and pale to dark brown, sometimes with a yellowish or reddish wash and with a variable degree of white portions in some localities. Abdomen in particular to a variable degree furnished with paler and darker mottling and speckles. Head with the cheeks ± distinctly white and one or two small black spots over the eyes; occasionally a further pair of black spots on the frons between the bases of the antennae. Antennae pale to mid brown dorsally and dull red ventrally. Spines of the thorax either brown, dull red or green. Tegmina and costal region of alae mid to dark brown, rarely with a greenish wash; either plain or to a variable degree furnished with faint pale brown or grey spots. Anal region of alae transparent with all major longitudinal and transverse veins broadly marked by dark brown (Fig. 248). Head: About 1.3x longer than wide and roughly oval in dorsal aspect. Vertex rounded and with a pair of very prominent but variably sized occipital spines; points either blunt or with a pointed black tip (Figs. 228–230). Occasionally one or two further pairs of small spiniform tubercles close to posterior margin of head capsule (Fig. 230). Eyes circular and contained about 2.5x in length of cheeks. Antennae ± reaching to posterior margin of median segment and consisting of 60–66 segments. Scapus almost 2x longer than wide and slightly narrowed towards the base. Pedicellus about half the length of scapus and roughly equal in length to III. Thorax: Pronotum a little longer but slightly narrower than head, about 1.6x longer than wide, roughly rectangular but the lateral margins with a deep roundly triangular emargination medially. Transverse median sulcus moderately distinct and almost reaching lateral margins of segment. Dorsal surface at least with a distinct pair of spines in anterior portion, but usually with a very variable number of additional spines and tubercles (Figs. 228–230). Probasisternum and profurcasternum with single granules or tubercles. Mesothorax 2.0–2.3x longer than head and pronotum combined. Mesonotum rather narrow anteriorly and very slightly gradually widened towards the posterior; surface armed with a very variable number of differently sized tubercles or blunt spines, a rather



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defined marginal row of evenly sized tubercles or small spines is present laterally. Meso- and metapleurae with an irregular longitudinal marginal row of spiniform tubercles or spines. Meso- and metasternum with a variable number of granules or low spiniform tubercles; may be no more than four on the metasternum. Metanotum less than 1/3 the length of mesonotum, unarmed. Tegmina oval, coriaceous, with the venation very distinct, dense and irregularly disposed, and slightly projecting over posterior margin of metanotum; median protuberance rather faint and roundly conical. Alae variable in length but always longer than tegmina; at least reaching ¾ along median segment but often projecting to anterior portion of abdominal tergum II (Fig. 223). Abdomen: Median segment 2x longer than wide and slightly narrowed medially. Segments II–V of roughly equal length, VI a little shorter than V, VII just a little more than ¾ the length of V; on average 1.6–1.8x longer than wide. Tergites II–IV often with a median pair of tubercles or spiniform processes close to posterior margin; most prominent on II (Figs. 234–235). VII with four very fine and low, longitudinal, sub-parallel carinae and lateral margins ± deflexed posteriorly or with a ± acutely triangular posterior lobe (Figs. 237–239). Praeopercular organ formed by a slightly raised dark oval marking in posterior margin of sternum Vii, which bears a rough, rounded tubercle posteriorly (Fig. 340). Tergum VIII slightly shorter than VII, gently narrowed medially and almost 2.5x longer than wide; sometimes with a pair of posteromedian tubercles. IX rectangular and roughly half the length of VIII. Anal segment with a distinct longitudinal carina, narrowed in posterior half and with a widely triangular posteromedian emargination. Epiproct prominent and at least 1/3 the length of anal segment; size and shape variable but mostly ranging from roundly triangular to almost semicircular and shield-like (Figs. 241–242). Cerci conical with a rather acute tip and slightly laterally compressed, hardly projecting over posterior margin of anal segment. Subgenital plate very long, lanceolate or cymbiform, longitudinally carinate and with a ± acute apex; extending greatly over apex of abdomen (Figs. 236–240). Legs: Profemora shorter than mesothorax, mesofemora about as long as metathorax and metafemora reaching ± half way along abdominal segment IV. Profemora occasionally with one or two small sub-apical spines on the medioventral carina. Protibiae often gently undulate dorsally. Anteroventral carina of meso- and metafemora with two, posteroventral carina with only sub-apical spine. Medioventral carina of mesofemora with 3–5, of metafemora with 5–7 ± prominent spines. Dorsal carinae each with a ± distinct roundly triangular tooth or expansion; much more prominent on the posterior carina. Posteroventral carina of meso- and metatibiae very minutely denticulate; anterodorsal carina occasionally with a roundly triangular tooth some ¼ off the base. Basitarsi about as long as following three tarsomeres combined. ♂ (Figs. 224–227, 388). Variable in size (body length 82.6–103.0 mm) and colouration but well recognized by the ± white cheeks and long alae. Although some morphological features vary to a certain degree, the variation is much less than in ♀♀. Alae well developed (length 40.5–51.0 mm) and at least reaching to abdominal tergum VI. Great parts of meso- and metapleurae, ventral body surface and legs green. Head either green or pale brown with the cheeks ± distinctly white (Figs. 231–233). Dorsal surface of thorax and abdomen straw to pale creamish brown. Spines of thorax either dull green or black, posterior portions of meso- and metapleurae yellow and the metapleurae with a pale brown longitudinal band along lower margin. Lateral margins of abdominal tergum VIII and IX broadly white. Tegmina and costal region of alae pale to creamish mid brown and to a variable degree furnished with irregularly disposed pale cream to white markings; anterior margin of tegmina broadly white and basal portion of alae black. Anal region of alae pink with all major veins brown (Fig. 247). Antennae, except scapus and pedicellus, ± distinctly red. Tarsi greenish brown. In addition to this more common green form described above brown forms occur in certain localities (Fig. 227). These generally agree in colouration to the green forms but lack all the green body parts described above. Head: Generally as in ♀♀ but with the cephalad horns often more prominent (Figs. 231–233). Eyes more prominent, projecting hemispherically and contained no more than 1.6x in length of cheeks. Antennae very robust and ± reaching posterior margin of abdominal segment III; with 60–66 segments. Thorax: Pronotum longer but narrower than head, general shape as in ♀♀. Armature usually limited to a prominent pair of spines in the anterior portion but sometimes a much smaller pair of spines may be present in the posterior portion (Figs. 231–233). Mesothorax about 2.0–2.2x longer than head and pronotum combined. Mesonotum with 4–10 distinct and ± pointed, roughly paired spines in the anterior 2/3 of the surface. Mesosternum with a few irregularly disposed granules or low spiniform tubercles; metasternum either smooth or with 2–4 paired tubercles in anterior portion. Tegmina oval and slightly projecting over posterior margin of metanotum, central protuberance roundly conical. Alae at least reaching 1/3 the way along abdominal segment VI.


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FIGURES 220–223. Haplopus micropterus (St. Fargeau & Audinet-Serville, 1828). 220. HT ♀: St. Thomas [RMNH]; 221. HT ♀ of synonym Haplopus obtusus Redtenbacher, 1908: St. Croix [NHMW, No. 835]; 222. ♀: Dominican Republic, Pedernales Province, Boca de la Cañada [USNM]; 223. ♀: Dominican Republic, Azua Province, El Curro (specimen with very long alae) [USNM].



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FIGURES 224–227. Haplopus micropterus (St. Fargeau & Audinet-Serville, 1828). 224. ♂: captive reared from Dominican Republic [coll. FH, No. 0243-19]; 225. ♂: Dominican Republic, Baní Peravia Province, Pueblo Nuevo [USNM]; 226. ♂: Dominican Republic, Monte Rio [USNM]; 227. ♂: Dominican Republic, Pedernales Province, Boca de la Cañada [USNM].

Abdomen: Segments II–IV of equal length and about 3.2x longer than wide. V–VII slightly decreasing in length with VII no more than 2.3x longer than wide. All tergites and sternites smooth. VII slightly expanded posteriorly or with a ± distinct triangular lobe (Figs. 244–245). VIII ¾ the length of VII and gently widening towards the posterior; IX shorter than VIII. Anal segment with a faint longitudinal median carina which gradually becomes more decided towards the apex. Posterior portion rounded and with a wide triangular median emargination (Figs. 244–246); posterior margin slightly swollen and on ventral surface armed with several small, black in-curving denticles (Fig. 360). Epiproct very small and roundly triangular (Fig. 246). Vomer broad with a large, rounded base and a fairly short terminal hook (Fig. 360). Cerci large, obtuse and almost as long as anal segment, slightly laterally compressed basally. Poculum moderately convex, cup-like and with a blunt basal hump (Fig. 243); posterior portion carinate longitudinally.


HENNEMANN ET AL.

FIGURES 228–235. Haplopus micropterus (St. Fargeau & Audinet-Serville, 1828). 228. Head and pronotum of ♀ from Dominican Republic, Independencia Province, Sierra de Neiba [USNM]; 229. Head and pronotum of ♀ from Dominican Republic, Pedernales Province, Boca de la Cañada [USNM]; 230. Head and pronotum of ♀, captive reared from Dominican Republic [coll. FH, No. 0243-15]; 231. Head and pronotum of ♂ from Dominican Republic, Baní Peravia Province, Pueblo Nuevo [USNM]; 232. Head and pronotum of ♂ from Dominican Republic, Baní Peravia Province, Pueblo Nuevo [USNM]; 233. Head and pronotum of ♂ from Dominican Republic, Santo Domingo Province, Santo Domingo [USNM]; 234. Abdominal tergum II of ♀ from Dominican Republic, Pedernales Province, Boca de la Cañada [USNM]; 235. Abdominal tergum II of ♀, captive reared from Dominican Republic [coll. FH, No. 0243-15].



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FIGURES 236–248. Haplopus micropterus (St. Fargeau & Audinet-Serville, 1828). 236. Apex of abdomen of ♀ HT (lateral view) [RMNH]; 237. Apex of abdomen of ♀ HT (dorsal view) [RMNH]; 238. Apex of abdomen of ♀ from Dominican Republic, Pedernales Province, Boca de la Cañada (dorsal view) [USNM]; 239. Apex of abdomen of ♀ PLT of synonym Haplopus ligius Westwood, 1859 (dorsal view) [NHMUK]; 240. Apex of abdomen of ♀ PLT of synonym Haplopus ligius Westwood, 1859 (lateral view) [NHMUK]; 241. Anal segment and epiproct of ♀ from Dominican Republic, Pedernales Province, Boca de la Cañada (dorsal view) [USNM]; 242. Anal segment and epiproct of ♀, captive reared from Dominican Republic) (dorsal view) [USNM]; 243. Apex of ♂ abdomen, captive reared from Dominican Republic (lateral view) [coll. FH, No. 0243-19]; 244. Apex of ♂ abdomen captive reared from Dominican Republic (dorsal view) [coll. FH, No. 0243-19]; 245. Apex of abdomen of ♂ LT of synonym Haplopus ligius Westwood, 1859 (dorsal view) [NHMUK]; 246. Anal segment of ♂, captive reared from Dominican Republic (dorsal view) [coll. FH, No. 0243-19]; 247. Left ala of ♂, captive reared from Dominican Republic [coll. FH, No. 0243-19]; 248. Left ala of ♀, captive reared from Dominican Republic [coll. FH, No. 024313].


HENNEMANN ET AL.

FIGURES 249–256. Haplopus micropterus (St. Fargeau & Audinet-Serville, 1828), Eggs. 249. Dominican Republic, Azua Province, El Curro (dorsal view) [USNM]; 250. Dominican Republic, Azua Province, El Curro (lateral view) [USNM]; 251. Captive reared from Dominican Republic (dorsal view) [coll. FH, No. 0243-E2]; 252. Captive reared from Dominican Republic (lateral view) [coll. FH, No. 0243-E2]; 253. Dominican Republic, Baní Peravia Province, Pueblo Nuevo (dorsal view) [USNM]; 254. Dominican Republic, Baní Peravia Province, Pueblo Nuevo (lateral view) [USNM]; 255. Captive reared from Puerto Rico (dorsal view) [coll. FH, 0243-E3]; 256. Captive reared from Puerto Rico (lateral view) [coll. FH, 0243-E3].

Legs: Profemora about as long as mesothorax, mesofemora slightly shorter than mesothorax and metafemora ± reaching to posterior margin of abdominal segment IV. Armature generally as in ♀♀ but less distinct and the mesoand metafemora lacking the sub-apical dorsal tooth or expansion seen in ♀♀. Tarsi relatively more elongate and basitarsi a little longer than following three tarsomeres combined. Nymphs: Very variable in colour, size and shape of the armature of the head, thorax, abdomen and legs, which generally is much stronger developed than in adult insects. Colour various shades of brown, often with a greenish wash and/or all over furnished with whitish speckles and markings. Egg (Figs. 249–256): A large number of eggs were examined from various localities throughout the Dominican Republic, Puerto Rico and the British Virgin Islands. Just like the insects, these show considerable variability which however is mainly limited to the size and colouration. Rather large for the genus, capsule ovoid and 1.5–1.7x longer than wide; dorsal surface more convex than ventral surface. Capsule surface ± coriaceous and granulose. Micropylar plate roughly 2/3 the length of capsule, narrowed in anterior portion, then gradually widened with the posterior portion broadly rounded. Posteromedially



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with a deep triangular to parallel-sided emargination. Micropylar cup small and cup-shaped. Operculum ± round in cross-section, convex and in centre with a large, squamiform, mushroom-shaped capitulum (height > 0.3 mm). Colouration of capsule highly variable and ranging from pale drab over ochre and grey to dull orange brown; usually to a variable degree furnished with dark brown to black markings, spots and speckles. Capitulum dull yellow, straw or pale ochre. Measurements [mm]: Length 3.1–3.7, length (including operculum) 4.3–4.8, width 2.1–2.6, height 2.4–2.7, length of micropylar plate 2.3–2.5. Variability: This is a very variable species which not only varies from locality to locality but also shows considerable variation within single colonies or culture-stocks reared in Europe. The most important variable characters are summarized separately for each sex below. Numerous features underlie considerable variability in ♀♀, some of which have already been mentioned in the description presented above. The body length ranges from 114.0–172.0 mm including the subgenital plate (103.0–152.5 mm excluding it), with the largest specimens occuring on Hispaniola. The cephalad horns are variable in size and may either be blunt (most specimens from Puerto Rico & Virgin Islands) or have a conspicuous, acute black point at the apex (most Hispaniolan specimens). The pronotum may be covered with numerous paired tubercles but in some cases merely bears a single pair of spines in the anterior portion. The armature is particularly distinct and well developed in specimens from certain localities in the Dominican Republic (e.g. Boca de la Cañada, Pedernales Province or Sierra Martín García, Azua Province). The mesothoracic armature is extremely variable with structures ranging from small spiniform tubercles to prominent spines. Some captive reared specimens from Hispaniola in the first author's collection (coll FH, No's 0243-2 & 17) merely have the mesonotum with a few small irregularly dispersed tubercles, while there are numerous spines of variable sizes in some wild ♀♀ at hand from the Dominican Republic. The ♀ from near Cabo Rojo (Pedernales Province, Dominican Republic) in FSCA is very small (body length 103.0 mm and 114.0 mm including the subgenital plate) but has the mesothoracic armature extremely prominent and the mesonotum and metapleurae all over armed with medium to large conical spines. Abdominal tergites II–IV usually bear a median pair of tubercles or spiniform processes close to posterior margin, which vary strongly in size and are most prominent on II (Figs. 234–235). Often there are only processes on II and more rarely these are entirely lacking. The alae may just reach ¾ the way along the median segment but in extremes also cover the anterior portion of abdominal tergum II (Fig. 223). Depending on the locality, the antennae are just slightly reddish in most Hispaniolan specimens but distinctly dull red in specimens from Puerto Rico and the Virgin Islands. Males are much less variable than ♀♀ with variability generally restricted to the size, colouration, size of the cephalad horns (Figs. 231–233), number of spines on the mesothorax and granules on the meso- and metasternum, shape of abdominal tergum VII as well as the number of spines on the medioventral carina of the femora. The latter shows the same range as in ♀♀. In addition to the features mentioned the pronotum may occasionally bear a further pair of spiniform tubercles in the posterior portion. They occur in a common green form (Figs. 224–226) and more rarely encountered brown form. The brown specimens found in certain localities in the Dominican Republic (e.g. Boca de la Cañada, Pedernales Province, Fig. 227) fully agree with the green form except for lacking any of the green body parts. The ventral body surface, meso- and metapleurae and legs are all greyish mid brown with irregular whitish mottling. The spines of the thorax however are dull green and the brown ♂♂ from the mentioned locality at hand also have the basal portion of the posterior margin of the tegmina white. Specimens at hand from Parque Nacional Sierra de Bahoruco (RD-191) in USNM deserve special mention. Females are considerably more stocky with the mesothorax relatively shorter and the armature much more prominently developed than in all other examined specimens. But despite the different appearance at first glance, the genital morphology of both sexes does not show any characters that would allow a clear distinction as a distinct subspecies or species. Only the epiproct of ♀♀ is on average larger than in other specimens but seems to be within the range of variability of H. micropterus. As in the insects, considerable variability is also seen in the eggs which however mainly is mainly limited to the colouration as well as the size of the capitulum (Figs. 219–226). Eggs laid by specimens from Puerto Rico (Figs. 225–226) or the British Virgin Islands appear to be generally more reddish and have less distinct mottling, while eggs at hand from localities throughout the Dominican Republic are often prettily mottled with dark brown (Figs. 219–222). However, remarkable variation is already seen within single colonies or culture-stocks and almost plain reddish brown eggs are also laid by some colonies in the Dominican Republic (e.g. from Pueblo Nuevo, Peravia Province).


HENNEMANN ET AL.

Comments: This species was first illustrated and described by Stoll (1788, pl. 21: 77) from a ♀ presumed from “Amboina” and subsequently (Stoll, 1813) named Phasma angulata. The name “Phasma angulata” had however already been used for another species by Fabricius in 1793 (→ Diapherodes angulata angulata (Fabricius, 1793)), and for this reason Saint Fargeau & Audinet-Serville (1828: 445) introduced Cyphocrana microptera as a replacement name for Stoll's species. There has been confusion concerning the published year of Haplopus micropterus (St. Fargeau & Serville) but according to Sherborn & Woodward (1899: 595) it should be 1828, not 1825 as recorded previously. The holotype of Phasma angulata Stoll (= Cyphocrana microptera St. Fargeau & Audinet-Serville, 1828) was not traced in RMNH by Bragg (1996) and since believed lost. Certainly, no author had ever seen Stoll's type specimen but an extensive search in the collection of RMNH in 2006 has revealed a ♀ which perfectly matches with the illustration provided by Stoll (1788, pl. 21: 77) and is here regarded as the previously missing holotype. The particular specimen is pinned with the same typical insect pin used in all of Stoll's types in RMNH and furthermore bears the same typical circular label seen on most of Stoll's specimens. Based on Stoll's figure the main diagnostic character of this species in the past has been the presence of a conspicuous white spot in the basal portion of the alae, which is however not seen in any known representative of Haplopus. Examination of the ♀ holotype shows this to have a large hole in each wing caused by a pin drawn through the wing by the preparator to spread these organs, hence are an artifact. The size and shape of these holes correspond to the “white spots” reproduced in Stoll's illustration and prove this artifact to have been misinterpreted by all subsequent authors dealing with the systematics of H. micropterus. H. micropterus shows a great deal of intraspecific variability and is widely distributed throughout Hispaniola, Mona Island, Puerto Rico and the Virgin Islands. The variability of certain characters is summarized above and has caused much confusion in the identity of this species and also resulted in several synonymies. Examination of the ♀ holotype of H. bituberculatum de Haan, 1842 in RMNH leaves no doubt this species is a synonym of H. micropterus, which might even have been collected alongside with the HT specimen described and illustrated by Stoll (n. syn.). It is pinned on the same kind of insect pin. Another synonym is represented by Haplopus obtusus Redtenbacher, 1908 whose ♀ holotype from “Sta. Kruz” (= Saint Croix) is a typical specimen of H. micropterus (n. syn.). The short and apically rounded subgenital plate of the holotype has proven to be an artifact, the apex being broken off and swollen due to an injury. Redtenbacher (1908: 432) described Haplopus ligiolus based on a ♂ and two ♀♀ from Mona Island in ZMUH and a ♀ from “Westindien” in NHMB. Although all three specimens are traced, H. ligiolus was omitted in the catalogue of type-material deposited in ZMUH by Zompro (2002). The ♂ in ZMUH is here designated as the lectotype of H. ligiolus Redtenbacher, 1908 in order to guarantee stability of the name and the synonymy here established. Careful examination of the four specimens has shown H. ligiolus to be a synonym of H. micropterus (n. syn.). Haplopus cytherea Westwood, 1859 was described from two ♂♂ from Santo Domingo in OXUM of which one (the specimen illustrated by Westwood, 1859, pl. 18: 5) is here selected as the lectotype. Both are typical specimens of H. micropterus, and for this reason this species also becomes a synonym (n. syn.). Examination of the type-specimens of Haplopus ligia Westwood, 1859 in NHMUK has shown these to represent merely a variety of this very variable species, consequently also H. ligia is a junior synonym of H. micropterus (n. syn.). The Santo Domingo specimen illustrated by Palisot de Beauvois (1805: 166, pl. 14: 4) as Phasma angulata is, as already noted by Westwood (1859: 87), a ♀ with the last three abdominal segments broken off and is clearly distinct from the species that was described as Mantis angulata by Fabricius (1793: 13, = Diapherodes angulata angulata (Fabricius) from Guadeloupe). Since Palisot de Beauvois's angulata is a junior homonym of Mantis angulata Fabricius, 1793, Gray (1835: 34) introduced Diapherodes spinipes as a replacement name for Palisot de Beauvois's species. The holotype of Phasma angulata Palisot de Beauvois, 1805 (= spinipes Gray, 1835) is lost but the figure of Palisot de Beauvois leaves no doubt it is conspecific with H. micropterus, hence D. spinipes as well is a junior synonym (n. syn.). Since the 1980's live eggs of H. micropterus have been imported to Europe on at least three occasions, from the Dominican Republic, Haiti and Tortola Island (British Virgin Islands). It was included on the Phasmid Study Group culture-list as culture No. 61 “Aplopus sp.” (later corrected to Haplopus micropterus) and is fairly easy to rear in moderately humid but well ventilated conditions and high temperatures (> 25°C). In captivity it readily accepts rose (Rosa spp., Rosaceae), bramble (Rubus fruticosus, Rosaceae), raspberry (Rubus idaeus, Rosaceae), oak (Quercus robur & Q. petraea, Fagaceae), guava (Psidium guajava, Myrtaceae), eucalyptus (Eucalyptus gunnii, Myrtaceae) and hawthorn (Pyracantha spp., Rosaceae) as alternative food plants. At least Malpighia gossipifolia



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(Malpighiaceae), Pictetia spinifolia (Fabaceae) and Bidens cynapiifolia (Asteraceae) are known to be part of the natural diet in Hispaniola. A REM-study of the egg was presented by Lipinski et al. (1999, fig. 25). Lu et al. (in press) present comprehensive information on the distribution, food plants, habitats, biology and nymphal states of H. micropterus in the Virgin Islands and report Piscidia carthagensis (Fabaceae), Pictetia aculeata (Fabaceae) and Pithecellobium unguis-cati (Fabaceae) to serve as host plants. Distribution (Fig. 380): Virgin Islands (St. Thomas [RMNH, MNHU, ZMUH]; St. Thomas (Loango [Werner, 1929: 7], Frenchman's Bay Esatate [Lu et al. in press], Estate Nazareth [Lu et al. in press]); St. Croix: Annaly [USNM]; St. Croix [NHMW]; St. John [MNHU]; St. John (Lameshur Ranger Station [Wenhua et al. In prep.]); Tortola, Long Bay, Belmont Estate [Lu et al. in press]; Anegada [USNM, Lu et al. in press]; Guana [NHMUK, USNM, Lu et al. in press]; Little Thatch [Lu et al. in press]; Moskito [Lu et al. in press]); Puerto Rico (Rincon [USNM]; Guánica [USNM]); Mona Island [USNM, ZMUH]; Hispaniola: Dominican Republic (Distrito Nacional [USNM]; Peravia Prov. [USNM]; Santo Domingo Prov. [IIBZ, MNHNSD, NHMUK, OXUM, USNM]; Pedernales Prov. [CMNH, FSCA, MNHN, MNHNSD, USNM]; Montecristi Prov. [USNM]; Barahona Prov. [FSCA, IIBZ, USNM]; Azua Prov. [MNHNSD, USNM]; Independencia Prov. [USNM]; Ocoa Prov. [USNM]; San Cristóbal Prov. [IIBZ, MNHNSD, USNM]; Macorís Prov. [MNHNSD]; La Vega Prov. [IIBZ]; Isla Beata [USNM]; Isla Saona [MNHNSD]); Haiti (Gros Morne [Moxey, 1972: 116, in litt.]; Furcy [ANSP]; Port-au-Prince [ANSP, NHMW, ZMUH]; Gonaive Island, Vicinity of Pointe-a-Raquettes [MCZC, Moxey, 1972: 116, in litt.]). Number of specimens examined: 261 TABLE 23. Measurements of Haplopus micropterus (St. Fargeau & Audinet-Serville, 1825) [mm] ♀, HT [RMNH]

♀ HT of obtusus [NHMW]

♀, HT of bituberculatum [RMNH]

♂, LT of ligia [NHMUK]

♂, PLT of ligia [NHMUK]

♀♀, PLT of ligia [NHMUK]


133.5

149.5

125.5

-

-

148.5–151.0

Body

119.5

143.0

112.5

87.0

-

126.5–129.0

Pronotum

5.9

5.8

4.9

3.9

3.7

5.2–5.4

Mesonotum

26.0

29.8

21.0

15.5

15.1

25.5–25.7

Metanotum

6.9

9.1

5.4

5.7

-

7.1–7.3

Median segment

8.9

10.2

9.5

7.4

-

9.5–9.8

Tegmina

9.2

-

7.6

9.8

9.8

8.8–9.0

Alae

10.1

10.8

8.5

41.3

41.0

7.8–8.7

Profemora

19.0

21.2

15.2

15.0

15.7

20.0–20.8

Mesofemora

16.8

19.1

13.7

13.8

13.3

16.1–16.8

Metafemora

21.2

17.2

18.1

17.7

18.4

21.0–23.4

Protibiae

18.0

22.8

14.0

14.1

16.3

19.9–20.1

Mesotibiae

16.2

18.3

12.5

12.3

12.3

16.8–17.0

Metatibiae

20.9

16.3

17.2

16.2

16.9

23.0–23.2

Antennae

ca. 40.0

> 20.4

> 16.1

> 43.0

-

> 43.0

TABLE 24. Measurements of Haplopus micropterus (St. Fargeau & Audinet-Serville, 1825) [mm] ♀♀ Hispaniola

♂♂ Hispaniola

♀♀ Puerto Rico, captive reared [coll. FH]

♂♂ Puerto Rico, captive reared [coll. FH]

♀ [NHMW]


121.5–172.0

-

115.5–122.0

-

149.0

Body

103.5–152.5

84.5–103.0

104.5–110.5

82.6–89.0

128.0



HENNEMANN ET AL.

TABLE 24. (Continued) ♀♀ Hispaniola

♂♂ Hispaniola

♀♀ Puerto Rico, captive reared [coll. FH]

♂♂ Puerto Rico, captive reared [coll. FH]

♀ [NHMW]

5.0–6.2

3.7–3.9

4.9–5.0

3.4–3.8

6.1

Mesonotum

22.8–31.0

13.9–20.0

20.4–21.8

14.5–16.0

28.5

Metanotum

6.8–9.2

7.1

6.9–7.7

6.1

7.0

Median segment

7.8–10.5

7.9

7.1–7.8

7.3

9.1

Tegmina

9.9–12.4

11.0–12.4

8.0

9.8–10.6

9.2

Alae

10.3–15.0

45.5–51.0

10.4–11.8

40.5–45.0

9.0

Profemora

19.8–25.6

17.8–18.4

19.2–19.7

18.8–19.7

-

Mesofemora

16.0–21.8

14.9–19.8

16.6–17.3

15.7–18.3

18.2

Metafemora

21.6–29.7

19.2–26.4

22.1–23.0

21.2–23.5

24.8

Protibiae

17.8–24.8

17.7–23.9

18.7–19.0

19.4–20.9

-

Mesotibiae

14.9–22.2

15.0–18.9

15.6–17.2

14.2–15.3

18.0

Metatibiae

21.8–31.2

19.0–25.3

22.0–22.8

19.6–22.0

23.8

Antennae

50.0–65.0

53.0–62.0

52.0–56.0

63.0–67.0

-

Pronotum

Haplopus scabricollis (Gray, 1835) (Figs. 257–275, 341, 361–362) Diapherodes scabricollis Gray, 1835: 34. LT (by present designation), ♂: no data [possibly from the Bahamas → see comments below] [LSUK]; PLT, ♀: no data [LSUK]. Moxey, 1972: 110 (in litt.). Haplopus scabricollis Westwood, 1859: 88. Kirby, 1904a: 364. Redtenbacher, 1908: 432. Otte & Brock, 2005: 152. Aplopus mayeri Caudell, 1905: 83. HT, ♂: Loggerhead Key, Dry Tortugas, Fla; Catal. No. 54; Aplopus mayeri Caud. ♂ TYPE; Type No. 42774 U.S.N.M. [USNM]; AT, ♀: Loggerhead Key, Dry Tortugas, Fla; Catal No. 54; Aplopus mayeri Caud. ♀ Type; Brooklyn Museum Coll. 1929; Allotype No. 42774 U.S.N.M. [USNM]; PT, ♂: Loggerhead Key, Dry Tortugas, Fla; Catal. No. 54; Brooklyn Museum Coll. 1929; Paratype No. 42774 U.S.N.M. [USNM]; PT, ♀: Loggerhead Key, Dry Tortugas, Fla; Catal. No. 54; Brooklyn Museum Coll. 1929; Paratype No. 42774 U.S.N.M. [USNM]; PT, ♀: Loggerhead Key, Florida, Dry Tortugas; Paratype No. 42774 U.S.N.M.; Aplopus mayeri Caudell ♀ Paratype [USNM]; PT, ♀ (penultimate instar): Loggerhead Key, Dry Tortugas, Fla; Catal. No.54; Brooklyn Museum Coll. 1929; Paratype No. 42774 U.S.N.M. [USNM]; PT, ♀ (nymph n4): Florida. Loggerhead Key; Catal. No. 54; Brooklyn Museum Coll. 1929; Paratype No. 42774 U.S.N.M. [USNM]; PT, ♂ (nymph n4): Loggerhead Key, Dry Tortugas, Fla; Catal. No. 54; Paratype No. 42774 U.S.N.M. [USNM]; PT, ♀ (nymph n5): Loggerhead Key, Dry Tortugas, Fla; Catal. No. 54; Brooklyn Museum Coll. 1929; Paratype No. 42774 U.S.N.M. [USNM]; PT, ♂ (in Ryker mount): Loggerhead Key Florida, Dry Tortugas, Dr. Mayer; Aplopus mayeri Caudell ♂ Paratype; Paratype No. 42774 U.S.N.M. [USNM]; PT, ♀: Loggerhead Key, Dry Tortugas, Fla. [AMNH]; PT, ♀ (nymph): Loggerhead Key, Dry Tortugas, Fla. [AMNH]. n. syn. Werner, 1929: 9, figs. B (♂) & C (♀). Stockard, 1908a: 239ff, figs. (♀, 1st instar nymph & egg). Stockard, 1908b: 43, figs. (♂, ♀ & egg). Caudell & Hebard, 1912: 159. [Designation of lectotype] Rehn & Hebard, 1914: 101. Arment, 2006: 18. Haplopus mayeri, Redtenbacher, 1908: 433. Werner, 1929: 9, figs. Otte & Brock, 2005: 152. Haplopus evadne, Caudell, 1904: 949, figs. 1 & 2 (♀). [Misidentification] Aplopus similis Rehn, 1904: 65. HT, ♂ (penultimate instar nymph): Swan Isl. Caribbean Sea; U.S.N.M. Acc. 19099, Cat. no. Aplopus similis Rehn TYPE; Type No.7343 U.S.N.M., wO192 [USNM], PT, ♀: acc 19099, Swan Isld., Caribbean Sea;



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Cat no. Aplopus similis Rehn TYPE; Type No. 7343 U.S.N.M. [USNM]. n. syn. Diapherodes similis, Moxey, 1972: 112 (in litt.). Haplopus similis, Kirby, 1904a: 364. Redtenbacher, 1908: 432. Otte & Brock, 2005: 152. Further material [53 ♂♂, 44 ♀♀, 64 nymphs]: DRY TORTUGAS: 16 ♂♂, 6 ♀♀, 10 ♂♂ (nymphs); 9 ♀♀ (nymphs): Loggerhead Key, Dry Tortugas, Fla., July 8, 1912, R&H [ANSP]; 1 ♀: Aplopus mayeri Caudell, Loggerhead Key, Fla. [ANSP]; 10 ♂♂, 7 ♀♀, 7 nymphs: Loggerhead Key, Dry Tortugas, FLA; R. E. Woodruff coll. 2.IX.61; At Suriana maritima L. [FSCA]; 1 ♀: Dry Tortugas, July 28, 1938, Jack Russel, No. 8271; Aplopus mayeri Caudell, det. R. E. Woodruff—70 [FSCA]; 22 ♂♂, 23 ♀♀, 31 nymphs (n4 to penultimate instar): Tortugas, FLA., Loggerhead Key, VII. [various dates] 1938, Jack Russell; Aplopus mayeri Caud., Det. By T.N. Hubbell 1956 [UMMZ]. FLORIDA KEYS: 1 ♂ (nymph n4): Key West, Fla. July 3–7, 1912, leg. R & H [ANSP]; 1 nymph (n2): Long Key, Fla. July 13, 1912, R&H [ANSP]; 1 nymph (n2): Key Largo, Florida, Monroe County, III, 18, 1910; A. mayeri Caud., Hebard Collection [ANSP]; 1 ♀: Bahia, Honda, Fla., VIII-14–38 [ANSP]; 1 ♀ (nymph—in Ryker mount): Key West Fla, 6.IV.03; EA Schwarz Collector; Aplopus mayeri Caudell ? ♀ nymph [USNM]. BAHAMAS: 1 ♀: Bahamas, J. L. Bouchote, 1902.-299, coll. J. L. Bouchote, Feb. 1902, Wood C. Andros, G. Smith [NHMUK]; 1 ♂, 2 ♀♀: Nassau, N. P., Bahamas, Veronica Higgs!; Diapherodes scabricollis Gray, det. C. F. Moxey 1972 [ANSP]; 1 ♀: “Pink House”, on road, vic FFS, WP 132, 13.V.03 [ANSP]; 1 ♂: Bahamas: Eleuthera, Rainbow Bay, I-VII-1988, R.W. & D.B. Wiley [FSCA]; 1 ♂ (penultimate instar), 1 ♀ (nymph n5): Bahamas: Andros Is., S. Fresh Creek nr. U.S. Naval Base; 26VII-2006, Trevor Smith; beating [FSCA]. NAVASSA ISLAND: 1 ♂: Navassa Island; near lighthouse, 80 m, 18°23.82’N, 75°00.74’W, 24 July 1998, Collrs. W. E. Steiner, J. M. Swearinger, et al. [USNM]. CAYMAN ISLANDS: 1 ♂: Roy. Soc.-CIG Expdn. Little Cayman, BWI, North Shore track, N. Blossomvillage, 2.8.1975, R. R. Askew [NHMUK]; 1 ♂: 17.IV.–26.VIII.1938, Oxf. Un. Cayman Is., Biol. Exped. Coll. By C. B. Lewis, G. H. Thompson, 2.VII.1938, Grand Cayman, East end of East end, Pres. Hope Dept. Oxford, B. M. 1967-147, East End Grand Cayman, VII-2-38, 7238 J. [NHMUK]; 1 ♀: 17.IV.–26.VIII.1938, Oxf. Un. Cayman Is., Biol. Exped. Coll. By C. B. Lewis, G. H. Thompson, 9.VII.1938, Grand Cayman, N. coast of North Side, G. C., VII-9-38, 7938, Pres. Hope Dept. Oxford, B. M. 1967-147, Diapherodes pulverulentus Gray det. Moxey 1972 [NHMUK]; 1 ♂ (nymph): 17.IV.–26.VIII.1938, Oxf. Un. Cayman Is., Biol. Exped. Coll. By C. B. Lewis, G. H. Thompson, 29.V.1938, Little Cayman, West end of S. W. Point area, Pres. Hope Dept. Oxford, B. M. 1967-147 [NHMUK].

Diagnosis: This, the northernmost distributed representative of the genus, is easily distinguished from all other species in the genus by the very short alae of both sexes, which are considerably shorter than the tegmina (Fig. 269). The short alae of ♂♂ resemble H. brachypterus n. sp. from Hispaniola and H. woodruffi n. sp. from Cayman Brac (Cayman Islands). From the first species ♂♂ clearly differ by: the more robust body; longitudinal white marking along the lateral margins of the pronotum (Fig. 266), and longitudinal white median markings on the median segment and abdominal tergites VIII–X; broader and less distinctly posteromedially indented anal segment (Fig. 242); differently shaped poculum (Fig. 275) and shorter terminal hook of the vomer (Fig. 353). From H. woodruffi n. sp. they differ by: the more slender body; longitudinal white marking along the lateral margins of the pronotum (Fig. 266), and longitudinal white median markings on the median segment and abdominal tergites VIII–X; presence of a posterolateral tooth or lobe on abdominal tergum VII (Fig. 275); more slender three terminal abdominal segments (Fig. 275); longer and more flattened anal segment (Fig. 275); larger basal projection of the poculum (Fig. 274) and differently shaped vomer, which has the terminal hook considerably longer than in H. woodruffi (Fig. 353). Females differ from those of H. woodruffi n. sp. by: the acutely pointed and black-tipped horns of the head (Fig. 268); differently shaped anal segment and larger, shield-shaped epiproct (Fig. 273); ± decided sub-apical dorsal tooth of the meso- and metafemora and sub-basal dorsal tooth of the corresponding tibiae and gently rounded dorsal carina of the probasitarsus.


HENNEMANN ET AL.

FIGURES 257–262. Haplopus scabricollis Gray, 1835. 257. ♀: Loggerhead Key (plain brown variety—dorsal view) [ANSP]; 258. ♀: Loggerhead Key (mottled variety—dorsal view) [ANSP]; 259. ♀: Loggerhead Key (plain brown variety—lateral view) [ANSP]; 260. ♀: Loggerhead Key (mottled variety—lateral view) [ANSP]; 261. ♂: Loggerhead Key (dorsal view) [ANSP]; 262. ♂: Loggerhead Key (lateral view) [ANSP].



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FIGURES 263–269. Haplopus scabricollis Gray, 1835. 263. ♂ (var.): Vanassa Island [USNM]; 264. Head and pronotum of ♂ (var.) from Vanassa Island [USNM]; 265. Tegmina and alae of ♂ (var.) from Vanassa Island [USNM]; 266. Head and pronotum of ♂ from Eleutheria Island, Bahamas [FSCA]; 267. Head and pronotum of ♂ from Loggerhead Key [ANSP]; 268. Head and pronotum of ♀ from Loggerhead Key [ANSP]; 269. Tegmina and alae of ♀ from Loggerhead Key [ANSP].


HENNEMANN ET AL.

FIGURES 270–275. Haplopus scabricollis Gray, 1835. 270. Apex of abdomen of ♂ (var.) from Vanassa Island (lateral view) [USNM]; 271. Apex of abdomen of ♂ (var.) from Vanassa Island (dorsal view) [USNM]; 272. Apex of abdomen of ♀ from Loggerhead Key (lateral view) [ANSP]; 273. Apex of abdomen of ♀ from Loggerhead Key (dorsal view) [ANSP]; 274. Apex of abdomen of ♂ from Eleutheria Island, Bahamas (lateral view) [FSCA]; 275. Apex of abdomen of ♂ from Loggerhead Key (dorsal view) [ANSP].

Description: ♀ (Figs. 257–260). Moderate to large (body length including the subgenital plate 125.0–164.0 mm) and moderately slender for the genus (maximum width of mesothorax 5.0–6.0 mm) with very short alae (5.3–8.0 mm); body surface very slightly glabrous. Colour variable and ranging from dark over mid and pale brown to pale grey; pale specimens are often all over furnished with mid to dark brown markings and speckles (Figs. 258, 260). Ventral body surface mostly with a yellowish or greenish wash. Head with one or two small elongate black markings above the eyes. Antennae greyish mid brown, sometimes with a slightly reddish hue. Eyes dark reddish brown. Spines of the thorax dark sepia with the points black. Tegmina and costal region of alae dark brown with the venations of a slightly paler colour; the latter black basally. Anal region of alae transparent pink and all major longitudinal and transverse veins marked with dark brown. Head: About 1.4x longer than wide and ovoid with the cheeks slightly convex. Vertex very gently rounded and armed with a pair of moderately sized, pointed cephalad spines (Fig. 268). Behind these often with a further pair of tubercles. Eyes circular and contained about 2.5x in length of cheeks. Antennae reaching about half way along median segment. Scapus about 2x longer than wide with the lateral margins gently rounded. Pedicellus roughly half the length of scapus and a little shorter than III. Thorax: Pronotum longer but narrower than head, about 2x longer than wide, roughly rectangular and with the lateral margins roundly emarginated medially. Transverse median sulcus faint, gently curved and not reaching lateral margins of segment. Dorsal surface with a pair of moderately sized, blunt spines in the anterior portion; otherwise with a variable number of small rather irregularly disposed spines (Fig. 268). Mesothorax about 2.4x longer than head and pronotum combined. Mesonotum very slightly gradually widened towards the posterior, the surface armed with a variable number of irregularly disposed spines or acute tubercles of variable sizes; usually a ± enlarged pair of spines is present in central portion. Lateral margins with a marginal row of about 8–12 small but pointed spines. Meso- and metapleurae armed with an irregular longitudinal marginal row of moderately sized spines. Mesosternum with a variable number of low spines or tubercles. Tegmina oval, coriaceous, with the venation very distinct, dense and irregularly disposed; hardly reaching to posterior margin of metanotum. The median protuberance very shallow and conspicuously displaced towards the apex of tegmen. Alae only 2/3 the length of tegmina and reaching only about 1/3 the way along median segment (Fig. 269). Abdomen: Median segment about 1.8x longer than wide and very gently widened towards the posterior. All segments unarmed, except for a pair of retrorse and compressed posteromedian spines on tergites II–IV; this most prominent on II but sometimes poorly developed on all three segments. Segments II–VI roughly equal in length and width, all rectangular and about 1.8x longer than wide. Tergum VII slightly shorter and narrower than previous,



139

about 2.2x longer than wide and slightly expanded posteriorly (Fig. 273). Praeopercular organ formed by a low, rounded swelling some distance off the posterior margin of sternum VII (Fig. 341). Tergum VIII slightly shorter than VII, narrowed anteriorly and roughly 2.5x longer than wide. IX slightly narrowed towards posterior and somewhat less than half the length of VIII. Anal segment with a very faint longitudinal median carina, narrowed in posterior half and with a very small triangular posteromedian incision. Epiproct of moderate size, roundly triangular and shield-shaped (Fig. 273). Cerci small and conical with a rather acute tip; slightly projecting over posterior margin of anal segment. Subgenital plate very long, lanceolate, longitudinally carinate and with a rather acute apex; extending over apex of abdomen by ± the combined length of tergites VIII–X (Figs. 272–273). Legs: Profemora about 4/5 the length of mesothorax, mesofemora ± reaching posterior margin of median segment and metafemora reaching about half way along abdominal segment IV. Profemora occasionally with two spines in the apical half of the medioventral carina, otherwise unarmed. Anteroventral carina of meso- and metafemora with two, posteroventral carina with one sub-apical spine; medioventral carina armed with 4–6 pointed spines. Both dorsal carinae with a ± distinct triangular tooth sub-apically. Anterodorsal carina of meso- and metatibiae with an angulate expansion sub-basally. Basitarsi roughly equal in length to following three tarsomeres combined; dorsal carina of probasitarsus gently rounded. ♂ (Figs. 261–262). Moderate to large (body length 83.0–121.0 mm) and fairly stocky (maximum width of mesothorax 2.8–3.1 mm) for the genus with conspicuously shortened alae (length 4.2–6.2 mm). Colouration variable and ranging from pale to mid greenish brown, the abdomen ranging from drab to dark brown. Ventral body surface green, the meso- and metapleurae green with a dull orange longitudinal stripe along lower margin. Head plain green but occasionally with a ± defined whitish postocular stripe. Dorsal spines of pro- and mesothorax mid brown with black tips. Lateral margins of pronotum and abdominal tergites II–IX broadly white. Median segment with a ± defined white longitudinal median stripe, often also with a white longitudinal median marking or stripe on abdominal tergites VII–X. Tegmina and costal region of alae dark brown, the latter black basally. Anal region of alae pink with all major veins brown. Antennae drab to pale brown. Tarsi mid brown. Head: Generally as in ♀♀ but eyes more prominent, projecting hemispherically and their length contained only about 2x in that of cheeks (Figs. 266–267). Antennae moderately robust and slightly projecting over abdominal segment II; with about 65 segments. Thorax: Pronotum slightly longer but a little narrower than head, general shape as in ♀♀; surface smooth except for a moderate pair of spines in the anterior portion (Figs. 266–267). Mesothorax only about 2.0–2.1x longer than head and pronotum combined. Mesonotum with 5–16 irregularly disposed and paired spines of moderate size in the anterior 2/3; the pair at anterior margin largest (Fig. 266–267). Mesosternum with ± ten and metasternum with two pairs of spiniform tubercles. Mesopleurae with a marginal row of granules, metapleurae smooth. Tegmina oval and very slightly projecting over posterior margin of metanotum, central protuberance very shallow. Alae small, shorter than alae and hardly reaching 1/3 the way along median segment. Abdomen: Segment II a little shorter than III–IV and equal in length to V, about 2.5x longer than wide. IV longest segment and roughly 2.8x longer than wide. V–VII gradually decreasing in length with VII no more than 1.7x longer than wide. All tergites and sternites smooth. VII with a ± distinct, roundly triangular tooth posterolaterally (Fig. 275). VIII almost equal in length to VII and gently widening towards the posterior; IX slightly shorter than VIII and narrowed in posterior 2/3. Anal segment with a very faint longitudinal median carina in basal portion and gradually widened toward the posterior; the posterior margin broadly rounded with a very shallow median indentation (Fig. 275); slightly swollen and on ventral surface armed with several small, black incurving denticles. Epiproct very small and roughly triangular. Vomer with a fairly small, roughly semi-circular base and a long, papillate, up-curving terminal hook (Figs. 361–362). Cerci large, obtuse and about equal in length to anal segment. Poculum moderately convex, cup-like and with a fornicate basal hump (Fig. 274); posterior portion with a fine longitudinal median keel and somewhat indented medially (Figs. 361–362). Legs: Profemora roughly equal in length to mesothorax, mesofemora about ¾ the length of mesothorax and metafemora just not reaching posterior margin of abdominal segment IV. All less carinate than in ♀♀ but armature generally alike, except for the dorsal sub-apical teeth of the meso- and metafemora very distinct. Tarsi relatively more elongate and basitarsi a little longer than following three tarsomeres combined. Nymphs: Immature ♀♀ usually have the retrorse posteromedian spines on abdominal tergites II–IV more prominent than adult insects and possess a well developed posterolateral tooth on tergum VII. Colour various shades of brown, grey and white.


HENNEMANN ET AL.

Variability: This very widely distributed species shows considerable variability in size, colouration (see description above), number and size of the thoracic spines and leg-armature. Specimens from the Bahamas on average are larger and somewhat more slender than specimens from the Florida Keys or Dry Tortugas, with ♂♂ usually longer than 90.0 mm and ♀♀ exceeding 140.0 mm including the subgenital plate. Furthermore, specimens from the Bahamas have the cephalad horns more acute and usually tipped with black (♀♀ in particular), whereas these are less developed, blunt and not tipped with black in ♀♀ from the Dry Tortugas or Florida Keys. Also the thoracic armature on average is more prominent in the Bahamas specimens with the pronotum usually bearing more than six spiniform tubercles to short spines. Examples from the Cayman Islands and Santanillas rather resemble specimens from the Bahamas in aspect of the cephalic and thoracic armature, having the cephalad horns distinct, acute and tipped with black and the prothoracic armature strongly developed. A fairly small ♂ (body length 84.5 mm) at hand from Navassa (Fig. 263), a small island only some kilometres southwest off Hispaniola, differs from typical specimens of H. scabricollis from the Dry Tortugas, Florida Keys and the Bahamas by lacking distinct tubercles on the meso- and metasternum, entirely smooth meso- and metapleurae, lack of a posterolateral lobe or tooth on abdominal tergum VII (Fig. 271), less prominent basal projection of the poculum (Fig. 270); broader basal portion of the vomer (Fig. 354) and having only 56 antennomeres (roughly 65 in typical scabricollis). Furthermore, the dorsal carinae of meso- and metafemora entirely lack a sub-apical tooth and there is also no sub-basal tooth on the anterodorsal carina of the corresponding tibiae. The lack of a posterolateral tooth on abdominal tergum VII and smooth dorsal carinae of the mid and hind legs resemble H. woodruffi n. sp. from Cayman Brac (Cayman Islands), but the acutely pointed and black tipped horns of the head (Fig. 264), white lateral margins of the pronotum (Fig. 264), white longitudinal median marking of the median segment, very short alae (Fig. 265) and morphology of the genitalia rather attribute it to H. scabricollis. However, more material from Navassa Island, including the unknown ♀ and egg, is required for any broader discussion on the identity of the present ♂. Comments: This species was first described by Gray (1835: 34) based on a ♂ and ♀ without locality now housed in the Linnean collection in London (LSUK). Both specimens were examined from detailed photographs kindly provided by Paul D. Brock (NHMUK) and in aspect of the large size (body lengths: ♂ lectotype 121.0 mm, ♀ paralectotype 164.0 mm including the subgenital plate; according to Redtenbacher, 1908: 432), acute and blacktipped cephalad horns and strong armature of the pronotum, most certainly are from the Bahamas. The ♀ paralectotype matches pretty well with a specimen from the island of Andros in NHMUK and the ♂ with a specimen from the island of Eleuthera in FSCA. Due to the strongly shortened alae, Gray (1835: 34) misinterpreted the ♂ to be a nymph, which is here selected as the lectotype of Diapherodes scabricollis in order to guarantee stability of the name and the new synonymy here established. Examination of the types of Aplopus mayeri Caudell, 1905 from Loggerhead Key in USNM leaves no doubt they are conspecific with Gray's scabricollis, hence Caudell's species is a junior synonym (n. syn.). Two further paralectotypes, an adult ♀ which lacks the terminal three segments of the abdomen and a large ♀ nymph, are housed in AMNH. These are the specimens that had previously been described and figured by Caudell (1904: 949, figs. 1, 2) as H. evadne Westwood, 1859, a distinct species from Hispaniola. Haplopus similis (Rehn, 1904) was described based on a ♀ and an immature ♂ from the Swan Islands (also Islas Santanilla or Islas del Cisne). Careful examination of these specimens show slight differences from typical H. scabricollis from the Dry Tortugas and Florida Keys. The prominent, acutely pointed and black-tipped cephalad horns and the prominently armed pronotum of the ♀ paratype resemble specimens from the Bahamas and Cayman Islands, but the specimen differs from H. scabricollis from all other localities by the more numerous spiniform tubercles of the mesonotum and meso- and metapleurae in particular, somewhat longer alae and having the praeopercular organ on abdominal sternum VII formed by two rough granules instead of one. The cephalic and thoracic armature are most certainly within the range of variability of H. scabricollis and the praeopercular organ may merely be an individual trait of this particular specimen. The immature ♂ holotype is not useful to distinguish it from H. scabricollis. Consequently, H. similis is here synonymised with H. scabricollis (n. syn.) although this synonymy deserves further evaluation by the availability of fresh material from this far off locality (see comments on distribution below). This is the northernmost distributed representative of the entire Haplopodini, and the only species of Haplopus known to occur on the Bahamas, Florida Keys and even some localities near the coast on mainland Florida. This remarkable disjunct distribution appears obscure and warrants special mention and liekely explanation. H. scabricollis is distributed throughout the Dry Tortugas, Florida Keys and Bahamas but there are also apparently REVISION OF HAPLOPODINI


141

disjunct records from Navassa, a small island southeast off Hispaniola, the Cayman Islands and even as far southeast as the Swan Islands, a group of three small islands some 150 km off the coastline of Honduras. Interestingly however, there have so far been no records from the coastlines of Hispaniola or Cuba. Since H. scabricollis is apparently restricted to coastlines and localities near the coast and is almost exceptionally peculiar to its main host-plant Suriana maritima (Simarubaceae), an occurrence in such habitats on Hispaniola and Cuba can not be fully excluded. The phasmatodean fauna of Cuba is still poorly prospected and perhaps this is also true for such habitats in eastern Hispaniola. Suriana maritima is a pantropic plant and found along coastlines throughout the entire Caribbean, including the Cayman Islands and Swan Islands and is also frequently found along the coasts of Central America. Since Suriana maritima is found on all the islands from which H. scabricollis has so far been recorded, a sufficient precondition is present for the occurrence of this species, a fact that is also true for Cuba and eastern Hispaniola. The coastal habitats of H. scabricollis may explain the wide and disjunct distribution of this particular species, since its distribution near the coast makes it prone to over-water dispersal on flotsam caused by frequent strong hurricanes within the Caribbean. As shown by a large series of specimens H. scabricollis is apparently very abundant on Loggerhead Key of the Dry Tortugas (Stockard, 1908a: 239; 1908b: 43), e.g. the collection of UMMZ containing 22 ♂♂, 23 ♀♀ and 31 nymphs of various sizes all collected during a single month on Loggerhead Key. Rehn & Hebard (1914: 387) also reported it from Bird Key and Garden Key, the other islands of the Tortugas group, as well as Key West and Long Key. Rehn & Herbard (1912) furthermore recorded it from Key Largo, the Everglades and Dade County, Florida, which are the only records from the United States mainland. In the wild H. scabricollis is known to feed almost exclusively on bay cedar (Suriana maritima, Simaroubaceae) and to be excellently camouflaged in these shrubs. Only a few specimens have so far been encountered on sea grape (Coccoloba uvifera, Polygonaceae), which however is known to be eaten by these insects. Stockard (1908a, 1908b) provided detailed studies on the habitats, biology and behaviour of H. scabricollis (as Aplopus mayeri). Distribution: Dry Tortugas (Loggerhead Key [USNM]; Bird Key [Rehn & Hebard, 1914: 387]; Garden Key [Rehn & Hebard, 1914: 387]); Florida Keys (Key West [USNM]; Long Key [ANSP]; Key Largo [ANSP]; Bahia Honda Key [ANSP]; Big Pine Key [photo by Alejandro Lopez-Couto]; Key Biscane, Bill Baggs Cape Florida State Park [photo by Elizabeth Golden]); Florida (Miami Dade County [Rehn & Hebard, 1912: 243]; Everglades [Rehn & Hebard, 1912: 243]); Bahamas (South Bimini [AMNH]; Eleuthera [FSCA]; Nassau, New Providence [ANSP]; Andros [NHMUK]); Cayman Islands (Grand Cayman [NHMUK]; Little Cayman [NHMUK]) and Swan Islands [USNM]. Number of specimens examined: 177 TABLE 25. Measurements of Haplopus scabricollis (Gray, 1835) [mm] ♀♀ Loggerhead Key

♂♂ Loggerhead Key

♀ Bahamas [NHMUK]

♂ Bahamas [FSCA]

♂ var. Navassa Id. [USNM]

♀, PT of similis [USNM]*


125.0–139.0

-

156.0

-

-

134.0

Body

113.0–127.0

83.0–93.5

141.0

94.3

84.5

-

-

-

7.1

4.3

4.2

6.5

Mesonotum

26.0–28.0

18.9–20.0

32.7

20.5

18.8

29.5

Metanotum

6.8–8.0

5.1–6.0

9.8

5.8

5.0

7.5

Median segment

7.3–8.2

6.0–6.9

9.1

6.2

6.4

9.5

Tegmina

7.9–8.7

6.5–7.0

9.3

7.3

5.7

10.0

Alae

5.3–6.5

4.2–5.0

6.0

5.2

3.9

10.0

Profemora

17.8–20.0

18.8–20.0

24.4

17.9

19.3

21.5

Mesofemora

15.5–17.0

15.0–16.0

20.4

15.3

17.5

18.7

Metafemora

21.0–22.0

19.7–21.0

26.5

20.0

21.4

24.5

-

-

26.0

18.8

19.6

22.5

Pronotum

Protibiae



HENNEMANN ET AL.

TABLE 25. (Continued) ♀♀ Loggerhead Key

♂♂ Loggerhead Key

♀ Bahamas [NHMUK]

♂ Bahamas [FSCA]

♂ var. Navassa Id. [USNM]

♀, PT of similis [USNM]*

Mesotibiae

-

-

21.0

14.9

15.3

-

Metatibiae

-

-

29.6

20.5

21.0

-

Antennae

45.0–50.0

50.0–53.0

54.0

52.0

ca. 50.0

-

FIGURES 276–278. Haplopus sobrinus n. sp.. 276. ♀ HT: Cuba, Camagüey [ANSP]; 277. ♀ HT: head and pronotum [ANSP]; 278. ♀ HT: tegmina and alae [ANSP].

Haplopus sobrinus n. sp. (Figs. 276–278, 342) HT, ♀: Isla de Turiguanó, Camagüey, Nov. 1935, V.M. Aguirene, Col.; wO176 [ANSP].

Diagnosis: Females (the only sex known) are very close to H. scabricollis (Gray, 1835) but differ by: the more slender body; relatively longer legs and tarsi; relatively longer mesothorax; just minutely tuberculate mesonotum; very faint and blunt cephalad tubercles (Fig. 277); smooth abdominal tergites and slightly longer alae (Fig. 278). Etymology: The specific name “sobrinus” (lat. = inconspicuous) refers to the rather sombre appearance of ♀♀ of this new species. Description: ♀ (Fig. 276). The holotype lacks both front legs, the apices of the antennae and the extending portion of the subgenital plate. Moderately sized (body length including subgenital plate 123.0 mm) and very slender for the genus with long and slender legs. Colour of body greyish dark brown with the head and pronotum pale brown; otherwise with some paler mottling. Legs greyish brown with darker mottling. Antennae reddish brown. Eyes dark reddish brown. Spines of the thorax with the tips black. Tegmina and costal region of alae very dark brown with the venations of a slightly paler colour. Anal region of alae transparent with all major veins dark brown. Head: About 1.3x longer than wide and ovoid with the cheeks slightly convex. Vertex gently rounded and posteriorly armed with a pair of blunt unequal tubercles; behind these a few small granules (Fig. 277). Eyes circular and contained almost 3x in length of cheeks. Antennae broken in the holotype. Scapus about 2x longer than wide and gently narrowed towards the base. Pedicellus round in cross-section and roughly half as long as scapus. REVISION OF HAPLOPODINI


143

Thorax: Pronotum longer but narrower than head, about 1.6x longer than wide and roughly rectangular. Transverse median sulcus rather indistinct, gently curved and not reaching lateral margins of segment. Surface with numerous small spiniform tubercles (Fig. 277). Mesothorax about 3x longer than head and pronotum combined. Mesonotum with numerous spiniform tubercles of variable sizes; a row of small tubercles is present along lateral margins. Meso- and metapleurae each with a marginal row of spiniform tubercles. Mesosternum irregularly tuberculate, the metasternum with two pairs of small tubercles. Tegmina ovate with a very elongate but shallow central protuberance and very slightly projecting over posterior margin of metanotum. Alae ¾ the length of tegmina and reaching about half way along median segment (Fig. 278). Abdomen: Median segment about 1.2x longer than wide and gently constricted medially; smooth. All tergites unarmed dorsally. Segments II–VI roughly equal in length and width, all rectangular and about 1.8x longer than wide. Tergum VII slightly shorter and narrower than previous, about 2.1x longer than wide and slightly expanded posteriorly. Praeopercular organ formed by a tuberculate swelling near the slightly bi-lobate posterior margin of sternum VII (Fig. 342). Tergum VIII slightly shorter and distinctly narrower than VII, gently constricted medially and 2.4x longer than wide. IX slightly narrowed towards posterior and slightly less than half the length of VIII. Anal segment with a very faint longitudinal median carina with the posterior margin sub-truncate and with a very small median indentation. Epiproct small, acutely triangular and carinate dorsally. Cerci small and conical with a rather acute tip. According to Moxey (1972: 96) the subgenital plate is very elongate and naviculate. Legs: Mesofemora about 4/5 the length of mesothorax and metafemora reaching posterior margin of abdominal segment IV. Medioventral carina of mesofemora with four and in metafemora with five spines. Anteroventral carina of meso- and metafemora with two, posteroventral carina with one very small sub-apical spine. Basitarsi slightly longer than the following three tarsomeres combined. According to Moxey (1972: 96) the profemora are slightly shorter than the mesothorax and bear two small spines on the medioventral carina. Comments: This rather sombre and slender species is so far only known from the unique ♀ holotype in ANSP, ♂♂ and eggs still being unknown. The specimen was mistaken for P. cubensis (Saussure, 1868) and obviously served for the description and set of measurements presented for this species by Moxey (1972: 95ff.) in his unpublished PhD-thesis. Comparison shows the specimen to have suffered from considerable damage, since some body parts described by Moxey are now lacking in the specimen, e.g. the front legs and subgenital plate. Distribution: N-Cuba: Isla de Turiguanó [ANSP]. Number of specimens examined: 1 TABLE 26. Measurements of Haplopus sobrinus n. sp. [mm] HT, ♀ [ANSP] Body (incl. sg. pl.)

123.0*

Body

118.5

Pronotum

4.7

Mesonotum

28.5

Metanotum

6.8

Median segment

9.0

Tegmina

7.0

Alae

5.5

Profemora

24.0*

Mesofemora

21.0

Metafemora

27.0

Protibiae

-

Mesotibiae

19.3

Metatibiae

26.0

* according to Moxey (1972: 96) since the subgenital is now missing


HENNEMANN ET AL.

FIGURES 279–286. Haplopus woodruffi n. sp.. 279. ♀ PT: Cayman Islands, Cayman Brac [FSCA]; 2480 ♂ HT: Cayman Islands, Cayman Brac [FSCA]; 281. Head and pronotum of ♀ PT [FSCA]; 282. Head and pronotum of ♂ HT [FSCA]; 283. Apex of abdomen of ♀ PT (dorsal view) [FSCA]; 284. Apex of abdomen of ♀ PT (lateral view) [FSCA]; 285. Apex of abdomen of ♂ HT (dorsal view) [FSCA]; 286. Apex of abdomen of ♂ HT (lateral view) [FSCA].

Haplopus woodruffi n. sp. (Figs. 279–286, 363) HT, ♂: Cayman Islands: Cayman Brac, Bight Road at Maj. Donald Rd., 24-V-2009, coll.: R. H. Turnbow [FSCA]. PT, ♂: Cayman Islands: Cayman Brac, Bight Road at Maj. Donald Rd., 24-V-2009, coll.: R. H. Turnbow [FSCA]. PT, ♀: Cayman Islands: Cayman Brac, Bight Road at Maj. Donald Rd., 24-V-2009, coll.: M. C. Thomas [FSCA].

Diagnosis: Very close to H. scabricollis (Gray, 1835) but differing by: the averagely smaller size; smaller and blunt horns of the head (Figs. 281–282); relatively larger eyes and lack of a sub-apical dorsal tooth on the meso- and



145

metafemora; lack of a sub-basal tooth on the anterodorsal carina of the meso- and metatibiae; slightly longer tegmina and alae, the first distinctly projecting over the posterior margin of the metanotum and the latter in ♀♀ reaching about half way along the median segment with the anal region not distinctly red as in H. scabricollis. ♀♀ also differ by the slender probasitarsus, differently shaped anal segment and much smaller, triangular epiproct (Fig. 283). ♂♂ can be distinguished from those of H. scabricollis by: the lack of a white longitudinal marking along the lateral margins of the pronotum (Fig. 282) and median portion of the median segment; lack of a posterolateral tooth or lobe of abdominal tergum VII (Fig. 285); club-like, much more prominently swollen and broadened abdominal segments VIII–X (Fig. 285); shorter and dorsally globose anal segment (Fig. 286); smaller but conical basal projection of the poculum (Fig. 286) and differently shaped vomer, which has the terminal hook considerably shorter than in H. scabricollis (Fig. 355). Etymology: This new species is dedicated to Dr. R. E. Woodruff (FSCA) for arranging loan of many specimens from the collection of FSCA and his private collection. Description: ♀ (Fig. 279). Of medium size (body length including the subgenital plate 142.0 mm) and moderately slender for the genus; body surface slightly glabrous. General colour creamish mid brown with faint darker speckles on body. Ventral body surface with a pale greyish wash. Head with two small, elongate black markings above the eyes. Cheeks irregularly furnished with white and dark brown (Fig. 281); two small black spots on frons. Antennae greyish straw, the apical portion with three dark transverse bands, otherwise about every fourth segment with a dark apex. Eyes pale ochraceous grey. Tubercles and spines of the thorax with black points. Tegmina and costal region of alae dark brown with the venations pale cream; the latter black basally. Anal region of alae transparent with a slight pinkish wash and all major longitudinal and transverse veins marked with black. Head: About 1.3x longer than wide and ovoid with the cheeks moderately convex. Vertex rounded and armed with a pair of moderately sized, blunt and conical cephalad tubercles (Fig. 281). A further pair of small acute granules posteriorly. Eyes circular and contained about 1.8x in length of cheeks. Antennae almost reaching posterior margin of median segment; consisting of 61 segments. Scapus about 2x longer than wide and slightly constricted towards the base. Pedicellus roughly half the length of scapus and 2/3 the length of III. Thorax: Pronotum longer but narrower than head, almost 2x longer than wide, roughly rectangular and with the lateral margins roundly emarginated medially. Transverse median sulcus rather decided, gently curved and not reaching lateral margins of segment. Dorsal surface with a two pairs of minute spines in the anterior portion and a pair of small spines near posterior margin; otherwise with a variable number of small rather irregularly disposed tubercles (Fig. 281). Anterior margin raised and with four spiniform tubercles. Mesothorax about 2.6x longer than head and pronotum combined. Mesonotum very slightly gradually widened towards the posterior and with a fine but acute longitudinal median carina; surface sparsely set with irregularly disposed small, spiniform tubercles and two pairs of small spines in anterior portion. Lateral margins with a marginal row of 10–12 spiniform tubercles. Meso- and metapleurae armed with an irregular longitudinal marginal row of small but pointed spines. Mesosternum with a few scattered granules and metasternum smooth except for two very small granules medially. Tegmina oval, coriaceous, with the venation very distinct, dense and irregularly disposed; reaching about ¼ along median segment. The median protuberance very shallow and conspicuously displaced towards the apex of tegmen. Alae a little shorter than tegmina and reaching only about half way along median segment. Abdomen: Median segment about 1.7x longer than wide and very gently widened towards the posterior. All segments unarmed, except for a pair of small posteromedian spines on tergum II. Segments II–VI roughly equal in length and width, all rectangular and about 2x longer than wide. Tergum VII slightly shorter and narrower than previous, about 2.2x longer than wide with lateral margins very slightly deflexed posteriorly. Praeopercular organ formed by a rounded sepia-coloured tubercle, which posteriorly extends into a longitudinal furrow, leaving a median gap in the posterior margin of sternum VII. Tergum VIII slightly shorter than VII, narrowed mediallly, almost 3x longer than wide and 2.5x longer than IX. IX roughly rectangular. Anal segment with a very faint longitudinal median carina, the posterior margin slightly deflexed, broadly rounded and with a small posteromedian indentation. Epiproct very small, triangular (Fig. 283). Cerci very small, conical and with an acute tip; not reaching to posterior margin of anal segment. Subgenital palte very long, lanceolate, longitudinally keeled over entire length and with a narrow but rounded apex; extending over abdomen by almost the combined length of tergites VIII–X (Figs. 283–284). Legs: Profemora about 4/5 the length of mesothorax, mesofemora slightly projecting over posterior margin of median segment and metafemora reaching about half way along abdominal segment IV. Profemora unarmed.


HENNEMANN ET AL.

Anteroventral carina of meso- and metafemora with two, posteroventral carina with one sub-apical spine; medioventral carina armed with 4–6 pointed and fairly strong spines. Both dorsal carinae of meso- and metafemora smooth but very gently elevated sub-apically. All tibiae unarmed. Basitarsi roughly equal in length to following three tarsomeres combined. ♂ (Fig. 280). Fairly small (body length 76.4–80.0 mm) and stocky for the genus with conspicuously shortened alae (length 4.7–5.5 mm). General colour of dorsal body surface pale ochre with a greenish hue. Head, meso- and metapleurae, most of ventral body surface and legs green. Lateral margins of abdominal tergites II–VII each with a white anterior marking, largest on II and gradually decreasing in size towards VII (Fig. 280). Tergites VIII and IX with entire lateral margins broadly white and anal segment with a small white marking anterolaterally. Points of thoracal spines black. Eyes pale reddish brown, antennae ochre with a slight reddish wash in basal portion. Tegmina greenish cream in central portion with the anterior and posterior margins broadly white. Costal region of alae greenish cream interiorly, whitish anteriorly and black towards the base. Anal region transparent with a slight pinkish wash and all major longitudinal and transverse veins marked with dark black. Tarsi pale brown. Head: Generally as in ♀♀ but eyes much more prominent, projecting hemispherically and their length contained only about 1.1x in that of cheeks. Vertex smooth except for the two small, blunt cephalad tubercles (Fig. 282). Antennae moderately robust and slightly projecting over abdominal segment II; otherwise as in ♀♀. Thorax: Pronotum slightly longer and about as wide as head, general shape as in ♀♀; surface smooth except for one (paratype) or two (holhotype, Fig. 282) moderate pairs of spines in the anterior portion. Probasisternum smooth. Mesothorax about 2.4–2.5x longer than head and pronotum combined. Mesonotum with 6–8 paired spines of moderate size in the anterior 2/3; the one (holotype, Fig. 282) or two (paratype) pairs near anterior margin largest. Mesosternum smooth except for 2–4 green granules in anterior portion and occasionally a further small pair of granules medially; metasternum smooth. Mesopleurae with a few small granules in anterior portion, metapleurae smooth. Tegmina broadly oval and very slightly projecting over posterior margin of metanotum, central protuberance very shallow. Alae small, slightly shorter than alae and almost reaching half the way along median segment. Abdomen: Segments II–V roughly equal in length and about 3.3x longer than wide. VI and VII decreasing in length with VII no more than 2.3x longer than wide; posterior portion of tergum VII very gently widened. All tergites and sternites smooth. Tergites VIII–X conspicuously swollen and about 1.7x broader than previous segments (Fig. 285). VIII about ¾ the length to VII, trapezoidal and strongly widening towards the posterior. IX about equal in length to VIII and gently narrowed towards the posterior. Anal segment shorter than IX, roundly rectangular with the dorsal surface roundly convex (Fig. 286). Posterior margin broadly rounded with a very shallow median indentation (Fig. 285); slightly swollen and on ventral surface armed with several small, black incurving denticles. Epiproct very small. Vomer with a broad base and a moderately elongate, conical, gently upcurving terminal hook which is about equal in length to basal portion (Fig. 363). Cerci large, obtuse and scarcely shorter than anal segment (Fig. 286). Poculum convex, cup-like with a small but acutely conical basal hump (Fig. 286). Legs: Profemora scarcely shorter than mesothorax, mesofemora about 4/5 the length of mesothorax and metafemora reaching to anterior of abdominal segment V. All less carinate than in ♀♀ but armature generally alike. Tarsi relatively more elongate and basitarsi a little longer than following three tarsomeres combined. Distribution: Cayman Islands (Cayman Brac: forest along Major Donald Drive) [FSCA]. Number of specimens examined: 3 TABLE 27. Measurements of Haplopus woodruffi n. sp. [mm] HT, ♂ [FSCA]

PT, ♂ [FSCA]

PT, ♀ [FSCA]

-

-

142.0

Body

80.0

76.4

137.5

Pronotum

4.0

3.9

6.0

Mesonotum

17.3

15.9

28.2





147

TABLE 27. (Continued) HT, ♂ [FSCA]

PT, ♂ [FSCA]

PT, ♀ [FSCA]

Metanotum

5.1

5.0

6.8

Median segment

5.9

5.8

8.8

Tegmina

6.0

5.0

11.0

Alae

5.5

4.7

9.0

Profemora

16.1

14.8

20.0

Mesofemora

14.3

12.9

16.2

Metafemora

18.8

18.0

22.9

Protibiae

16.0

15.0

20.8

Mesotibiae

12.2

11.0

16.1

Metatibiae

17.4

16.0

24.0

Antennae

43.5

44.0

56.0

5.6. Genus Paracranidium Brock, 1998 Type-species: Diapherodes (Cranidium) pumilio Westwood, 1843: 50, pl. 61: 2 (♀), by original designation of Brock, 1998b: 28. Paracranidium Brock, 1998b: 28. Bragg, 2001: 639. Zompro, 2004: 317. Otte & Brock, 2005: 246. Zompro, 2005: 135. Cranidium (?) Kirby, 1904a: 361 (in part). Diapherodes (Cranidium) Westwood, 1843: 50 (in part). Monandroptera, Westwood, 1859: 80 (in part). Phereboea Moxey, 1972: 153 (in litt).

Description: ♀♀ (Fig. 287). Very small apterous Haplopodini (body length 50.0–54.0 mm). Body moderately broad with mesothorax laterally expanded, triangular in cross-section. Body surface rugose and granulate. Head longer than wide, prognathous, vertex flat and granulose. Antennae longer than head and complete thorax combined. Scapus dorsoventrally compressed, pedicellus subcylindrical. Pronotum longer than wide, rectangular and distinctly shorter and narrower than head. Posterior margin raised medially, otherwise granulose. Mesothorax elongate, about 1.3x longer than head and pronotum combined. Mesonotum strongly dilated just behind the anterior margin and with a very prominent, longitudinal median carina which is semicircularly elevated in anterior half of segment (Fig. 287). Mesopleurae simple, granulose. Metathorax slightly tapering towards the posterior. Metanotum and median segment with a longitudinal median carina; transverse suture between these segments absent. Mesosternum smooth. Abdomen with all tergites medially carinate. Segments II–VI roughly parallel-sided, subequal in length and slightly transverse. VII shorter than previous and narrowed posteriorly. VIII–X tapering to the apex. Anal segment slightly longer than IX and rounded apically (Fig. 289). Epiproct very small, triangular. Cerci small and slender, tapering towards the apex. Subgenital plate elongate and scarcely projecting over posterior margin of anal segment, apex acuminate (Fig. 288). Legs slender and of moderate length; profemora slightly longer than mesothorax, metafemora reaching about half way along tergum IV. Profemora strongly compressed and curved basally, the medioventral carina slightly displaced towards the anteroventral carina. Meso- and metafemora minutely spinulose ventrally; otherwise legs unarmed. Basitarsi elongate, as long as or slightly longer than remaining tarsomeres combined. ♂♂ and eggs unknown. Differentiation: Females differ from all other genera of Haplopodini by: the small size, elongate, prognathous head, and triangular body cross-section having the complete dorsal body surface with a distinct longitudinal median keel which is semicircularly elevated in the mesonotum (Fig. 287).


HENNEMANN ET AL.

Comments: The thickened antennae, simple mesosternum, ventrally spinulose meso- and metafemora, not conspicuously laterally dilated abdomen and short gonapophyses VIII clearly place Paracranidium in Haplopodini. Several characters show similarity to Venupherodes n. gen. from Cuba, which also has a distinct longitudinal median carina running along the dorsal body surface. The elongate and flat head resembles Aploploides Rehn & Hebard, 1938 also from Cuba. Males and eggs unknown. Distribution (Fig. 378): Mountainous regions of central Jamaica (Newton 3000 ft. [ANSP, Moxey, 1972: 153] & Cinchona 5000 ft. [ANSP, Moxey, 1972: 153]), endemic.

Species included: 1. Paracranidium pumilio (Westwood, 1843: 50, pl. 61: 2 (♀)) [Diapherodes (Cranidium)].

Paracranidium pumilio (Westwood, 1843) (Figs. 287–289, 378) Diapherodes (Cranidium) pumilio Westwood, 1843: 50, pl. 61: 2 (♀). HT, ♀: no data “Africa tropicali?” [BMAG]. Cranidium (?) pumilio, Kirby, 1904a: 361. Monandroptera pumilio, Westwood, 1859: 80. Paracranidium pumilio, Brock, 1998: 28, fig. 2. Otte & Brock, 2005: 246. Phereboea pumilio, Moxey, 1972: 153 (in litt.). Further material [1 ♀]: JAMAICA: 1 ♀: 240, 47–62 Jamaica, Diapherodes (Cranidium) pumilio Westw. are Ent t 61 f.2, Cranidium pumilio Westw. Jamaica [NHMUK].

FIGURES 287–289. Paracranidium pumilio (Westwood, 1843). 287. ♀: Jamaica [NHMUK]; 288. Apex of ♀ abdomen (lateral view) [NHMUK]; 289. Apex of ♀ abdomen (dorsal view) [NHMUK].

Comments: Diapherodes (Cranidium) pumilio Westwood, 1843 was omitted in the monograph by Redtenbacher (1908). Brock (1998) transferred this distinctive species to the newly described genus Paracranidium and clarified the depository and identity of Westwood's holotype, which was previously believed lost. Kirby (1904a: 361) erroneously referred to the NHMUK ♀ as Westwood's type specimen. This was however collected in 1847, four REVISION OF HAPLOPODINI


149

years after Westwood's description was published and hence cannot be the type. The holotype in BMAG was only examined from photos kindly provided by Paul D. Brock (NHMUK). The two ♀♀ stated to be in ANSP by Moxey (1972: 153) were not traced. Measurements of the ♀ in NHMUK are provided below. Males and eggs unknown. Number of specimens examined: 2 TABLE 28. Measurements of Paracranidium pumilio (Westwood, 1843) [mm] ♀ [NHMUK] Body

54.5

Pronotum

2.8

Mesonotum

12.2

Metanotum

3.8

Median segment

3.2

Profemora

13.5

Mesofemora

9.3

Metafemora

12.1

Protibiae

12.4

Mesotibiae

8.0

Metatibiae

12.5

5.7. Genus Parhaplopus n. gen. Type-species: Haplopus cubensis Saussure, 1868: 68, by present designation. Haplopus, Westwood, 1859: 85, pl. 18: 6, 6a & b (in part). Saussure, 1868: 68. Kirby, 1904a: 363, 364 (in part). Redtenbacher, 1908: 429 (in part). Otte & Brock, 2005: 150 (in part).

Description: ♂♂, ♀♀. Large (body length ♀♀ 117.5–164.5 mm, ♂♂ 78.0–99.2 mm), slender and elongate Haplopodini, ♀♀ brachypterous, ♂♂ with fully developed alae. Body sub-cylindrical (♀♀) or cylindrical (♂♂) in cross-section. Colouration of ♀♀ various shades of brown and grey. ♂♂ rather colourful insects, mostly green or greenish brown with body surface slightly glabrous. Head distinctly longer than wide, vertex very gently convex and moderately bi-spinose. Antennae moderately thickened; in ♀♀ reaching to median segment, in ♂♂ at least to abdominal segment II. Pronotum slightly shorter than the head, rectangular and sparsely tuberculose; entirely unarmed in ♂♂. Mesothorax moderately elongate and at best 2.3x longer than head and pronotum combined; parallel-sided in ♂♂ and medially swollen in ♀♀. Mesonotum of ♀♀ convex medially and rather irregularly armed with prominent, spiniform tubercles or blunt spines of various sizes; those in the raised central portion of the mesonotum largest and with a very broad base. Mesonotum of ♂♂ armed with 8–12 pointed spines. Mesopleurae with a longitudinal row of ± distinct tubercles and mesosternum irregularly granulose (♂♂) or minutely tuberculose (♀♀). Metasternum sparsely granulose in ♀♀ and smooth in ♂♂. Tegmina elongate, oval and with a moderately distinct central hump in ♂♂; reaching almost half way along median segment. Alae of ♀♀ indistinctly longer than tegmina; of ♂♂ well developed and reaching to abdominal segment VI. Anal region reticulate with distinct brown to black radial and transverse veins in ♀♀, translucent orange or pink with only the marginal portion slightly reticulate in ♂♂. Abdomen considerably longer than head and thorax combined. Median segment longer than metanotum, smooth. Segments II–VII distinctly longer than wide; II–VI ± rectangular. Tergites unarmed in ♂♂, II–IV sometimes with a ± distinct pair of posterior spines in ♀♀. VII with lateral margins ± rounded and elevated posteriorly (♀♀ in particular). VIII–X distinctly narrower (♀♀) or slightly wider (♂♂) than previous segments. Sternites II–VII smooth, except for a very few small granules on II–III of ♀♀. Praeopercular organ of


HENNEMANN ET AL.

♀♀ indistinct and formed by a faint and short longitudinal median ridge close to posterior margin of sternum VII. Anal segment longitudinally carinate and the posterior margin with a small median notch; tapered in posterior portion in ♀♀. Epiproct very small, roundly triangular and almost completely hidden under anal segment in ♀♀. Vomer of ♂♂ prominent, broadly triangular and with a moderately long, acute terminal hook. Cerci very small, straight and tapered towards the pointed apex in ♀♀; obtuse, about 2/3 the length of anal segment and laterally compressed in ♂♂. Poculum of ♂♂ moderately convex, cup-like. Subgenital plate of ♀♀ very long, naviculate with the apex rounded or broadly truncate; projecting over apex of abdomen by at least the combined length of tergites VIII–X. Legs of moderate length, rather robust in ♀♀; pro- and mesofemora shorter (♀♀) or slightly longer (♂♂) than mesothorax. Hind legs ± reaching (♂♂) or clearly not reaching (♀♀) to the apex of abdomen. Two outer ventral carinae of meso- and metafemora each with 1–2 sub-apical spines and very minutely denticulose, the medioventral carina of these femora armed with a longitudinal row of five rather prominent spines. Dorsal carinae of all femora unarmed, those of the meso- and metafemora sometimes with a shallow sub-apical tooth in ♀♀. Tibiae unarmed except for a few small teeth in the apical portion of the medioventral carina; anterodorsal carina of ♀♀ undulate and lobate. Basitarsi of moderate length, longer than following two tarsomeres combined. Dorsal carina of probasitarsus with a rounded lobe. Eggs (Figs. 320–321): Of moderate size (capsule length 3.8 mm), barrel-shaped and cylindrical in crosssection; capsule about 2x longer than wide. In lateral aspect, dorsal surface of capsule considerably more convex than ventral surface. Polar-area impressed. Capsule surface strongly granulose and covered with irregular ridges. Micropylar plate small heart-shaped and < 1/3 as long as capsule; posterior end with a wide median gap. Operculum flat, circular and in the centre with prominent, raised tubercles and hump-like projections; no conspicuous central capitulum. Colouration brown. Differentiation (Table 29): Very closely related and certainly the sister-taxon of Haplopus Burmeister, 1838. The eggs represent the most striking distinctive features, being barrel-shaped with the capsule surface distinctly tuberculose and covered with irregular ridges, the polar-area impressed and the operculum lacking the hat or knoblike extension seen in Haplopus. Adult insects may be distinguished by: the more elongate and distinctly less convex head, which merely bears two small cephalad spines or tubercles; medially swollen mesothorax; apically rounded or truncate subgenital plate and dorsally lobed probasitarsus of ♀♀; as well as the plain anal region of the alae and smaller, laterally compressed cerci ♂♂. For a detailed comparison and distinction from Haplopus see Table 29 below. Comments: The status of a subgenus of Haplopus Burmeister, 1838 was originally considered for the three species here contained in Parhaplopus n. gen.. In addition to a number of morphological characters of the insects the very distinctive eggs however appear to justify the status of Parhaplopus as a generic unit. Unfortunately, only a very few specimens are yet known. Distribution (Fig. 378): Cuba, Gonaive Island and Hispaniola. Etymology: Masculine. A combination of the prefix “Par(a)-“ (gr. = very similar) and the generic name Haplopus Burmeister, 1838 to emphazise the close relation to this genus. Species included: 1. Parhaplopus cubensis (Saussure, 1868: 68) [Haplopus]. n. comb. [Distribution: Cuba] 2. Parhaplopus evadne (Westwood, 1859: 85, pl. 18: 6, 6a & b (♂)) [Haplopus]. n. comb. [Distribution: W-Hispaniola] 3. Parhaplopus navarroi n. sp. [Distribution: Hispaniola]

TABLE 29. Comparison and differentiation of Parhaplopus n. gen. and Haplopus Burmeister, 1838

Head

Parhaplopus n. gen.

Haplopus

Longer than wide; vertex at best gently rounded and armed with two small spines or tubercles

Globose and indistinctly longer than wide; vertex ± convex and distinctly bi-cornute




151

TABLE 29. (Continued) Parhaplopus n. gen.

Haplopus

Unarmed

With a ± distinct pair of anterior tubercles / spines

Slightly swollen medially; armed with several large and blunt tubercles

Slender, not swollen medially; surface granulose or tuberculose

Plain

Reticulate

Laterally compressed; apex narrowed

Cylindrical; apex rounded to club-like

Poculum (♂♂)

Moderately convex; at best with a small hump basally

Strongly convex and with a ± distinct basal spine

Subgenital plate (♀♀)

Apex rounded to broadly truncate

Apex narrow and ± pointed

With a rounded dorsal lobe

Slender

Barrel-shaped

Ovoid

Distinctly coriaceous / tuberculose and covered with irregular ridges

Minutely granulose / rugulose to almost smooth

Impressed

Convex and rounded

Pronotum (♂♂) Mesothorax (♀♀) Anal region of alae (♂♂) Cerci (♂♂)

Probasitarsus (♀♀) Egg (capsule) Egg (capsule surface) Egg (polar area) Egg (micropylar plate)

Small; < 1/3 the length of capsule

> ½ the length of capsule

Egg (operculum)

Flat and with an irregularly raised circular rim of tubercles

Convex, either conical or knob-like

Keys to the species of Parhaplopus n. gen. ♀♀ 1. 2. -

Subgenital plate naviculate to lanceolate with the apex bluntly rounded to pointed; Hispaniola. . . . . . . . . . . . . . . . . . . . . . . . . 2 Subgenital plate gradually broadened towards a roundly truncate apex (Fig. 293); Cuba . . . . . . . . . . . . . . . . . . . . . . . cubensis Body length > 140.0 mm; cephalad horns prominent and acutely pointed (Fig. 303); pronotum multi-spinose (Fig. 303); mesonotum slender (Fig. 298); subgenital plate lanceolate with apex pointed (Fig. 308) . . . . . . . . . . . . . . . . . . . . . . . . . evadne Body length 117.5 mm; cephalad horns small, conical (Figs. 312–313); pronotum sparsely tuberculose (Fig. 312); mesonotum widened medially (Fig. 310); subgenital plate with apical half gently widened and apex bluntly rounded (Fig. 316) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . navarroi n. sp.

♂♂ 1. 2. -

Vertex with a pair of small spines; tegmina and costal region of alae green with a brown stripe along anterior margins . . . . . . 2 Cephalad spines prominent, conical (Figs. 301–302); tegmina and alae yellow with all longitudinal veins green (Fig. 298); Hispaniola & Gonaive Island . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . evadne Head unicolourous (Fig. 297); abdominal tergum VII deflexed laterally; Cuba . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . cubensis Cheeks white (Fig. 314); abdominal tergum VII parallel-sided; Hispaniola. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . navarroi n. sp.

Parhaplopus cubensis (Saussure, 1868) n. gen., n. comb. (Figs. 290–297, 344) Haplopus cubensis Saussure, 1868: 68. HT, ♀: Cuba; Haplopus cubensis Sauss. [MHNG]. Saussure, 1871–1872: 192. Bolívar, 1888: 40. Kirby, 1904a: 364 (in part). Redtenbacher, 1908: 433. Otte & Brock, 2005: 151. Aplopus cubensis, Rehn, 1909: 200 (in part). Zompro & Brock, 2003: 10. Diapherodes cubensis, Moxey, 1972: 95 (in litt; in part—not ♀ from Turiguano, Cuba). [Description of ♂]


HENNEMANN ET AL.

Further material [1 ♂, 1 ♀]: CUBA: 1 ♀: Museum Paris, Cuba, La Havane, P. Serre 1910; Haplopus cubensis Sauss., L. Chopard & L. Bertrand 191 [MNHN]; 1 ♂: Cuba, Mr. H. d. Saussure; Haplopus evadne Westw. [MHNG].

Diagnosis: Similar to P. navarroi n. sp. from Hispaniola but differing by: the larger size and relatively longer mesothorax of both sexes; less distinct pair of tubercles on the vertex (Fig. 296); tuberculate abdominal segments; more decidedly posterolaterally expanded abdominal tergum VII (Fig. 293); broadly truncate apex of the subgenital plate (Fig. 293) and more undulate dorsal carinae of the protibiae of ♀♀. Males differ by the plain greenish brown head (Fig. 297) and more robust legs. Description: ♀ (Fig. 290–291). Large (body length including the subgenital plate 164.5 mm) and characteristic for the broadly truncate apex of the subgenital plate. Colour greyish to ochraceous brown, with irregular pale grey mottling on dorsal surface of abdomen, meso- and metasternum. Metapleurae and ventral surface of abdomen pale greyish. Antennae pale brown dorsally and reddish mid brown ventrally. Eyes dull ochraceous. Spines of the thorax dull yellow to ochre, those of the mesonotum and pleurae with dark brown tips. Tegmina and costal region of alae greyish mid to dark brown with slight darker mottling in the basal and lateral portions; the venations ochre. Anal region of alae transparent drab and all major longitudinal and transverse veins broadly marked with dark brown. Head: Ovoid, about 1.3x longer than wide; vertex gently convex and armed with a pair of small cephalad tubercles (Fig. 296); a few small tubercles present near posterior margin. Eyes fairly prominent, circular and contained slightly more than 2x in length of cheeks. Antennae broken in both specimens available, but at least reaching to posterior of metanotum and consisting of > 50 segments. Scapus 2.2x longer than wide with the lateral margins gently rounded and slightly constricted towards the base. Pedicellus about half the length of scapus and longer than III, roughly cylindrical. Thorax: Pronotum about equal in length but slightly narrower than head, 1.6x longer than wide, roughly rectangular and with the lateral margins gently emarginated medially. Transverse median sulcus distinct, gently curved and almost reaching lateral margins of segment. Dorsal surface with a pair of spiniform tubercles just in front of the median sulcus and in posterior portion with several small, irregularly disposed tubercles (Fig. 296). Probasisternum with a transverse row of four pointed tubercles. Profurcasternum with a few small tubercles along lateral margins. Mesothorax about 2.2x longer than head and pronotum combined, constricted at the anterior and conspicuously swollen medially. Mesonotum slightly widening towards the mid of segment, median portion swollen and in posterior portion almost parallel-sided. Surface with a fine longitudinal median carina and with a good number of very prominent, irregularly paired, conical spines of variable sizes; usually with two strongly enlarged pairs of spines in anterior half (Fig. 296). Along lateral margins with a marginal row of about 16 pointed spines. Mesopleurae armed with a longitudinal row of about 20 rather prominent and acute spines, metapleurae merely with a marginal row of eight tubercles. Mesosternum all over covered with irregularly disposed short spines, metasternum only with a few tubercles. Tegmina broadly ovate, coriaceous, with the venation very distinct, dense and irregularly disposed; roughly reaching to posterior margin of metanotum. The median protuberance very shallow. Alae a little longer than tegmina and reaching about 2/3 the way along median segment. Abdomen: Median segment 1.8x longer than wide and very gently narrowed medially, in posterior portion with two small granules. Tergites II and III each with a few scattered tubercles, II–V with a transverse row of four tubercles along posterior margin and sternites II–IV each with a small median pair of tubercles; remaining segments unarmed. Segment II about 1.2x longer than median segment but shorter than III, rectangular and about 1.7x longer than wide. III–VI roughly equal in length and slightly longer than II, on average about 1.6x longer than wide. Tergum VII shorter and narrower than VI and about 1.7x longer than wide, lateral margins roundly deflexed posteriorly (Fig. 293). Sternum VII with the praeopercular organ formed by an elongate wart-like structure near posterior margin (Fig. 344). Tergum VIII about ¾ the length of VII, gently narrowed medially and roughly 1.4x longer than wide. IX almost rectangular and about half the length of VIII. Anal segment with a very faint longitudinal median carina, narrowed in posterior half and with a deep but narrow posteromedian emargination; epiproct very small (Fig. 295). Cerci very small and conical. Subgenital plate very long, naviculate, longitudinally carinate, gradually flattened and widened towards a broad and roundly truncate apex; extending over abdomen by more than the combined length of tergites VIII–X (Figs. 293–294).



153

FIGURES 290–292. Parhaplopus cubensis (Saussure, 1868) n. gen., n. comb.. 290. ♀: Cuba, Havanna [MNHN]; 291. ♀ HT: Cuba [MHNG]; 292. ♂: Cuba [MHNG].


HENNEMANN ET AL.

FIGURES 293–297. Parhaplopus cubensis (Saussure, 1868) n. gen., n. comb.. 293. ♀ apex of abdomen: Cuba, Havana (dorsal view) [MNHN]; 294. ♀ apex of abdomen: Cuba, Havana (lateral view) [MNHN]; 295. Anal segment and epiproct of ♀ HT (dorsal view) [MHNG]; 296. Head, pro- and mesothorax of ♀ HT [MHNG]; 297. Head and pronotum of ♂ [MHNG].

Legs: Profemora about 3/5, mesofemora about 2/3 the length of mesothorax and metafemora just not reaching posterior margin of abdominal segment III. Medioventral carina of profemora with 7–9 small spines, in meso- and metafemora armed with five very prominent and strong, back-curving spines. Anterodorsal carina of profemora strongly raised and slightly undulate, the anterodorsal carina of the protibiae lamellate and furnished with four rounded lobes. Anteroventral carina of meso- and metafemora with two, posteroventral carina with one strong subapical spine; both carinae minutely granulose. Both dorsal carinae with a slight triangular lobe-like tooth subapically. Anterodorsal carina of meso- and metatibiae with a roundly triangular elevation sub-basally and subapically. Probasitarsus about as long as the following two tarsomeres combined, dorsal carina rounded. Meso- and metabasitarsi simple and just a little longer than 2nd tarsomere. ♂ (Fig. 292). Unfortunately, the terminal abdominal segments are broken off and missing in the only specimen available. Of moderate size and fairly slender for the genus. Colour greenish brown, the posterior portion of the pronotum and bases of the profemora dull green. Ventral surface of body and profemora mostly whitish to pale grey with darker speckles, metapleurae green. Cheeks with a slight dark brown wash (Fig. 297). Apices of all femora mid brown. Dorsal spines of pro- and mesothorax with black tips, those of the mesopleurae and sternum dark olive. Tegmina and costal region of alae with anterior margin white, then a broad brown longitudinal band, and innermost portion green. Anal region of alae pale transparent pink with a slight greyish hue along the outer margin. Antennae dark ochracheous with a few dark brown transverse bands, tarsi pale brown. Head: Generally as in ♀♀ but vertex flattened and smooth except for two small cephalad spines (Fig. 297); eyes much more prominent, projecting hemispherically and their length contained only about 1.8x in that of cheeks. Antennae moderately robust and projecting over posterior margin of median segment; consisting of 61 segments. Otherwise as in ♀♀. Thorax: Pronotum slightly shorter and narrower than head with the lateral margins gently concave medially; surface smooth (Fig. 297). Transverse median sulcus shallow, gently rounded and just not reaching lateral margins of segment. Probasisternum with a small median pair of granules. Mesothorax about 1.9x longer than head and pronotum combined. Mesonotum armed with about ten distinct paired spines in the anterior 2/3. Mesopleurae with a longitudinal row of acute tubercles, metapleurae only with a few small granules. Mesosternum with numerous irregular low spiniform tubercles; metasternum smooth. Tegmina oval and reaching about ¼ the way along median



155

segment, central protuberance roundly conical. Alae almost reaching to posterior margin of abdominal segment VI (46.0 mm). Abdomen: Segments II-V roughly equal in length and about 3.6x longer than wide; all unarmed. VI slightly shorter than previous and hardly 3x longer than wide. Remaining segments missing in the unique specimen available. Legs: Legs all fairly short and moderately robust. Profemora very slightly longer, and mesofemora a little shorter than mesothorax, metafemora projecting a little over posterior margin of abdominal segment III. All less carinate than in ♀♀ and lacking any dorsal elevations or teeth. Armature of medioventral carina of the femora as in ♀♀ but considerably less developed. Tarsi relatively more elongate, probasitarsus almost equal in length to following three tarsomeres combined, meso- and metabasitarsus about as long as combined length of following two segments. Dorsal carina of probasitarsus gently rounded. Comments: Brief examination of the ♀ recorded by Moxey (1971: 95, in litt.) from Isla de Turiguano in ANSP has shown this to be a distinct species, hence this is an erroneous record (→ Haplopus sobrinus n. sp.). Eggs unknown. Distribution: Cuba (Camagüey & Oriente, Santiago de Cuba [USNM; Moxey, 1972: 97, in litt.]); NW- Cuba (Havanna) [MNHN]. Number of specimens examined: 3 TABLE 30. Measurements of Parhaplopus cubensis (Saussure, 1868) n. gen., n. comb. [mm] ♀, HT [MHNG]

♂ [MHNG]

-*

-

139.6

-

Pronotum

5.7

3.3

Mesonotum

28.3

13.8

Metanotum

3.9

6.2

Median segment

12.8

8.1

Tegmina

9.0

10.2

Alae

10.0

46.0

Profemora

19.1

14.1

Mesofemora

18.0

13.2

Metafemora

23.9

17.8

Protibiae

16.0

13.8

Mesotibiae

15.1

10.4

Metatibiae

21.7

14.7

Antennae

ca. 38.0

ca. 38.0


* ♀ in MNHN measures 164.5 mm

Parhaplopus evadne (Westwood, 1859) n. gen., n. comb. (Figs. 298–309, 355, 379–398) Haplopus evadne Westwood, 1859: 85, pl. 18: 6, 6a & b (♂). HT, ♂: Type, St. Dom. 55.1., Haplopus Evadne Westw., St. Domingo, evadne Westw. Mon. Phasm. Pl. 18 fig. 6, Haplopus evadne Westwood, HT; BMNH(E) #844943 [NHMUK]. Brock et al., (in press) Kirby, 1904a: 363. Redtenbacher, 1908: 432. Otte & Brock, 2005: 151. Diapherodes evadne, Moxey, 1972: 97 (in litt.).


HENNEMANN ET AL.

[Not: Haplopus evadne Caudell, 1904: 949, figs. 1 & 2 (♀)—misidentification. Later described as Aplopus mayeri Caudell, 1905 = A. scabricollis (Gray, 1835)] Further material [6 ♂♂, 1 ♀]: HISPANIOLA (HAITI): 1 ♂: Gonaive Is., Haiti, Aug., Eyerdam coll.; wO130 [ANSP]; 1 ♂: Haiti: Dept. Sud'est, Formond, alrededores Kay, Michel 2026684 mN, 603254 mE, 1100 m, 4–8 February 2006, coll. R. Bastardo; 33514 (MHND) [USNM]; 1 ♀: Haiti: Department du Sud, Formon, alrededores de Kay Michel, 1100 m, 603254 mE, 2026684 mN, 9.II.2006, R. Bastardo; 35424 (MHND) [USNM]; 1 ♂: Haiti: Department du Sud, Formon, alrededores de Kay Michel, 1100 m, 603254 mE, 2026684 mN, 9.II.2006, R. Bastardo; 35403 (MHND) [USNM]; 1 ♂: Haiti: Department du Sud, Formon, alrededores de Kay Michel, 1100 m, 603254 mE, 2026684 mN, 9.II.2006, R. Bastardo; 38600 (MHND) [USNM]; 1 ♂: Haiti: Department du Sud, Formon, alrededores de Kay Michel, 1100 m, 603254 mE, 2026684 mN, 9.II.2006, R. Bastardo; 33594 (MHND) [USNM]; 1 ♂: Haiti: Department du Sud, Formon, alrededores de Kay Michel, 1100 m, 603254 mE, 2026684 mN, 9.II.2006, R. Bastardo; 35484 (MHND) [USNM].

Diagnosis: Males readily differ from the other two species in the genus by: the large size; much more prominent cephalad horns; pale green colouration and yellow tegmina and alae which have all longitudinal veins broadly marked with bright green. Females resemble those of P. navarroi n. sp. but differ by: the larger size; much more prominent cephalad horns; more prominently spinose pronotum and mesothorax; more decided spines of the metapleurae; larger, semi-circular and shield-shaped epiproct and slender apically pointed subgenital plate. Description: The colouration is described from photos of live specimens taken at Parc National Pic Macaya, Massif de la Hotte. ♀ (Fig. 298). Fairly large (body length including the subgenital plate 149.0 mm) and fairly slender for the genus. Colour greenish mid brown all over mottled with irregular dark brown markings. Abdominal tergites III–VI with a pale cream roughly square marking anteriorly. Legs dull greenish brown. Entire thoracic armature tipped with black. Antennae dark reddish to greenish brown with most antennomeres, except for the scapus, dull brown to black apically. Eyes cream with reddish brown mottling. Tegmina and costal region of alae dull ochraceous with faint darker mottling. Anal region of alae transparent grey with all major longitudinal and transverse veins broadly marked with black. Head: Ovoid, about 1.3x longer than wide; vertex gently convex and armed with a pair of fairly large conical, apically pointed and black tipped cephalad horns; the dextral one bifid in the only specimen at hand. A further pair of spiniform tubercles towards the frons and two pairs of low spiniform tubercles in the posterior portion. Cheeks with two small granules (Fig. 303). Eyes rather small, circular in outline and contained almost 2.5x in length of cheeks. Antennae reaching about half way along median segment. Scapus some 1.5x longer than wide with the lateral margins gently rounded. Pedicellus less than half the length of scapus and about equal in length to III, roughly cylindrical. Thorax: Pronotum slightly longer and somewhat narrower than head, 1.6x longer than wide, roughly rectangular and with the lateral margins very gently emarginated medially (Fig. 303). Transverse median sulcus distinct, gently curved and almost expanding over entire width of segment. Anterior portion with two pairs of blunt spines, the anterior pair being larger and more distant than the posterior pair; a further pair of small spiniform tubercles on anterior margin. Posterior half with ten almost equally sized paired spines. Profurcasternum with two black-tipped spiniform tubercles at lateral margins and two tubercles near posterior margin. Mesothorax about 2.2x longer than head and pronotum combined, slender and somewhat constricted in the anterior portion. Mesonotum all over armed with numerous spines of variable sizes and a longitudinal row of about 12 pointed spines along lateral margins. Mesopleurae armed with a longitudinal row of about 16 medium sized spines and metapleurae with a marginal row of about 8–9 short spines. Mesosternum with roughly 12 fairly distinct spiniform tubercles and the metasternum sparsely tuberculose. Tegmina broadly ovate, coriaceous, with the venation very distinct, dense and irregularly disposed; slightly projecting posterior margin of metanotum; the median protuberance shallow. Alae some 1.3x longer than tegmina and almost reaching to posterior margin of median segment.



157

FIGURES 298–300. Parhaplopus evadne (Westwood, 1859) n. comb.. 298. ♀: Haiti, Department du Sud, Formon [USNM]; 299. ♂ Haiti, Department du Sud, Formon [USNM]; 300. ♂ HT: Dominican Republic, Santo Domingo [NHMUK].

Abdomen: Median segment 1.6x longer than wide and roughly rectangular, surface smooth. Tergites I–IV each with posteromedian pair of small retrorse spines, which are most prominent on II; remaining segments unarmed. Segments II–VII roughly equal in length and about 1.3x longer than wide. II–III somewhat widening, IV roughly parallel-sided and V–VII very slightly but gradually narrowing. Tergum VII with the lateral margins very gently rounded (Fig. 308). Sternum II with a pair of granules posteromedially, remaining sterna smooth. Praeopercular organ formed by a shallow, longitudinal median granule in posterior portion of sternum VII (Fig. 309). Tergum VIII somewhat less than ¾ the length of VII and roughly 1.5x longer than wide. IX almost rectangular and about half the length of VIII. Anal segment parallel-sided with posterior portion narrowed and the posterior margin with aconcave median emargination; epiproct small almost semi-circular and shield-shaped with a fine longitudinal median carina (Fig. 308). Cerci very small, conical and scutely pointed; hardly projecting over posterior margin of


HENNEMANN ET AL.

anal segment. Subgenital plate very long, naviculate, longitudinally carinate and gradually narrowing towards an acute apex; extending over abdomen by more than the combined length of tergites VIII–X (Figs. 307–309).

FIGURES 301–309. Parhaplopus evadne (Westwood, 1859) n. comb.. 301. Head and pronotum of ♂ HT (lateral view) [NHMUK], 302. Head and pronotum of ♂ Haiti, Department du Sud, Formon [USNM]; 303. Head and pronotum of ♀ Haiti, Department du Sud, Formon [USNM]; 304. ♂ apex of abdomen: Haiti, Department du Sud, Formon (lateral view) [USNM]; 305. ♂ apex of abdomen: Haiti, Department du Sud, Formon (ldorsal view) [USNM]; 306. ♂ apex of abdomen: Haiti, Department du Sud, Formon (ventral view) [USNM]; 307. ♀ apex of abdomen: Haiti, Department du Sud, Formon (lateral view) [USNM]; 308. ♀ apex of abdomen: Haiti, Department du Sud, Formon (ldorsal view) [USNM]; 309. ♀ apex of abdomen: Haiti, Department du Sud, Formon (ventral view) [USNM];

Legs: Pro- and mesofemora shorter than mesothorax and metafemora almost reaching half way along abdominal segment III. Medioventral carina of profemora with six small spines, in mesofemora armed with four and on metafemora with five rather prominent spines. Anterodorsal carina of profemora and tibiae strongly raised, the latter slightly undulate. Anteroventral carina of meso- and metafemora with two, the posteroventral carina with



159

one sub-apical spine. Both dorsal carinae slighty elevated sub-apically. Anterodorsal carina of mesotibiae gently rounded sub-basally and sub-apically. Probasitarsus about as long as the following three tarsomeres combined, the dorsal carina gently rounded. Meso- and metabasitarsi simple and slightly shorter than following three tarsomeres combined. ♂ (Figs. 299–300, 379–398). Large (body length 89.0–99.2 mm) and fairly slender for the genus with well developed alae (length >36.0–49.0 mm). Colour of body and legs pale yellowish green, the mesonotum, metanotum and three basal abdominal tergites pale creamish brown; body surface slightly glabrous. Cheeks with a faint yellow postocular stripe, eyes pale creamish brown and the antennae drab to pale ochraceous. All spines of thorax with black tips. Metapleurae with a white longitudinal stripe along upper margin and lateral margins of abdominal tergites VIII and IX broadly yellow. Cerci cream to pale brown and greenish basally. Tegmina and costal region of alae bright yellow with five bright green longitudinal streaks along the main veins. Anal region of alae plain transparent orange. Tarsi pale greenish brown. Head: Ovoid and about 1.3x longer than wide, vertex slightly convex and armed with a pair of obtuse tubercles to acutely pointed spines; usually two small granules behind these (Figs. 301–302). Eyes sub-circular, projecting hemispherically and their length contained less than 2x in that of cheeks. Antennae moderately robust and reaching to anterior of abdominal segment III; consisting of about 56 segments. Thorax: Pronotum about equal in length but narrower than head, roughly rectangular and with the lateral margins gently concave; surface smooth (Figs. 301–302). Transverse median depression moderately distinct, slightly rounded and not reaching lateral margins of segment. Mesothorax about 2.3x longer than head and pronotum combined. Mesonotum armed with 9–12 rather distinct, obtusely conical spines which are roughly arranged in two longitudinal rows along anterior 2/3 of dorsal surface; either black-tipped or almost completely black. Mesopleurae with a longitudinal row a minute granules, metapleurae unarmed. Mesosternum with about four low spines in anterior portion and a few small tubercles behind; metasternum smooth. Tegmina oval and reaching about 1/3 along median segment, central protuberance rather distinct, roundly conical and positioned premedially. Alae very long and projecting over posterior margin of abdominal segment V. Abdomen: Segments II–IV roughly of equal length and about 4x longer than wide. V–VII gradually decreasing in length with VII only about ¾ the length of II–IV and no more than 3.5x longer than wide. All tergites and sternites smooth, tergites II–VII parallel-sided. VIII about 3/5 the length of VII and gradually widening towards the posterior; IX very slightly shorter than VIII and narrowed towards the posterior. Anal segment with a faint longitudinal median carina, slightly converging towards the posterior and posterior margin with a roundly triangular median emargination (Fig. 305); on ventral surface armed with several small, black in-curving denticles. Epiproct very small and fully concealed by anal segment. Vomer with a broad and rounded base and a short but acutely pointed terminal hook (Fig. 355). Cerci elongate, a little longer than anal segment, laterally compressed, carinate dorsally and ventrally (Fig. 355) and gradually constricted towards a narrow apex (Figs. 304–306). Poculum moderately convex, scoop-like and with a small, rounded tubercle at the base (Fig. 304); posterior portion longitudinally carinate and the posterior margin indented medially (Fig. 355). Legs: Profemora a little longer and mesofemora slightly shorter than mesothorax, metafemora reaching about 2/3 the way along abdominal segment IV. Medioventral carina of profemora with three small spines in apical portion, of meso- and metafemora armed with five strong spines. Anteroventral carina of meso- and metafemora with two, the posteroventral carina with one sub-apical spine. Basitarsi a little longer than following three tarsomeres combined. Variability: Males show variability in the number and size of the mesothoracal spines as well as the size and shape of the cephalad horns. The mesonotal spines range from 9–12 in number and are mostly black-tipped but may be almost completely black in specimens which have them strongly developed. The cephalad horns are rather blunt and conical in shape (Fig. 301) but in some specimens at hand are prominent, elongate and acutely pointed with the tip if the dextral horn black (Fig. 302). Comments: Westwood (1859: 85, pl. 18: 6) originally described and illustrated this species based on a unique ♂ from Santo Domingo (Dominican Republic) in NHMUK. Eggs unknown. Distribution (Fig. 379): Hispaniola. Dominican Republic: Santo Domingo [NHMUK]; Departamento de Vida Silvestre [photograph taken by Eladio Fernández]. SE-Haiti: Parc National Pic Macaya, Massif de la Hotte, Formon, alrededores de Kay Michel 1100 m [USNM]. Gonaive Island [ANSP]. Number of specimens examined: 8


HENNEMANN ET AL.

TABLE 31. Measurements of Parhaplopus evadne (Westwood, 1859) n. gen., n. comb. [mm] ♂ HT [NHMUK]

♂ [ANSP]

♂♂ [USNM]

♀ [USNM]

-

-

-

149.0

Body

99.2

ca. 96.0

89.0–94.5

135.0

Pronotum

3.8

3.6

3.2–3.3

6.0

Mesonotum

17.3

20.0

15.6–18.2

27.8

Metanotum

6.3

6.2

-

6.7

Median segment

7.9

7.8

-

10.0

Tegmina

12.0

12.0

9.8–10.8

10.5

> 36.0

49.0

46.0–48.5

13.0

Profemora

17.8

18.0

16.2–16.9

19.0

Mesofemora

16.5

17.0

14.8–17.0

18.2

Metafemora

23.4

22.0

19.0–22.1

24.6

Protibiae

16.3

17.0

16.6–17.8

19.4

Mesotibiae

14.0

15.6

12.5–13.5

17.3

Metatibiae

19.3

20.1

16.6–18.5

23.0

Antennae

> 37.0

> 39.0

50.0–52.0

55.0


Alae

Parhaplopus navarroi n. gen., n. sp. (Figs. 310–321, 343, 356, 379) HT, ♂: Dominican Republic, RD-095 Rodeo, ~ 0.5 km E Presa de Blanco, Bonao, Monseñor Nouel Prov., 20.III.2003, D. Perez, R. Bastardo, B. Hierro. (night) [USNM]. PT, ♀: Dominican Republic, Jardín Botánico Nacional, Santo Domingo, 20.IX.2000, 18°29.7’N 69°57.3’W, D. E. Perez, R. H. Bastardo [USNM]. PT, egg (removed from ovipositor): Dominican Republic, Jardín Botánico Nacional, Santo Domingo, 20.IX.2000, 18°29.7’N 69°57.3’W, D. E. Perez, R. H. Bastardo [USNM].

Diagnosis: Similar to the Cuban P. cubensis (Saussure, 1868) but differing by: the smaller size and relatively shorter mesothorax of both sexes; more distinct pair of spines on the vertex (Figs. 304–306); less deflexed abdominal tergum VII (Fig. 308); narrowly rounded apex of the subgenital plate (Fig. 308) and less undulate dorsal carina of the protibiae of ♀♀, as well as the distinct white cheeks (Fig. 306) and more slender legs of ♂♂. Etymology: This pretty new species is dedicated to Santo Q. Navarro (Universidad Autónoma de Santo Domingo) for his invaluable support and assistance throughout the years to the collection efforts of the third author in the Dominican Republic. Description: ♀ (Fig. 310). Of medium size (body length including the subgenital plate 117.5 mm) and fairly stocky for the genus. Colour greyish mid brown but the mesothorax and abdomen strongly darkened during preservation in the unique specimen at hand; abdominal sternites dull green posteriorly. Legs with a slight greenish wash and basal portion of profemora pale greyish green. Dorsal spines of the mesonotum with black tips, all other tubercles and spines of the head and thorax tipped with dull green. Antennae dark reddish brown ventrally and rather greyish mid brown dorsally. Eyes sepia with dull orange mottling. Tegmina and costal region of alae very dark brown and almost black laterally. Anal region of alae transparent grey with all major longitudinal and transverse veins broadly marked with dark grey or black. Head: Ovoid, about 1.3x longer than wide (Fig. 312); vertex gently convex in the posterior portion and armed with a pair of moderately sized, conical cephalad spines as well as a number of smaller tubercles; two small granules are present on the cheeks (Fig. 313). Eyes rather small, circular and contained almost 3x in length of



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cheeks. Antennae broken in the only specimen available. Scapus 2.3x longer than wide with the lateral margins gently rounded. Pedicellus less than half the length of scapus and longer than III, roughly cylindrical.

FIGURES 310–314. Parhaplopus navarroi n. gen., n. sp.. 310. ♀ PT: Dominican Republic, Santo Domingo Province, Santo Domingo [USNM]; 311. ♂ HT: Dominican Republic, Monseñor Nouel Province, Bonao [USNM]; 312. Head and pronotum of ♀ PT (dorsal view) [USNM]; 313. Head and pronotum of ♀ PT (lateral view) [USNM]; 314. Head and pronotum of ♂ HT (lateral view) [USNM].

Thorax: Pronotum slightly shorter and narrower than head, 1.4x longer than wide, roughly rectangular and with the lateral margins gently emarginated medially (Fig. 312). Transverse median sulcus distinct, gently curved and almost reaching lateral margins of segment. Dorsal surface with a slightly enlarged pair of spiniform tubercles just in front of posterior margin and otherwise with several irregularly disposed pointed tubercles (Fig. 312). Probasisternum with a median pair of tubercles, and two granules at each lateral margin. Profurcasternum with two small tubercles at lateral margins and two further tubercles at posterior margin. Mesothorax about 2x longer than


HENNEMANN ET AL.

head and pronotum combined, rather broad and slightly down-curving in lateral aspect, constricted at the anterior and conspicuously swollen medially. Mesonotum gradually widened towards the mid of segment, then very gently rounded and in posterior portion almost parallel-sided. Surface all over armed with numerous prominent, irregularly paired, conical spines of variable sizes; in the median portion with a single conspicuously enlarged pair of spines. Along lateral margins a longitudinal row of about 10–14 pointed spines. Mesopleurae armed with a longitudinal row of about 12 small but acute spines, metapleurae with a marginal row of eight spiniform tubercles. Mesosternum with short, sparse tubercles, the metasternum with three pairs of small granules. Tegmina broadly ovate, coriaceous, with the venation very distinct, dense and irregularly disposed; roughly reaching to posterior margin of metanotum. The median protuberance rather shallow. Alae a little longer than tegmina and reaching about ¾ the way along median segment. Abdomen: Median segment 1.8x longer than wide and roughly rectangular, surface smooth. Tergites II and III each with a few small tubercles, II–IV each with posteromedian pair of retrorse spines, which are most prominent on II; remaining segments unarmed. Segments II–VI roughly equal in length and about 1.8x longer than wide. Tergum VII shorter than previous and with the lateral margins gently rounded posteriorly (Fig. 316). VIII less than ¾ the length of VII and roughly 1.5x longer than wide. IX almost rectangular and about half the length of VIII. Anal segment almost parallel-sided but somewhat narrowed in posterior portion, with a small posteromedian emargination; epiproct very small and triangular (Fig. 316). Cerci very small and conical with a pointed apex; hardly projecting over posterior margin of anal segment (Fig. 315). Subgenital plate very long, naviculate, longitudinally carinate and with a gently expanded, rounded apex; extending over abdomen by more than the combined length of tergites VIII–X (Figs. 315–316). Legs: Pro- and mesofemora shorter than mesothorax and metafemora slightly projecting over posterior margin of abdominal segment III. Medioventral carina of profemora with three small spines, in mesofemora armed with four and on metafemora with five rather prominent spines. Anterodorsal carina of profemora and tibiae strongly raised and slightly undulate. Both outer ventral carinae of meso- and metafemora with two sub-apical spines, although smaller on the posterior carina; otherwise both carinae very minutely tuberculose. Both dorsal carinae slighty elevated sub-apically. Anterodorsal carina of mesotibiae gently rounded sub-basally and sub-apically. Probasitarsus about as long as the following three tarsomeres combined, dorsal carina rounded. Meso- and metabasitarsi simple and slightly shorter than following three tarsomeres combined. ♂ (Fig. 311). Of moderate size (body length 78.0mm) and fairly slender for the genus. Colour of body and front legs greenish mid brown, the metapleurae, metasternum and three terminal abdominal segments bright apple green. Mid and hind legs green with apices of the femora brown. Mesosternum whitish and the ventral surface of the abdomen dull green with white mottling. Cheeks mostly white (Fig. 314), eyes dark reddish brown. Spines of mesothorax with black tips. Tegmina and costal region of alae with anterior margin white, then a broad brown longitudinal band, and innermost portion bright apple green; in alae the brown longitudinal band with several washed greyish markings. Anal region of alae pale transparent pink with a slight greyish hue along the outer margin (Fig. 319) Antennae dark ochracheous and becoming darker towards the apex. Head: Generally as in ♀♀ but only 1.2x longer than wide, with the vertex gently convex and smooth except for a pair of small but acute cephalad spines (Fig. 314). Eyes much more prominent, projecting hemispherically and their length contained only about 1.8x in that of cheeks. Antennae projecting over posterior margin of abdominal segment II; consisting of 59 segments. Otherwise as in ♀♀. Thorax: Pronotum slightly shorter and narrower than head with the lateral margins very gently concave; surface smooth (Fig. 314). Transverse median sulcus shallow, gently rounded and almost reaching lateral margins of segment. Prosternum with a small median granule at each lateral margin. Mesothorax about 2x longer than head and pronotum combined. Mesonotum armed with four pairs of moderately sized but pointed spines in the anterior 2/3. Mesopleurae with a few small granules, metapleurae smooth. Mesosternum with three pairs of small tubercles, metasternum smooth. Tegmina oval and reaching almost half the way along median segment, central protuberance roundly conical. Alae reaching to posterior margin of abdominal segment VI (45.0 mm). Abdomen: Segments II–IV roughly equal in length and almost 4x longer than wide; all unarmed. V slightly shorter than previous and V–VII gradually decreasing in length with VII hardly 3x longer than wide. VIII 3/5 the length of VII and widened towards the posterior, IX about equal in length to VIII and constricted medially. Anal segment a little shorter than IX, widened towards the posterior and posterior margin with a deep triangular median emargination (Fig. 318). Epiproct very small and fully concealed by anal segment. Cerci slightly shorter than anal



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segment, laterally compressed and gradually narrowed towards a slender apex (Fig. 309). Vomer very broad, roundly triangular and with a very short, straight terminal hook (Fig. 356). Poculum moderately convex, scoopshaped and longitudinally carinate in posterior portion; reaching to posterior margin of tergum IX (Figs. 317–318).

FIGURES 315–321. Parhaplopus navarroi n. gen., n. sp.. 315. Apex of abdomen of ♀ PT (lateral view) [USNM]; 316. Apex of abdomen of ♀ PT (dorsal view) [USNM]; 317. Apex of abdomen of ♂ HT (lateral view) [USNM]; 318. Apex of abdomen of ♂ HT (dorsal view) [USNM]; 319. Right tegmen and ala of ♂ HT [USNM]; 320. Egg PT (dorsal view) [USNM]; 321. Egg PT (lateral view) [USNM].

Legs: Pro- and mesofemora almost equal in length and slightly longer than mesothorax, metafemora reaching about half way along abdominal segment IV. All less carinate than in ♀♀ and lacking any dorsal elevations or teeth. Armature of medioventral carina of the femora as in ♀♀ but considerably less developed. Posteroventral carina of meso- and metafemora with only one small sub-apical spine. Tarsi relatively more elongate, probasitarsus almost equal in length to following three tarsomeres combined, meso- and metabasitarsus about as long as combined length of following two segments. Dorsal carina of probasitarsus very gently raised. Egg (Figs. 320–321): A single fully developed egg was removed from the ovipositor of the paratype. Description as for the genus (see above). Measurements [mm]: Length 3.8, length (including operculum) 4.0, width 2.0, heigth 2.1, length of micropylar plate 1.2. Comments: The two type specimens are from separate localities but undoubtedly represent both sexes of the same species. The fact that only two specimens were encountered during as many as nine expeditions and collections at 280 sites throughout the Dominican Republic suggest this species to be rare. The conspicuous colouration of the unique known egg seems to be unusual and is perhaps an artifact or abnormality. Distribution (Fig. 379): Hispaniola (Dominican Republic: Monseñor Nouel Prov. & Santo Domingo) [USNM]. Number of specimens examined: 2


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TABLE 32. Measurements of Parhaplopus navarroi n. gen., n. sp. [mm] HT, ♂ [USNM]

PT, ♀ [USNM)]

-

117.5

Body

78.0

103.5

Pronotum

2.8

4.0

Mesonotum

12.8

18.8

Metanotum

6.1

6.2

Median segment

6.9

8.0

Tegmina

10.3

7.6

Alae

45.0

8.9

Profemora

13.8

15.1

Mesofemora

13.9

14.2

Metafemora

17.3

20.6

Protibiae

12.8

14.4

Mesotibiae

10.7

13.2

Metatibiae

15.5

18.3

Antennae

38.5

>


5.8. Genus Venupherodes n. gen. Type-species: Platycrana venustula Audinet-Serville, 1838: 242, by present designation. Platycrana, Audinet-Serville, 1838: 242. Diapherodes, Saussure, 1872: 185. Redtenbacher, 1908: 434 (in part). Westwood, 1859: 84 (in part). Otte & Brock, 2005: 120 (in part). Diapherodes (?), Kirby, 1904a: 362. Haplopus, Redtenbacher, 1908: 429 (in part). Otte & Brock, 2005: 150 (in part). Phasma, Westwood, 1859: 34, pl. 22: 7.

Description: ♀♀, ♂♂ (Figs. 322–325). Small Haplopodini (body length < 80 mm) with general appearance similar to Diapherodes Gray, 1835; ♂♂ indistinctly shorter than ♀♀. ♂♂ slender with scale-like tegmina and welldeveloped alae. ♀♀ apterous and rather broad insects with the body distinctly oval in cross-section and the thorax and abdomen conspicuously broadened. ♀♀ usually bright green but sometimes with washed blackish-brown portions, ♂♂ a mixture of mid green, dull green, creamish brown and white. Head of ♂♂ with a conspicuous white streak along the cheeks. Head globose, indistinctly longer than wide. Vertex roundly convex and indistinctly bicornute; sinistral tubercle often lacking. Antennae slender and as long as (♀♀), or considerably longer than head and thorax combined (♂♂). Scapus dorsoventrally flattened, longer than pedicellus. Pronotum with a ± prominent pair of blunt spines or spiniform tubercles in anterior half. Mesothorax of ♀♀ strongly broadened towards the posterior and constricted towards the anterior; roughly 1.5x the combined length of head and pronotum. Mesonotum conspicuously lyriform, being strongly broadened towards about 2/3 of its length and gently constricted in the posterior portion, laterally overlapping mesopleurae. Dorsal surface of mesonotum with a distinct, longitudinal median keel and anterior margin with a small pair of spiniform tubercles. Meso- and metapleurae granulose in ♀♀, unarmed in ♂♂. Mesosternum very minutely granulose in ♀♀ and smooth in ♂♂, metasternum smooth in both sexes. Abdomen of ♀♀ hardly longer than head and thorax combined and swollen



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medially; segment II–II increasing and V–X decreasing in width, IV widest segment. Segments II–VI wider than long. All tergites of ♀♀ finely multi-carinate, VII parallel-sided. Abdomen of ♂♂ slender and cylindrical with the terminal three segments just very slightly broader than previous. Praeopercular organ of ♀♀ formed by three small, wart-like tubercles which are arranged in a triangle close to posterior margin of sternum VII. Cerci very small (♀♀ in particular) and laterally compressed; almost straight. Epiproct very small and mostly hidden under anal segment. Subgenital plate of ♀♀ moderately, longitudinally keeled and tapered towards an acute apex; projecting over apex of abdomen by hardly more than the length of the anal segment. Vomer of ♂♂ very broad and roughly triangular, poculum moderately convex and with a blunt central spine. Legs of both sexes moderately slender and rather short in ♀♀, profemora about as long as (♂♂) or shorter than mesothorax (♀♀), hind legs not reaching apex of abdomen. All carinae of legs unarmed, except for spines on the medioventral carina and 1–2 small, sub-apical spines on the two outer ventral carinae of the meso- and metafemora. Basitarsi short, about as long as following two tarsomeres combined. Eggs (Figs. 332–333): Fairly small, capsule about 1.6x longer than wide and with dorsal surface more convex than ventral surface; polar area slightly impressed if seen in lateral aspect. Capsule surface irregularly but very prominently and unevenly sculptured being all over covered with raised ridge-like structures, tubercles and granules; a very blunt tuberculate longitudinal keel runs from the micropylar plate towards the polar-area. Micropylar plate impressed and lower than capsule surface, less than 1/3 the length of capsule, roughly oval, shield-shaped and 1.3x wider than long. Central portion of micropylar plate granulose, micropylar cup distinct and placed in a small posteromedian gap of plate. Operculum with an outer rim of irregularly raised tubercles; centre slightly lowered. General colour mid to dark brown, sometimes with a greyish wash, micropylar plate mid brown. Differentiation: The single Cuban representative of this new genus is easily distinguished from the closely related Diapherodes Gray, 1835 (→ Table 33). ♀♀ at once differ by being completely apterous, having a distinct longitudinal median keel on the mesonotum, a lyriform mesothorax which is distinctly widened laterally and overlaps the mesopleurae, and having a medially swollen and roundly expanded abdomen with IV being the widest segment. ♂♂ exhibit a very characteristic colouration, having the cheeks with a distinct white stripe and eggs are clearly distinguished by the broad and shield-shaped micropylar plate. The absence of wings in the ♀♀ resembles the second exclusively Cuban genus Aploploides Rehn & Hebard, 1938, which may be sympatric. This new genus however differs at first glance by: the globose and bi-cornute head, shorter cerci and armed thorax of both sexes, which is longitudinally keeled in ♀♀, as well as the more bulgy poculum of ♂♂. Distribution (Fig. 374): Cuba, endemic. Etymology: Neuter. The generic name is a combination of “Venu(s)” (venus gr. = goddess of love), overtaken from the specific name “venustula” of the type-species, and the ending “-pherodes” to indicate the close relation to Diapherodes Gray, 1835. Species included: 1. Venupherodes venustula (Audinet-Serville, 1838: 242) [Platycrana]. n. comb. = Phasma havaniense Westwood, 1859: 34, pl. 22: 7, 7a, 7b (♀). = Haplopus juvenis Redtenbacher, 1908: 430, pl. 20: 4 (♂). n. syn.

TABLE 33. Comparison and differentiation of Diapherodes Gray, 1835 and Venupherodes n. gen. Diapherodes

Venupherodes n. gen.


64.0–126.0 mm

54.5–68.0 mm


> 90.0 mm

< 85.0 mm

Unicolorous or with a faint dark green marking on cheeks

Cheeks with a distinct white marking or stripe (Fig. 327)

Broadened towards the posterior; mesopleurae dilated and projecting laterally over mesonotum

Distinctly lyriform; mesonotum laterally overlapping mesopleurae (Fig. 322)

Head (♂♂) Mesonotum—shape (♀♀)

.....continued on the next page


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TABLE 33. (Continued) Diapherodes

Venupherodes n. gen.

Smooth or with an indistinct longitudinal median carina; with 2–6 ± distinct anterior spines

With a blunt, longitudinal median keel; at best with a small anterior pair of tubercles (Fig. 322)

With a ± prominent marginal row of spines, teeth or acute tubercles

Granulose

Brachypterous (alae may be vestigial)

Apterous (Figs. 322–324)

Broadened basally or sub-basally; II or III broadest segment; considerably longer than head and thorax combined

Strongly widened medially; IV broadest segment; hardly longer than head and thorax combined (Fig. 322)

Lateral margins ± expanded laterally

Gradually narrowing with lateral margins straight (Fig. 329)

Praeopercular organ (♀♀)

A single, oval or longitudinal wart-like tubercle (Figs. 292–294)

A cluster of three wart-like tubercles, arranged in a triangle (Fig. 345)

Subgenital plate (♀♀)

Long and ± lanceolate; projecting over apex of abdomen by at least the combined length of tergites IX and X

Acutely spatulate; projecting over apex of abdomen by hardly more than length of anal segment (Figs. 328–329)

> 3.7 mm

< 3.5 mm

Elongate to oval, often longer than wide and at least 1/3 as long as capsule

Shield-shaped, wider than long and < 1/3 as long as capsule (Figs. 332–333)

Lesser Antilles & Greater Antilles (Puerto Rico & Jamaica) Fig. 375

Cuba (Fig. 374)

Mesonotum—surface (♀♀)

Meso- and metapleurae (♀♀) Wings (♀♀) Abdomen (♀♀)

Abdominal tergum VII (♀♀)

Egg (capsule length) Egg (micropylar plate) Distribution

Venupherodes venustula (Audinet-Serville, 1838) n. gen., n. comb. (Figs. 322–333, 345, 364, 374, 384–387) Platycrana venustula Audinet-Serville, 1838: 242. HT, ♀: Cuba [MNHN—lost]. NT (by present designation), ♀: 800; Havaniense (M. Leay) Westw., Cuba, Otto [MNHU]. Diapherodes venustula, Westwood, 1859: 84. Saussure, 1872: 185. Bolivar, 1888: 140. Redtenbacher, 1908: 435. Rehn, 1909: 200. Moxey, 1972: 117 (in litt.). Otte & Brock, 2005: 121. Rabaey & Simoens, 2007: 6, figs. [Culture report] Diapherodes (?) venustula, Kirby, 1904a: 362. Phasma (Diapherodes) venustulum, de Haan, 1842: 109. Phasma havaniense Westwood, 1859: 34, pl. 22: 7 (♀). HT, ♀: Ins. Havannah; Type—Westwood, Phasma havaniense, Cat. Phasm. 1859, p. 24, pl. 22 f. 7; Pleseophyllum Havaniense; Type Orth: 615, Phasma havaniense Westw., Hope Dept. Oxford [OXUM, No. 615]. [Synonymised by Saussure 1871–1872: 185] Moxey, 1972: 117 (in litt.). Otte & Brock, 2005: 121. Haplopus juvenis Redtenbacher, 1908: 430, pl. 20: 4 (♂). LT (by present designation), ♂: Coll. Br. v. W., ex. Coll. Sommer, Cuba; det. Br. v. W., Haplopus juvenis (specimen with open wings) [NHMW, No. 795]; PLT, ♂: Coll. Br. v. W., ex. Coll. Sommer, Cuba; det. Br. v. W., Haplopus juvenis [NHMW, No. 795]; PLT, ♂: Coll. Br. v. W., ex. Coll. Sommer, Cuba; det. Br. v. W., Haplopus juvenis, 77.; 7647, jamaicensis, Cuba [NHMW, No. 795]; PLT, ♂: Coll. Br. v. W., ex. Coll. Sommer, Cuba; det. Br. v. W., Haplopus juvenis; 7647; 7647 Phasma jamaicensis Fabr. Drur., Spectrum viride Stoll, capta T. Müller, Cuba, Phas. Inv. Nr. 795 [NHMW, No. 795]. n. syn. Rehn, 1909: 200. Moxey, 1972: 117 (in litt.). [Listed as a synonym of Diapherodes venustula (Audinet-Serville, 1838)—unpublished]



167

Otte & Brock, 2005: 151. Aplopus juvenis, Brock, 1998a: 57. Further material [78 ♂♂, 53 ♀♀, 3 nymphs, eggs]: CUBA: 1 ♀: Coll. Br. v. W., Kuba, Saussure; det. Br. v. W. Diapherodes venustula; 11.165; 11.165 Diapherodes venustula Serv. (Havaniensis) ♀ [NHMW, No. 840]; 1 ♀: Diapherodes venustula, Serv.; Cuba (Lembeye) [MNCN]; 1 ♀: Diapherodes venustula, Cuba, Gundlach [MNCN]; 1 ♂ (penultimate instar): Cuba, Cabrara [MNCN]; 1 ♀, 2 ♂♂: Cuba, achété par M. Poey; Diapherodes venustula Serv. (MHNG); 2 ♀♀, 2 ♂♂: Cuba, Mr. H. de Saussure; Diapherodes venustula Serv. [MHNG]; 1 ♂: Cuba, coll. Guerin; Diapherodes venustula Serv. [MHNG]; 1 ♂, 1 ♀ (nymph): Cuba; Diapherodes venustula Serv. [MHNG]; 2 ♂♂: 836; venustula (Westw.), Platycr. venustula Serv., Cuba, Otto [MNHU]; 1 ♀, 1 ♂: ex Zucht: K. Rabaey (Belgien), Herkunft: W-Cuba, leg. I. Fritzsche, VII.2006 [coll. FH, No’s 0599- 1 & 3]; 1 ♂: ex Zucht: F. Hennemann, urspr.: CUBA, leg. I. Fritzsche, VII.2006 [coll. FH, No. 0599-2]; 13 ♀♀, 18 ♂♂, 1 ♂ (penultimate instar), eggs: ex Zucht: F. Hennemann 2007/2008, Herkunft: W-Cuba, Pinar del Rio Provinz, leg. I. Fritzsche 2005 [coll. FH, No’s 0599-4 to 35 & E]; 1 ♀, 2 ♂♂: ex Zucht F. Hennemann 2008/09, Herkunft: W-Cuba, Pinar del Rio Provinz, leg. I. Fritzsche 2005 [coll. FH, No’s 0599-36 to 38]; 1 ♂: Sierra Rangel, de Rio—1934; Roberts Coll.; wO172 [ANSP]; 1 ♂, 1 ♀: Somorrostro, Havana, Aug. 20, 35, Schramuzza, y. Brunner [ANSP]; 1 ♂, 1 ♀: Marianao, Cuba, Aug. (From. T. Barbouy) [ANSP]; 30 ♀♀, 46 ♂♂, eggs: ex Zucht O. Conle, Herkunft: W-Cuba, Pinar del Rio Provinz [coll. OC].

Description: ♀ (Figs. 322–324, 384, 386–387). Small (body length including subgenital plate 65.0–80.0 mm) and robust Haplopodini (maximum body width at abdominal segment IV 8.0–10.0 mm) with a strongly broadened mesothorax and medially swollen abdomen. Colour of body and head bright apple green with the ventral body surface dull green. Sometimes specimens occur which have certain parts of the dorsal body surface furnished with irregular washed brownish or blackish markings (Figs. 324, 387); more rarely blackish markings are seen on the ventral surface of the abdomen. Rather frequently specimens have the head dull brownish green or almost entirely brown. Anterior margin of pronotum yellow. Anterior portion of the lateral margins of mesonotum ochre, mesoand metapleurae ochre with a broad, white longitudinal stripe along ventral margin. Coxae reddish brown. Cerci pale to dark brown. Front legs greenish brown with the basal portion of the femora pale green. Mid and hind legs pale to mid green with the base of the femora slightly brownish. Basitarsi green, remaining tarsomeres brown. Scapus and pedicellus mid to dark brown. Remainder of antennae pale brown and each antennomere with a fine transverse dark band at the apex; this turning to black towards apex of antennae. Eyes reddish brown with pale cream markings. Head: Globose, indistinctly longer than wide, and considerably broader than prothorax. Vertex convex, with a few small, scattered granules and in centre mostly armed with a blunt, conical dextral tubercle; the sinistral one faint or obsolete (Fig. 326). Eyes convex, almost circular and of moderate size; their length contained about 1.8x in that of cheeks. Antennae slender and hardly reaching posterior margin of median segment; with about 45 antennomeres. Scapus dorsoventrally compressed, about 2x longer than wide and constricted at the base. Pedicellus cylindrical and about half as long as scapus. Thorax: Pronotum about as long but distinctly narrower than head and strongly tapered towards the posterior; anterior margin about 1.3x wider than posterior margin and median portion very gently constricted. Transverse median sulcus very distinct, slightly curved but just not reaching lateral margins of segment. Anterior half with a pair of prominent, blunt spines and close to posterior margin with two small tubercles; otherwise smooth (Fig. 326). Mesothorax about 1.5x longer than head and pronotum combined and strongly constricted anteriorly; posterior margin about 3x wider than anterior margin. Mesonotum roughly 2x longer than wide and distinctly lyriform, being strongly broadened towards about 2/3 of its length and gently constricted in the posterior portion. Lateral margins strongly rounded in medial portion and overlapping the mesopleurae. Entire dorsal surface of mesonotum with a distinct longitudinal median keel, close to the anterior margin with a small pair of ± distinct tubercles (Fig. 326) and with a few rounded granules along the lateral margins; otherwise smooth. Lateral margins very minutely denticulate. Mesopleurae with a fine longitudinal keel close to anterior margin and with a median row of granules. Metapleurae slightly tectiform with an acute longitudinal median keel, unarmed. Mesosternum smooth or with a very few scattered granules, metasternum smooth. Metanotum less than 1/3 the length of mesonotum and roughly quadrate; dorsal surface with a faint longitudinal median keel.


HENNEMANN ET AL.

FIGURES 322–325. Venupherodes venustula (Audinet-Serville, 1839) n. gen., n. comb.. 322. ♀: captive reared from NWCuba, Pinar del Rio [coll. FH, No. 0599-13]; 323. ♀: captive reared from NW-Cuba, Pinar del Rio [coll. FH, No. 0599-3]; 324. ♀: captive reared from NW-Cuba, Pinar del Rio [coll. FH, No. 0599-20]; 325. ♂: captive reared from NW-Cuba, Pinar del Rio [coll. FH, No. 0599-1].



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FIGURES 326–333. Venupherodes venustula (Audinet-Serville, 1839) n. gen., n. comb.. 326. Head and pronotum of ♀: captive reared from NW-Cuba, Pinar del Rio [coll. FH, No. 0599-6]; 327. Head and pronotum of ♂: captive reared from NWCuba, Pinar del Rio [coll. FH, No. 0599-2]; 328. Apex of ♀ abdomen: captive reared from NW-Cuba, Pinar del Rio (lateral view); 329. Apex of ♀ abdomen: captive reared from NW-Cuba, Pinar del Rio (dorsal view); 330. Apex of ♂ abdomen: captive reared from NW-Cuba, Pinar del Rio (lateral view); 331. Apex of ♂ abdomen: captive reared from NW-Cuba, Pinar del Rio (dorsal view); 332. Egg: captive reared from NW-Cuba, Pinar del Rio (dorsal view) [coll. FH, No. 0599-E]; 333. Egg: captive reared from NW-Cuba, Pinar del Rio (lateral view) [coll. FH, No. 0599-E].

Abdomen: Median segment slightly longer than metanotum, a little longer than wide and very gently widening towards the posterior. Dorsal surface with a faint longitudinal median keel, otherwise smooth. Abdomen swollen medially with II–III widening, V–X narrowing and IV broadest segment (Fig. 322). Tergites II–IV transverse with IV almost 2x wider than long, VII slightly longer than wide. Tergites II–IX each with four distinct, longitudinal and roughly parallel carinae; two close to the median line and one on each side about half way towards the lateral margin. The two median carinae are melted together to form a single blunt carina on VIII–X. Sternites II–VII smooth. Praeopercular organ formed by three small, wart-like granules close to posterior margin of sternum VII; these roughly arranged as a triangle (Fig. 345). Tergum VIII about ¾ the length of VII and strongly convex. IX slightly wider than long and about 2/3 the length of VIII. Anal segment about as long as IX, flattened and gradually tapered towards the posterior; posterior margin rounded and with a small median notch and lateral margins somewhat emarginated medially (Figs. 328–329). Epiproct very small, slightly projecting over anal segment, semicircular and with a keel dorsally. Cerci mostly hidden underneath anal segment; finely bristled. Subgenital plate


HENNEMANN ET AL.

longitudinally keeled and tapered towards an acute apex; projecting over apex of abdomen by about 1.5x the length of the anal segment (Figs. 328–329). Legs: All fairly short, profemora and mesofemora shorter than mesothorax and metafemora slightly projecting over posterior margin of abdominal segment IV. Two outer ventral carinae of meso- and metafemora each with 1–2 small but pointed sub-apical spines. Medioventral carina of profemora with 1–2 small sub-apical spines, of mesoand metafemora with five moderately strong spines, which increase in size towards the apex of femur. Dorsal carina of the probasitarsus very gently rounded, all basitarsi roughly as long as following two tarsomeres combined. ♂ (Figs. 325, 385–386). Fairly small (body length 54.5–68.0 mm) and moderately slender insects with well developed alae (29.5–35.0 mm) and a very characteristic colouration. General colour of head and dorsal body surface mid to dull green. Ventral body surface creamish mid brown, sometimes with a slight greenish wash. Head with a bold white longitudinal stripe along the cheeks (Fig. 327). Pronotum ochre with the central portion green. Mesonotum mid to dull green with the anterior and lateral margins broadly ochre to mid brown. Greatest parts of metapleurae bright apple green. Tegmina and costal region of alae mid to dull green with the anterior margin brown and towards the interior bordered by a longitudinal whitish stripe. Central hump of tegmina sometimes dull yellow. Anal region translucent orange. Eyes reddish brown with pale cream markings. Antennae generally as in ♀♀. Fore legs brownish green, mid and hind legs mid to dull green with the bases of the femora pale green. Tarsi greenish brown with the basitarsus green. In addition to the typical colour-form here described, considerably darker specimens frequently occur amongst captive reared specimens. These have the mid and hind legs, tegmina and costal region of the alae very dark green and the mesothorax almost entirely brown, even the metapleurae are dull instead of apple green. It is not known whether such varieties also occur in the wild. Head: Generally as in ♀♀ but vertex slightly less convex and the two cephalad tubercles more distinct (Fig. 327). A small C-shaped impression between the bases of the antennae. Eyes of moderate size, circular and strongly convex; their length contained less than 1.5x in that of cheeks. Antennae moderately thickened and ± reaching to posterior margin of abdominal segment III. Otherwise as in ♀♀. Thorax: Pronotum slightly shorter and narrower than the head, slightly trapezoidal and gradually narrowed towards the posterior; posterior margin rounded. Median transverse depression very distinct, gently curved and almost reaching lateral margin of segment; armature as in ♀♀ but posterior pair of tubercles indistinct to obsolete (Fig. 327). Mesothorax elongate, roughly 2x longer than head and pronotum combined, cylindrical and slightly widened at the posterior. Anterior margin of mesonotum usually with one or two pairs of blunt spine-like tubercles or spines (Fig. 327); sometimes lacking. Surface otherwise with a few irregular, minute granules. Mesopleurae with a few small granules arranged in a longitudinal row, metapleurae as well as meso- and metasternum smooth. Tegmina oval, reaching about 1/3 the way along median segment and with a shallow central hump. Alae reaching to abdominal tergum VI. Abdomen: Median segment considerably longer than metanotum, gently narrowed towards the posterior; smooth. Segments II–VII, elongate, cylindrical, parallel-sided and gradually decreasing in length; II almost 4x and VII just 2.6x longer than wide. Tergites II–VI and sternites II–VII smooth; tergites VI–X with a fine longitudinal median carina. VIII–X indistinctly broader than previous. VIII ¾ the length of VII and very slightly widened towards the posterior. IX a little shorter than VIII, slightly longer than wide and very gently constricted medially. Anal segment 3/5 the length of IX slightly widened towards the posterior and the posterior margin rounded with a broad triangular median indentation (Fig. 331). Epiproct very small and hidden underneath anal segment. Vomer large, flattened and broadly triangular with a very short terminal hook (Fig. 364). Cerci about 2/3 the length of anal segment, slender, strongly laterally compressed and very gently in-curving. Poculum moderately convex, cup-like, acutely carinate longitudinally and with the posterior margin notched medially; reaching about ¾ along tergum IX (Figs. 330, 364). Legs: Generally as in ♀♀ but relatively longer and more slender. Profemora about equal in length to mesothorax, mesofemora shorter than mesothorax, metafemora reaching about 2/3 the way along abdominal segment IV. Nymphs: Newly hatched nymphs are slender and have a body length of ± 16.0 mm. The body, fore legs and antennae are greenish brown, the head, mid and hind legs bright green and the tarsi brown. Later instars vary from dull green over various shades of brown, often irregularly furnished with pale or dark mottling. The legs are pale green to straw. It is interesting to note that nymphs generally resemble those of various Diapherodes and Haplopus-



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species, exhibiting rounded lateral lobes on abdominal tergum VII which disappear in the adult stage. Furthermore, the typical body shape of ♀♀ does not occur before the final ecdysis, even penultimate instar nymphs having a rather slender body. Variability: In adults insects considerable variability is limited to the size and colouration. Slight variability is also seen in the length/width-relation of the mesothorax of ♀♀ as well as the size of the spines on the vertex, pronotum and anterior portion of the mesonotum (the latter ones may be missing). In addition to the usual plain green specimens ones may occur that are washed with dark brown, or have certain body-parts brownish (Fig. 316). Egg (Figs. 332–333): Description as for the genus (see above). Measurements [mm]: Length 3.1–3.5, length (including operculum) 3.7–3.9, width 1.9–2.2, height 2.2–2.6, length of micropylar plate 1.1–1.2. Comments: Audinet-Serville (1838: 242) described Platycrana venustula based on a single ♀ which he stated to be deposited in the “Collection de la Société entomologique de France”. This is now held in MNHN but extensive search has not revealed a single specimen of this species in the entire collection. Therefore, the holotype must be regarded lost (Otte & Brock, 2005: 121) and a ♀ in MNHU is here selected as the neotype to guarantee stability of Audinet-Serville's taxon. Phasma havaniense Westwood, 1859 was described based on a ♀ from Havana in OXUM and was correctly synonymised with P. venustula by Saussure (1872: 185). The corresponding ♂♂ were described by Redtenbacher (1908: 429) as Haplopus juvenis which consequently becomes a synonym of V. venustula (n. syn.). Culture-stock of V. venustula originating in the Pinar del Rio Province was imported to Germany by I. Fritzsche (Wernigerode, Germany) in 2005. This has proven fairly easy to rear in captivity and was included on the Phasmid Study Group culture-list as culture No. 283. A warm, rather dry and well ventilated climate is preferred in captivity. Eucalyptus (Eucalyptus spp., Myrtaceae), oaks (Quercus robur, Q. rubra and Q. ilex, Fagaceae), salal (Gaultheria shallon, Ericaceae) and St. John's Wort (Hypericum patulum, Hypericaceae) are accepted as alternative food plants. Eggs have a rather short incubation period of only 10–12 weeks at an average temperature of 25°C. About 3–4 eggs are laid per day and ♀, each of which produces a total of 150–200 eggs. Mating takes place exclusively at night and lasts for one to three hours. Distribution (Fig. 374): NW-Cuba (Havanna [OXUM]; Mariano [ANSP]; Somorrostro [ANSP]; Cabrera [MNCN];Sierra Rangel, Pinar del Rio [ANSP]; Pinar del Rio [coll. FH, coll. OC]). Number of specimens examined: 140 TABLE 34. Measurements of Venupherodes venustula (Audinet-Serville, 1838) n. gen., n. comb. [mm] ♀, NT [MNHU]

♂, LT of juvenis [NHMW]

♂, PLT of juvenis [NHMW]

♀♀ [coll. FH]

♂♂ [coll. FH]


74.5

-

-

65.0–80.0

-

Body

70.0

65.5

59.2

61.0–76.0

54.5–68.0

Pronotum

3.8

2.7

2.5

3.8–4.0

2.4–3.1

Mesonotum

16.0

12.9

11.4

12.9–15.5

10.0–13.2

Metanotum

5.5

3.8

-

4.8–5.5

3.2–4.0

Median segment

4.9

5.8

-

3.9–5.3

5.0–5.8

Tegmina

-

7.2

6.0

-

5.0–7.7

Alae

-

35.0

30.9

-

29.5–35.0

Profemora

12.3

13.4

12.0

11.7–13.1

11.1–13.9

Mesofemora

10.8

11.9

10.2

9.0–11.8

9.4–12.0

Metafemora

14.0

15.1

13.8

12.3–15.7

12.6–15.0

Protibiae

11.8

13.5

11.7

10.8–12.9

10.5–13.8

Mesotibiae

8.7

9.7

8.6

8.8–10.8

7.8–10.0

Metatibiae

13.0

13.0

11.9

12.0–14.4

11.5–13.0

Antennae

35.0

> 32.0

-

28.0–34.0

33.0–38.5


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5.9. Species incertus sedis One species, formerly attributed to Haplopus Gray, 1835 (Westwood, 1859: 59) and retained in the genus by all subsequent authors is here removed. Unfortunately, the original description of Haplopus dubius (Gray, 1835) is very brief and does not allow any confirmed generic affiliation “Foemina, cylindrica, nigrescens, pulvere albo superinduta; pro mesothoraque spinis minoribus” (Gray, 1835: 34) and does not even include citation of the body length. Gray (1835: 34) however stated the ♀ holotype to be from “Cayenne” (French Guiana), which makes it most unlikely to belong in Haplopodini at all. Westwood (1859: 59) was unable to locate the specimen and also recent searches in the LSUK have been unsuccessful, hence it must be presumed lost. The type-locality and cylindrical body suggest it to belong in Diapheromeridae: Diapheromerinae: Diapheromerine “Phanoclesgroup”and the minutely spinose thorax would affiliate it with Phanocloidea muricata (Burmeister, 1838), a species commonly found in French Guiana. Although, based on the very insufficient original description and lack of measurements any decision on the generic placement or the identity of Gray's species remain impossible, it is here provisionally transferred to Phanocloidea Zompro, 2001 in order not to leave this species without a generic affiliation (n. comb.).

6. BIOGEOGRAPHY The West Indies are a group of differently sized tropical islands in the Caribbean Sea east of Central America, which form a fringing archipelago between Central and South America. They are highly complex historically and geographically and several aspects of their geobiotical history and the distributional mechanisms are still poorly understood and discussed controversially (see e.g. Hedges, 2001; Rickfels & Bermingham, 2007). The West Indies are sufficiently isolated to develop and sustain endemic forms but on the other hand are close enough to sources of colonists to develop a dynamic interaction with surrounding continental regions, this is northern South America in the south and Central America in the west. The depauperate higher taxonomic level fauna but enormous adaptive radiations of certain taxa in the West Indies suggest there has been a kind of selective filter allowing only parts of the biota to colonize these islands, a fact also true for the phasmatodean fauna of the West Indies. For biogeography, it is most important to know which areas were above sea level during the geological history of the West Indies, but unfortunately that is one of the most poorly known aspects of Caribbean geological history (Hedges, 2001: 18). Another fundamental issue in West Indian biogeography concerns the origins of the fauna of the Greater Antilles in particular (Rickfels & Bermingham, 2007: 2395). The Lesser Antilles are accepted to have been colonized entirely by over-water dispersal from northern South America and the Greater Antilles, which is also supported by the phasmatodean fauna of these islands (see below). For the Greater Antilles however, two possible models of colonization have been proposed, this is (i) the over-water dispersal also termed the dispersal model, and (ii) the overland dispersal with subsequent vicariance and separation of biotas resulting from continental drift, also termed the vicariance model. The latter model required land bridges in the distant past, which are now submerged, but there seems to be no supporting evidence for any such land bridges. The resolution of any of these two mechanisms depends mostly on the timing of colonization (Rickfels & Bermingahm, 2007: 2396). If colonization took place close to the beginning of the Tertiary, a land route to the proto-Greater Antilles might have been possible through what is now southern Mexico, a scenario that might explain the present-day distribution of one of the phasmatodean tribes discussed herein i.e. Hesperophasmatini which in addition to numerous representatives mainly on the Greater Antilles also has two genera in southeastern Mexico and Central America (Hennemann & Conle, 2012). Any later colonization via Central America would have meant crossing of a widening water gap. A more recent land connection however, the Greater Antilles-Aves Ridge (Gaarlandia) dating to ca. 33–35 Ma between northern South America and the Greater Antilles was postulated (Iturralde-Vinent & MacPhee, 1999). A primary alternative to overland dispersal for non-flying animals, and generally believed to be the more plausible scenario (e.g. Hedges, 2001), is to raft on flotsam discharged by the outflow of continental rivers and it appears to be evident, that before the Mid-Oligocene some 25 Ma, the Amazon Basin drained to the north into the Caribbean Sea in what is now western Venezuela (Rickfels & Bermingham, 2007: 2396). This hypothesis is congruent to the almost unidirectional northwestward current flow in the West Indies. In general most lineages of biota are estimated to have arrived in the West Indies during the Cenozoic at times when there were no land connections to the surrounding continental areas (Hedges, 2001), but a few groups appear to have arrived very REVISION OF HAPLOPODINI


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early and may represent ancient relicts of the proto-Antillean fauna. However, any broader discussion on the timing of the phasmatodean colonization of the West Indies will require extensive molecular data and phylogenetic studies, which potentially could fully clarify the still speculative relationships of most of the Antillean Phasmatodea. Furthermore, only a fraction of the phasmatodean diversity of the West Indies is known so far and molecular or phylogenetic studies will have to await a more comprehensive knowledge and treatment of the true existing diversity, which is much larger than estimated previously. Phylogenetic analyses (e.g. Whiting et al., 2003; Buckley et al., 2009; Buckley et al., 2010) have provided support for a sister-group relationship between the predominantly Antillean taxa discussed herein (tribes Haplopodini and Hesperophasmatini) and the tribe Stephanacridini Günther, 1953, an Old World clade that is today mostly restricted to some of the Pacific Islands, New Guinea and parts of Wallacea. This suggests a Gondwanan origin for the clade currently recognized as the subfamily Cladomorphinae and furthermore suggests invasion of its ancestors via Gondwanan land connections from the Australasian region through Antarctica to South America perhaps during the late Cretaceous or late Tertiary. This is, for instance, supported by the phylogenetical relationships of certain New World marsupials, which have been shown to have their closest relatives in Australia (Karanth, 2001; Nilsson et al. 2010). Furthermore, up to date five insect species known from Dominican amber have proven to be of Australasian origin, having their closest relatives nowadays distributed throughout Australia, New Guinea, Wallacea and also the Oriental Region (Woodruff, 2009). Invasion of potential ancestors of the phasmatodean clades discussed herein is also possible from Africa during the early Oligocene, e.g. from WestAfrica to northeastern Brazil, but this hypothesis is most speculative since no potentially closely related African taxa (e.g. members of the subfamily Palophinae Kirby, 1896) have so far been included in phylogenetic studies for clarifying possible relationships with Neotropical phasmatodeans. This aforementioned hypothesis should however be taken into account in future phylogenetic or molecular studies, since morphologically there are striking parallels between the Neotropical Cladomorphinae and African Palophinae, which include several characters generally regarded to be of importance to discuss phylogenetic relationships within Phasmatodea. Several Neotropical rodents, primates, bats and turtles are known to be most closely related to African taxa (e.g. Flynn & Wyss, 1998), hence there is a possibility this might also prove true for certain phasmatodean clades. On the basis of geology and size, the West Indies can be divided into three groups of islands: (i) the Greater Antilles (Cuba, Jamaica, Hispaniola, Puerto Rico and Cayman Islands), which are primarily old fragments of continental crust originally formed as islands to the west of the present-day Central American isthmus and carried to their present position by plate tectonics, (ii) the Lesser Antilles (the principal ones are Grenada, Saint Vincent, Barbados, Saint Lucia, Martinique, Dominica, Guadeloupe, Montserrat, Antigua, Saint Kitts (= St. Christopher), Saint Bartholomé, Saint Thomas, Saint John, Tortola and Saint Croix), which form a volcanic arc along the the eastern edge of the eastward moving Caribbean plate, and (iii) the Bahamas Islands, which are a group of 30 larger and about 3000 smaller, low-lying and rather dry islands on the Bahamas platform north of Cuba and Hispaniola (Rickfels & Berminham, 2007). Trinidad, a continental island lying just north of Venezuela is often included with the Lesser Antilles, but its fauna is essentially continental and shows almost no relation to the West Indies, hence biogeographically belongs to the Northwestern Southamerican dominion and together with Tobago is termed the Trinidad-Tobago province (Morrone, 2006: 479). The Caribbean subregion is part of the Neotropical region and is defined to extend through southern Mexico, Central America, the Antilles and northwestern South America (Morrone, 2007: 478). The true West Indies possess a very typical fauna with an apparently high degree of endemism and are regarded as the Antillean dominion of the Caribbean subregion (Morrone, 2006: 479). According to Morrone (2006: 479) the Antillean dominion comprises eight provinces, most of which contain a single island with its satellite islands in the case of the Greater Antilles, or archipelagos as in the Lesser Antilles and Bahamas, and also contains the eastern coastal portions of the Yucatan peninsula below an altitude of 200 metres. These eight biogeographical provinces are defined as follows: (i) the Yucatan Peninsula province (the states of Campeche, Yucatan and Quintana Roo) at altitudes below 200 metres, (ii) the Bahama province, which comprises the Bahamas archipelago as well as the Turks and Caicos Islands, (iii) the Cuba province, which corresponds to the island of Cuba and the nearby Isla de la Juventud, (iv) the Cayman Islands province, which comprises the archipelago of the Grand Cayman, Little Cayman and Cayman Brac islands, (v) the Jamaica province, (vi) the Hispaniola province (Dominican Republic and Haiti), (vii) the Puerto Rico province, and (viii) the Lesser Antilles province, which comprises the archipelagos of the Lesser Antilles in the south and Virgin Islands in the north (see Morrone, 2007: 49).


HENNEMANN ET AL.

The climate of the West Indies is primarily tropical, although the Bahamas and northern parts of the Greater Antilles are affected by northern weather systems during the winter. The higher elevations of mountain ranges in the Greater Antilles posess cooler climates that support in part relict temperate faunas and floras (Rickfels & Bermingahm, 2007: 2395). Precipitation is moderate and fairly constant over the year throughout much of the West Indies, but heavy at mid elevations on the windward sides of higher elevation islands and considerably more sparse on the leeward sides. The Greater Antilles are old islands, believed to be of oceanic origin (Draper et al., 1994) and composed of volcanic rock and marine limestone. Throughout their evolution as islands, the Greater Antilles appear to have fragmented and rejoined in various combinations, which results in that the present-day islands have highly complex geological histories. Although these are broadly understood nowadays, geologists still remain uncertain about many details. Cladistic biogeographic analyses of various biota have shown close relationships between Cuba, Hispaniola and Puerto Rico (Crother & Guyer, 1996) but Cuba and northwestern Hispaniola in particular (Iturralde-Vinent & MacPhee, 1999). All of the Greater Antilles are mountainous, the highest elevation being the Pico Duarte in Hispaniola with an altitude of 3098 m (Orvis, 2003). The largest of these islands is Cuba with an area of 110.860 km². Great parts consist of lowlands and low hilly regions with a central mountainous region. In the very west of the island the Sierra de los Organos reaches 772 m, the central Trinidad highlands reach 1156 m and the Sierra Maestra in the extreme southwest of Cuba rises to a maximum elevation of 1974 m (Pico Turquino). Most of the mountainous regions are covered in rain forest, making some 17% of the island area. With an area of 76.480 km² Hispaniola is just slightly smaller than Cuba but is relatively more mountainous. The highest elevation in the eastern half of the island (Dominican Republic) is the Pico Duarte (3098 m, see Orvis, 2003), while the highest elevation in the western half (Haiti) is only 2680 m. The Dominican Republic is rather more humid than Haiti with some 40% of the area covered in tropical forest, while in Haiti forests are naturally only represented in the central and southern mountain ranges. The entire lowlands of Hispaniola are dominated by dry savannas or semi-arid scrub-land and great parts of altitudes above 850 m are dominated by pine forests. Jamaica is considerably smaller than the former two islands, only 10.991 km², and has a tropical and humid climate. Some 20% are covered with dense tropical rain forest that is mainly restricted to the Blue Mountains in the eastern portion of the island, which reach a maximum altitude of 2257 m. The island represents a fragment of the Central American crust, which was later dragged into the Caribbean basin by plate movement (Rickfels & Bermingahm, 2007: 2394). Puerto Rico is the smallest island of the Greater Antilles with an area of only 8.897 km² if the small satellite islands are included. It is also less mountainous than the other three islands, the highest elevation of the central Sierra Luquillo (Cerro de Punta) reaching only 1338 m. The northern part of the island including the central mountains is very humid mostly covered in tropical rain forest and with plenty of rain, while the southern lowlands are rather dryer and dominated by scrub-land. The Cayman Islands south of Cuba are satellites of the Sierra Maestra in the extreme southwest of Cuba and are regarded as a separate biogeographical province that comprises the three islands Grand Cayman, Little Cayman and Cayman Brac (Morrone, 2001; Morrone, 2006). The low-lying Bahamas Islands (highest elevation on Little San Salvador 123 m), including the Caicos and Turks Islands, are all situated on the Bahamas platform, which appears to have held its geographical position relative to North America throughout the Tertiary being a stable geological feature, that is quite different from the rest of the West Indies (e.g. Meyerhoff & Hatten, 1974). The fauna is comparatively poor, less diverse and exhibits little endemism if compared to the Greater Antilles and is recently derived from North America and the Greater Antilles (Rickfels & Bermingham, 2007: 2395). The islands of the Lesser Antilles are geologically younger, of volcanic origin and considerably smaller in size than the Greater Antilles, the largest being Guadeloupe with an area of 1509 km². They comprise the archipelagos of the Lesser Antilles in the south (Grenada, Saint Vincent, Barbados, Saint Lucia, Martinique, Dominica, Guadeloupe, Montserrat, Antigua, Saint Kitts (= St. Christopher) and Saint Bartholomé) and the Virgin Islands in the north (the principal ones being Saint Thomas, Saint John, Tortola and Saint Croix) and are considered as the Lesser Antilles province (Morrone, 2006: 479). The Virgin Islands are situated no more than 80 km east of Puerto Rico and are in the east separated from the Lesser Antilles archipelago by the Anegada Passage. Although they are generally considered as belonging to the Lesser Antilles their phasmatodean fauna shows influences from both the Lesser and Greater Antilles; for instance a principally Greater Antillean species represented on the Virgin Islands is Haplopus micropterus (St. Fargeau & Audinet-Serville, 1828; see below). The largest island is Saint Croix with an



175

area of 213 km² which also exhibits the highest elevation of the archipelago, represented by Mount Eagle (355 m). Most of the islands exhibit a tropical climate with plenty of rain and at least the larger islands Saint Croix, Saint Thomas, Saint John and Tortola have areas of tropical rain forest in the central mountains. The Lesser Antilles archipelago forms a chain of moderately sized, volcanic islands along the western border of the Caribbean plate most of which have reasonably high mountainous central regions and exhibit a tropically hot and humid climate with plenty of rainfall. The highest elevations are the Mount Soufrière on Guadeloupe with 1467 m, Morne Diablotin on Dominica with an altitude of 1447 m, Mount Pelée on Martinique with 1397 m, Mount Gimie on Saint Lucia with 958 m and Mount Soufrière on Saint Vincent raising to 1234 m. The Lesser Antilles have never had continental connections and, except for Antigua-Barbuda, Saint Kitts-Nevis-Saint Bartholomé and GrenadaGrenadines, which are joined by shallow banks, the islands have not been connected to each other (Rickfels & Bermingham, 2007: 2395). In studying the Phasmatodea of the West Indies it becomes obvious that almost all genera and species are restricted to a single island, archipelago or a cluster of islands, which particularly applies to the members of Haplopodini. Hence, analyzing the distribution patterns and bringing them in connection with the complex geology of the West Indies is of great importance to define genera and distinguishing species. In his unpublished PhDthesis, Moxey (1972: 183) grouped the phasmatodean genera known to occur on the West Indies into three categories: endemic Antillean, principally Antillean and extra Antillean. However, Moxey’s observations were partially based on taxa attributed to wrong genera, most of which are in fact restricted to South or Central America and extinct in the West Indies. In several cases already, the Antillean species attributed to genera of Moxey's latter two categories, have shown to represent distinct genera that in fact are endemic to the West Indies (e.g. Bacteria Berthold, 1827 or Malacomorpha Rehn, 1906). It follows that, most of the genera that Moxey (1972) stated to be principally Antillean or extra Antillean can be regarded as misinterpreted. As a result, the number of genera endemic to the West Indies is apparently much higher than presumed previously, a fact well emphasized by the here presented study of the Haplopodini and not surprising if the aforementioned observations are taken into account. The biogeographical origin of the Haplopodini is difficult to define and due to the lack of sufficient concrete phylogenetic or molecular data and still controversely discussed colonization of the West Indies any broader discussion would be highly speculative (see above). Also, any discussion on the biogeographical origins or distributional patterns of the other tribes of the subfamily Cladomorphinae discussed above would be premature and shall not be subject to the present study. Hence, such discussion is here restricted solely to the Haplopodini. A few observations and rather obvious hypotheses shall however be mentioned. The tribe is principally endemic to the West Indies and Florida Keys (the latter are generally regarded as belonging to the Nearctic region) and has colonized almost the entire Antillean dominion as defined by Morrone (2006: 478), with only one of the eight genera represented in Central America (eastern Honduras). In the Antilles, the distribution of the tribe is as follows (with the number of recorded species/subspecies in parentheses): Swan Islands (1), Cayman Islands (2), Jamaica (3), Cuba (5), Florida Keys (1), Bahamas (1), Navassa Island (1), Hispaniola (9), Mona Island (1), Puerto Rico (2), Virgin Islands (1), Saint Croix (1), Guadeloupe (1), Dominica (1), Martinique (1), Saint Lucia (1), Saint Vincent & Grenada (1). Seen from these numbers, the greatest diversity of species is found on Hispaniola and Cuba, all the other islands merely having one or two species. The Lesser Antilles are interesting in the aspect that each of the major islands, except for the Grenadines, have their own endemic species or subspecies (see comments on Diapherodes below). Based on the relationships of the known genera and species there appear to have been at least two possible but presumably very early invasions into the West Indies which gave raise to the present Haplopodini, perhaps as early as during the late Tertiary. Consequently, and since the Haplopodini are principally endemic to the West Indies with no particularly close relatives in the surrounding continental areas the clade is likely to represent an ancient relict of the proto-Antillean fauna. The single known Central American taxon Apteroplopus n. gen. in eastern Honduras appears to be of Antillean origin and perhaps colonized this area by over-water dispersal via the Greater Antilles rather than indicating a Central American origin of Haplopodini. This hypothesis is supported by the almost unidirectional northwestward current flow in the West Indies (see above). The closely related Haplopus, Cephaloplopus n. gen. and Parhaplopus n. gen., all of which occupy rather dry lowland habitats, are restricted to the Greater Antilles, Cayman Islands and Bahamas with one species in the Virgin Islands and have radiated into as many as nine known species on Hispaniola alone. Diapherodes has radiated most significantly on the Lesser Antilles with one species on Puerto Rico and two on Jamaica but none on Hispaniola. The closely related


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Venupherodes n. gen. is endemic to northern Cuba. Representatives of these two genera almost exclusively inhabit humid mountainous forests in the central portion of each settled island, leaving the dryer lowland and coastal regions to other taxa. This separation is very obvious on Puerto Rico, an island which harbors one species each of Diapherodes and Haplopus (see below). Although a South American ancestor and invasion into the West Indies from South America appears the most likely scenario for Haplopodini, there are no extant close relatives in northeastern South America, which supports that the clade might have colonized the West Indies very early and is an ancient relict of the proto-Antillean fauna. The only taxon that virtually resembles species of Diapherodes in appearance is the monotypical tribe Cranidiini, but as discussed in detail above close relation between these two tribes as erroneously postulated by Zompro (2004), can be discarded due to fundamental differences in the morphology of the genitalia and eggs (→ 4.2.3). Hence, if the ancestral stock had been more widespread and also reaching into northeastern South America, it is likely that Diapherodes could have originated from such a lineage, which is generally confirmed by the relationships of the known species indicating a dispersal from the Grenadines to as far northwest as Jamaica. The two closely related and rather derived genera Aploploides and Venupherodes n. gen. are endemic on Cuba and more closely related to Diapherodes than to the other four genera, which suggests that they have evolved from the same lineage that gave rise to Diapherodes. On Cuba these two genera are separated by climatic, vegetational and perhaps also dietary preferences, with Venupherodes inhabiting the humid mountainous habitats typical for Diapherodes on other islands, and Aploploides occuring in the dryer lowland regions which in other localities are occupied by Haplopus, Cephaloplopus n. gen. and Parhaplopus n. gen.. The distributional patterns and preferred natural habitats of the eight genera here recognized are discussed in more detail below. Aploploides (Fig. 374). The single known species of this genus, A. stenocephalum Rehn & Hebard, 1938, is restricted to western Cuba (Pinar del Rio Province), where it occurs in lowland regions of semi-arid pine and bamboo-forests or open scrub-land with palmettos. Morphologically it differs considerably from all other genera of Haplopodini by e.g. the completely unarmed, smooth and shiny body as well as the flat, elongate and unarmed head of both sexes. ♀♀ are entirely apterous, a feature only shared with the second Cuban endemic Venupherodes n. gen.. Apteroplopus n. gen. (Fig. 377) This distinctive genus is only known from a unique ♂ of the type-species A. grossedentatus (Brunner v. Wattenwyl, 1907) with no more precise locality than “Honduras”. It is at once distinguished from all other ♂♂ of the tribe by being completely apterous, whereas at least small, scale-like tegmina and alae are present in all other known genera. It is the only member of Haplopodini that is known from Central America, and the type-locality “Honduras” suggests it may, similar to Haplopus in the Greater Antilles, inhabit the flat eastern costal regions of Honduras (and perhaps Nicaragua) facing the Caribbean Sea. According to Morrone (2006: 479) these regions biogeographically belong to the Eastern Central America province, which comprises a narrow strip along the east coast of Central America from Mexico in the north to Guatemala in the south, and is part of the Caribbean Subregion. Unfortunately, nothing is known about the natural habitats of Apteroplopus and the ♀ and egg still await discovery. Cephaloplopus n. gen. (Figs. 377, 379). This fairly derived genus presently contains four known species, three of which occur on Hispaniola and one on Cuba. Habitats in the Dominican Republic include semi-arid scrub-land and mountainous forest in the south (Santo Domingo Province & Azua Province) as well as lowland rain forest in the southwest (northern Azua Province, Pedernales Province and Independencia Province). The altitudinally highest record for the genus is at 800 m in the Sierra Martín García, northern Azua Province. As can be seen from the map (Fig. 316) the distributional pattern is still very poorly known but believed to include most of the lower regions throughout Hispaniola and Cuba. Cephaloplopus appears to prefer habitats similar to those occupied by the closely related Haplopus and in some localities e.g. in the southern Dominican Republic even occurs alongside with Haplopus micropterus (St. Fargeau & Audinet-Serville, 1828). This suggests they are in direct competition, which however may be ruled out by different food-preferences. Unfortunately however nothing is known about the host plants of any of the species of Cephaloplopus. The fact that as many as four new species from a very few rather defined localities are described herein, indicates the genus may have radiated equally successfully in the same habitats as did the closely related Haplopus, which makes the discovery of further still unknown species of Cephaloplopus highly probable once more extensive collections are carried out in so far lesser prospected areas of northern Hispaniola and throughout Cuba. The genus is closely related to Haplopus and Parhaplopus n. gen. but all known species are smaller on average (body length < 10 cm) and remarkably procryptic for the tribe. They are



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well recognized by e.g. the very prominent cephalic horns and widely expanded abdominal tergum VII of both sexes as well as the distinct leg armature of ♀♀. Diapherodes (Fig. 375, 381). This genus divides into two distinct groups of species, here recognised as the gigantea species-group and jamaicensis species-group. These two groups not only differ by morphological features of the insects but are also separated geographically, being either restricted to the Lesser (gigantea species-group, Fig. 381) or Greater Antilles (jamaicensis species-group). Four distinct species and one subspecies are presently recognised in the gigantea species-group. All are endemic to a certain island and hence well separated by geography. Only the large southern D. gigantea (Gmélin, 1789) simultaneously occurs on two islands, namely Grenada and Saint Vincent, and has developed a subspecies on Saint Lucia, the island north of Saint Vincent, here described as D. gigantea saintluciae n. ssp.. The three northward islands have one species each, this is D. martinicensis Lelong & Langlois, 2005 on Marinique, D. dominicae Rehn & Hebard, 1938 on Dominica and D. angulata (Fabricius, 1793) on Guadeloupe (Fig. 381). The latter species on Guadeloupe is only found in the humid forests that cover the slopes of Mount Soufrière in the central portion of Basse Terre. The jamaicensis species-group is restricted to the Greater Antilles but has a remarkably disjunct distribution with one species on Puerto Rico (D. achalus (Rehn, 1904)) and two sympatric species on Jamaica (D. jamaicensis (Drury, 1773) and D. laevicollis Redtenbacher, 1908). The fact that the genus is present on Puerto Rico and Jamaica but is apparently absent on the intervening Hispaniola is interesting and suggests the common ancestral stock of Diapherodes-species may have been more widespread and perhaps also present on Hispaniola at one time. The reasons however why Diapherodes is now extinct on Hispaniola are difficult to define but a possible scenario could be, that it was not sufficiently equipped to compete against the apparently more numerous competitors in its preferred habitats on Hispaniola. All species of the genus occupy the central mountainous forests of the island they are endemic to, leaving the domain of dryer lowland or coastal regions to other taxa. On Jamaica both known species are exclusively found in the humid and tropical forests of the Blue Mountains in the eastern portion of the island and on Puerto Rico D. achalus is found throughout almost the entire tropical and mountainous regions in the northern half of the island. On the Lesser Antilles members of the genus are found in different habitats, including humid mountainous forest, lowland rain-forest and cultivated areas in semi-humid environments (Lelong et al., 2003; Langlois et al., 2006). Neither species exhibits high population densities with specimens occurring rather isolated or at best in small groups on an individual host-plant. The diet of Diapherodes-species includes several common jungle plants in their natural habitats as well as cultivated trees in agricultural areas and plantations, amongst them Anacardium occidentale (Anacardiaceae), Heliconia sp. (Heliconiaceae), Cinnamomum spp. (Lauraceae), Eugenia alpina (Myrtaceae), Guarea ramiflora (Meliaceae), Inga vera (Mimosaceae), Ixora sp. (Rubiaceae), Psidium guajava (Myrtaceae) and Sterculia caribea (Sterculiaceae). Haplopus (Figs. 376, 380). This genus has a wide distribution but has particularly radiated on Hispaniola which harbors four of the eight known species. It extends from the Dry Tortugas, Florida Keys and Bahamas, over Hispaniola to as far east as Mona Island, Puerto Rico and the Virgin Islands, and even has two species on the Cayman Islands and Swan Islands (= Santanillas). All species inhabit lowlands, the habitats ranging from dry coastal regions, over savannahs and dry scrub-land to semi-humid low-land forests. Neither species is found in mountainous regions or particularly humid habitats. While five of the seven known species are rather locally distributed and endemic to a certain island, two species namely H. scabricollis (Gray, 1835) and H. micropterus (St. Fargeau & Audinet-Serville, 1828) have remarkably large distributional ranges which include several rather widely separated islands and archipelagos. In regards to their distribution both species represent exceptions within the entire Haplopodini. H. scabricollis occurs on the Dry Tortugas, Florida Keys and coastal regions of extreme southern Florida, on the Bahamas, Navassa Island, the Cayman Islands and even as far southwest as the Swan Islands (also Islas Santanilla or Islas del Cisne) some 150 km off the coastline of Honduras. The occurrence of H. scabricollis on islands of the Caribbean Sea, which are generally regarded a different biogeographical province (Morrone, 2001), but absence on Cuba is very remarkable and deserves further evaluation. It is not unlikely to occur also along coastlines of Cuba and eastern Hispaniola but so far there have been no definite records from these areas. In its extremely dry and sandy, coastal-near natural habitats H. scabricollis is known to feed almost exclusively on bay cedar (Suriana maritima, Simaroubaceae) and to be excellently camouflaged in these shrubs. Suriana maritima is peculiar to coral reefs or sandy soil and widely spread throughout the coastlines of the Caribbean. Its habitats near the coast make H. scabricollis predestinated for over-water dispersal on flotsam during the frequent heavy hurricanes in the Caribbean, which may explain the remarkably wide and disjunct distributional


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pattern of this particular species. Judging from the number of specimens in museum collections H. scabricollis seems to be fairly abundant in some localities, e.g. Loggerhead Key (Dry Tortugas). Although a very specialized feeder in its natural habitats, H. scabricollis will accept bramble (Rubus spp., Rosaceae), roses (Rosa spp., Rosaceae) or oaks (Quercus spp., Fagaceae) as alternative food plants in captivity. H. micropterus is widespread throughout the lowland regions of Hispaniola but also occurs on Mona Island, Puerto Rico and the Virgin Islands (Saint Thomas, Saint John, Saint Croix & Tortola). Throughout the various recorded localities this species shows a great deal of intraspecific variability in several morphological characters of the insects (Figs. 190–197) but also the eggs (Figs. 219–226). As in all other localities, on Puerto Rico it occupies the lower regions and coastlines, leaving the domain of the high mountainous forests to Diapherodes achalus (Rehn, 1904), hence any direct competition is precluded by altitudinal separation. In the Dominican Republic Haplopus-species are known to feed at least on guava (Psidium guajava, Myrtaceae), “Palo de Tabaco” (Pictetia sulcata & P. spinifolia, Fabaceae) and West Indian Cherry (Malpighia glabra, M. gossipifolia & M. punicifolia, Malpighiaceae). In the Virgin Islands Lu et al. (in preparation) report H. micropterus to feed on Pictetia aculeata (Fabaceae), Piscidia carthagensis (Fabaceae) and Pithecellobium unguis-cati (Fabaceae). Certainly however, several other plants are eaten since these insects are fairly polyphagous in captivity in Europe, accepting oaks (Quercus spp., Fagaceae), roses (Rosa spp., Rosaceae), bramble (Rubus spp., Rosaceae), eucalyptus (Eucalyptus spp., Myrtaceae) or salal (Gaultheria shallon, Ericaceae) as alternative food plants. Paracranidium (Fig. 378). The single known species of this very distinctive genus is small in size (body length < 7 cm), with the body strikingly triangular in cross-section, entirely unarmed and apterous in the ♀♀ (the only sex known). The mesothorax has the longitudinal median keel strongly semi-circularly raised (Fig. 251). Paracranidium seems more closely related to either Aploploides or Diapherodes rather than Haplopus, but takes on a rather isolated position in the Haplopodini, which would tend to imply a long separation from the rest of the fauna. P. pumilio (Westwood, 1843) is restricted to the high mountainous regions of eastern Jamaica (Blue Mountains, e.g. Newton 3000 ft. & Cinchona 5000 ft.), but unfortunately nothing is known about its dietary habits. It must however be presumed, that at one time, the ancestral stock of Paracranidium and its closest relative was more widespread in the West Indies. Perhaps Paracranidium survived only in the high mountains of Jamaica, either because it was too ill-equipped to compete against other competitors (e.g. Diapherodes), or because it is a specialized feeder whose preferred host-plant(s) only survived in its present range. The so far unknown ♂♂ and eggs still await discovery. Parhaplopus n. gen. (Figs. 378, 379). This new genus is very closely related to Haplopus and is here interpreted as the sister-taxon. It however differs by having eggs which resemble those of Cephaloplopus n. gen.. The still very fractionally known distribution generally corresponds to that of Cephaloplopus n. gen., being represented only on Hispaniola (and Gonaive Island) and Cuba. The habitats range from semi-arid and low altitude scrub-land, over agricultural areas to humid mountainous forest below 1000 m, hence it generally inhabits the same dominions as Haplopus and Cephaloplopus n. gen.. The Cuban P. cubensis (Saussure, 1868) even occurs in the same localities as Cephaloplopus alope n. sp. and for this reason they can be presumed sympatric species. Food preferences probably rule out any direct competition between these taxa, but unfortunately nothing is known about the dietary habits of Parhaplopus-species. Venupherodes n. gen. (Fig. 374). The single known species of this genus is closely related to Diapherodes but at once differs by features like the small size, unarmed body and ♀♀ being apterous. These features are only shared with the other Cuban endemic of Haplopodini, the genus Aploploides. Unfortunately, only a very few definite records are known for V. venustula (Audinet-Serville, 1838) but the distribution seems to be restricted to the mountainous rain forests of northwestern Cuba, e.g. the Sierra del Rosario. It obviously leaves the domain of lowland forests and scrub-land to Aploploides, hence these two genera do not seem to be in direct competition because of altitudinal and climatic separation.

7. BIOLOGY & ECOLOGY Members of Haplopodini occur in a wide range of different habitats throughout the Caribbean subregion, including primary forest, secondary forest, montane and lowland forest, semi-arid to dry scrub-land, savannas, agricultural or even urban areas such as public parks, private gardens or small areas of secondary forest or scrub-land towards the



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outskirts of cities. The elevations of recorded localities range from sea-level to about 1500 m, but the greater number of species is restricted to lowland regions often close to the coast rather than montainous habitats. The domain of montane forests is mainly occupied by members of the genus Diapherodes, but the only species exclusively restricted to high altitude montane forest appears to be the striking Paracranidium pumilio (Westwood, 1843) in the Blue Mountains of eastern Jamaica. At the opposite extreme Haplopus scabricollis (Gray, 1835) has settled the very low, dry and sandy coastal regions of the Dry Tortugas, Florida Keys, Bahamas and southern Florida, where it feeds exclusively on the bay cedar Suriana maritima (Simaroubaceae). In general Haplopodini occur in heights between 0.5–3 metres off the ground, usually in undergrowth of forests, on low growing shrubs or small trees and there appears to be no particular specialization to the canopy layer. Principally, these insects are not abundant in their natural habitats and none have so far been mentioned to occur in pest-like numbers, hence they are not of agricultural importance in the West Indies. In most cases the insects are found singularly or as small groups on an individual host-plant. The only species known to be particularly numerous in some localities is Haplopus scabricollis (Gray, 1835). For instance (Stockard, 1908b: 47) encountered six adult ♀♀ and two ♂♂ on a group of Suriana-bushes only some 3,5 metres in diameter on Loggerhead Key, Dry Tortugas, and the collection of UMMZ contains a series of 76 specimens all collected during a single month on this tiny island, which has an area of only 260.000m2. The natural diet of Haplopodini includes several common native plants, as well as cultivated trees in agricultural areas, plantations or public parks. Plants known to serve as food are e.g. Anacardium occidentale (Anacardiaceae), Heliconia sp. (Heliconiaceae), Cinnamomum spp. (Lauraceae), Eugenia alpina (Myrtaceae), Guarea ramiflora (Meliaceae), Inga vera (Mimosaceae), Ixora sp. (Rubiaceae), Malpighia glabra (Malpighiaceae), Malpighia punicifolia (Malpighiaceae), Malpighia gossipifolia (Malpghiaceae), Pictetia sulcata (Fabaceae), Pictetia spinifolia (Fabaceae), Pictetia aculeata (Fabaceae), Piscida carthagensis (Fabaceae), Pithecellobium unguis-cati (Fabaceae), Bidens cynapiifolia (Asteraceae), Psidium guajava (Myrtaceae), Sterculia caribea (Sterculiaceae), Cichona ledgeriana (Rubiaceae), Coccoloba uvifera (Polygonaceae) and Suriana maritima (Simaroubaceae). Alternative food plants accepted in captivity in Europe include oaks (Quercus spp., Fagaceae), roses (Rosa spp., Rosaceae), bramble and raspberry (Rubus spp., Rosaceae), beech (Fagus sylvaticus, Fagaceae), eucalyptus (Eucalyptus spp., Myrtaceae), St. John's Wort (Hypericum patulum & H. androsaemum, Hypericaceae) or salal (Gaultheria shallon, Ericaceae). The fact that these plants are eaten in captivity makes it highly probable that also West Indian representatives of the Rosaceae, Ericaceae or Hypericaceae serve as food plants for certain species, but more detailed observations are needed for confirmation. As for most Phasmatodea, members of Haplopodini are nocturnal in their natural habitats, the insects making use of their camouflage and resting almost motionless on the host plants during the day. Feeding, moulting and copulation mostly takes place at night. Defensive reactions include remaining motionless, dropping to the ground and quickly walking or flying (winged ♂♂) away. Some species (e.g. Diapherodes gigantea or D. jamicensis) even exhibit conspicuous startle-displays which include flashing the wings and pinching at the predator with the spiny hind-legs. Mostly adults are fairly long-lived Phasmatodeans with ♀♀ of some species (e.g. Diapherodes gigantea) reaching up to one year of age, whereas ♂♂ usually live 5–8 months. Eggs are simply dropped to the ground singularly, but ♀♀ usually make use of their elongated subgenital plate to flick the eggs away and ensure optimum dispersal. The number of eggs produced per day and ♀ depends on the relative size of the egg in relation to the adult insects. Some species of Haplopus can produce as many as ten eggs per day (Adams & Adams, 1982), while species of Diapherodes or Venupherodes n. gen. produce an average of only 3–4 days per day and ♀. The total number of eggs produced by a single ♀ in an entire lifetime ranges from 200–800, the hatching rates usually being fairly high. Adams & Adams (1982) reported adults of Haplopus bicuspidatus de Haan, 1842 to emit a punget smell from the prothoracic defensive glands when disturbed. Two very detailed studies on the habits, reactions and biology of Haplopus scabricollis (Gray, 1835) were published by Stockard (1908a, 1908b).

8. SUMMARY The present paper provides a detailed taxonomic revision of the Caribbean tribe Haplopodini Günther, 1953 at the species-level, which is here re-established as a valid taxon. Four new genera, nine new species and one new subspecies are described (→ 8.2); six of the newly described species are from Hispaniola, two from Cuba and one


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from Cayman Brac, the northernmost of the Cayman Islands. Seven of the eight known genera are restricted to the West Indies and only one poorly known genus (Apteroplopus n. gen.) occurs along the east-coast of Honduras, being the only representative in Central America. The distributional pattern of the tribe and the eight contained genera is discussed in detail and shows the tribe to have radiated most strongly on Hispaniola and Cuba which harbour the largest number of known species (→ 6). The results show the phasmid fauna of the West Indies is still poorly studied and the faunas of Cuba and northern portions of Hispaniola in particular are certainly up to now just fractionally known. Despite the fact that Haplopodini are the largest and most striking phasmatodeans of the West Indies, the poor degree of exploration of these regions is emphasized by the descriptions of numerous new and previously unknown taxa. Only three valid species of Haplopodini (all others are here shown to be synonyms) had previously been recorded from Hispaniola, and so the six new Hispaniolan species described herein increase the number of known species from Hispaniola by 200%. Although the present paper studies the materials from collections at over 300 sites throughout the area of the Dominican Republic, it is very likely that further new species will be discovered once less prospected areas are surveyed in more detail. The western and northern portions of Hispaniola in particular should harbor further unknown species, since only a few collections have so far been conducted there. The fauna of Cuba, the largest of the Antillean Islands, is even less known, since as many as three genera of Haplopodini are represented (two of which are endemic) but only five species of the tribe have been described from the island so far. With certainty several new species await discovery and scientific recognition especially throughout the eastern portions of Cuba. According to this revision 26 valid species and one subspecies of Haplopodini have so far been recognized (→ 8.1), six of which are only known from a single sex or the original type-specimen(s), with the opposite sex still unknown. Also the eggs of as many as 13 species still await discovery and formal description. One species, Haplopus similis (Rehn, 1904) from the island of Santanilla, is only available from an immature type-specimen and remains entirely obscure. In addition to the taxonomy and diversity also our present knowledge of the biology, habits, habitats and natural food plants of Haplopodini are still rather limited, with sufficient information so far only available for a very few species. It is therefore hoped that future studies will not only be restricted to the recognition of new taxa but will also provide descriptions of the so far unknown sexes and eggs, new records to understand the distributional pattern of the tribe and its sub-ordinate taxa, and also data to broaden our knowledge on the biology as well as the ecological and dietary preferences of these insects, which are the largest and perhaps most beautiful phasmatodeans of the entire Caribbean subregion. A survey of the subfamily Cladomorphinae (→ 4) has shown several of the taxonomic changes proposed by Zompro (2004) to be wrong and poorly or not at all supported, and conclusions that correspond to the results of phylogenetic studies and molecular data (Whiting et al., 2003; Bradler, 2009; Buckley et al., 2009; Buckley et al., 2010). Cladomorphinae currently comprises six tribes, which form two well defined subordinate groups: Cladomorphini + Cladoxerini + Cranidiini (here termed the Cladomorphinae sensu stricto) and Pterinoxylini n. trib. + Hesperophasmatini + Haplopodini. Pterinoxylini n. trib. + Hesperophasmatini are likely to be the sistergroup of Haplopodini. The three tribes Hesperophasmatini + Pterinoxylini n. trib. + Haplopodini are very closely related and might form a monophyletic clade that possibly is the sister-group of Cladomorphinae sensu stricto (Fig. 409), because they all differ by the lack of a median line in the eggs, dorsally furcate meso- and metabasitarsi as well as the lack of a gonoplac and lack of the elongated gonapophyses VIII of ♀♀ seen in other Cladomorphinae. The tribe Baculini Günther, 1953 is shown to be a synonym of Cladoxerini Karny, 1923. Cladoxerini differs from all other Cladomorphinae, including the suggested Hesperophasmatini + Pterinoxylini n. trib. + Haplopodini clade, by a good number of morphological features such as the dorsally serrate profemora and conspicuously shortened antennae of ♀♀, which consist of no more than 30 antennomeres (→ 4.2.1). Cranidiini Günther, 1953 is shown to be a monotypical and exclusively South American tribe that differs from all other Cladomorphinae by a good number of morphological features, such as the tectiform mesosternum and entirely unarmed legs of both sexes, strongly laterally dilated and rhombic cross-section of the body of ♀♀ as well as the strongly enlarged poculum and minutely spinulose phallus of ♂♂ (→ 4.2.3). Hence, all Caribbean genera wrongly attributed to Cranidiini by Zompro (2004) are here removed from that tribe and Haplopodini Günther, 1953 is re-established to comprise these misplaced Caribbean genera. The new tribe Pterinoxylini n. trib. is introduced to separate the typegenus Pterinoxylus Audinet-Serville, 1838 from the remainder of Hesperophasmatini (→ 4.2.5). This is based on a good number of morphological characters such as the much larger size, more slender body and strongly lobed and lamellate protibiae of both sexes, presence of a tympanal region in the basal portion of the alae of ♀♀, dorsally



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lobed probasitarsus of ♂♂ and elongate, alveolar eggs, which bear peripheral opercular and polar extensions. Pterinoxylini n. trib. is believed to be the sister-taxon of Hesperophasmatini, since e.g. both tribes possess conspicuous sensory areas on the profurcasternum otherwise lacking in all other taxa discussed and currently attributed to Cladomorphinae (Fig. 409). The tribe Hesperophasmatini Bradley & Galil, 1977 is removed from Pseudophasmatidae: Xerosomatinae and transferred back to Cladomorphinae (→ 4.2.4). Phylogenetic analyses (e.g. Whiting et al., 2003; Buckley et al., 2009; Buckley et al., 2010) have provided support for a sister-group relationship between the Cladomorphinae and the tribe Stephanacridini Günther, 1953, an Old World clade that is today mostly restricted to some of the Pacific Islands, New Guinea and parts of Wallacea. Some characters supporting this sister-group relationship are the strongly elongated gonapophyses VIII and presence of gonoplacs in ♀♀, which are shared by Stephanacridini and the subordinate Cladomorphinae taxa Cladomorphini, Cranidiini and Cladoxerini. This postulates a Gondwanan origin for the Cladomorphinae and suggests invasion of its ancestors via Gondwanan land connections from the Australasian region through Antarctica to South America perhaps during the late Cretaceous or late Tertiary (see above). Close relation of Cladomorphinae to African taxa of the subfamily Palophinae Kirby, 1896 hypothesized above definitely deserves further evaluation but invasion of potential ancestors from Africa perhaps during the early Oligocene, e.g. from West-Africa to northeastern Brazil, is a hypothesis that should be taken into account in future phylogenetic or molecular studies, since there are striking morphologically parallels between the Neotropical Cladomorphinae and African Palophinae. Possible support for this hypothesis is represented by certain Neotropical rodents, primates, bats and turtles that are known to be most closely related to African taxa (e.g. Flynn & Wyss, 1998). The actual relationships within Cladomorphinae still remain somewhat obscure and also the definition of the subfamily is still not clarified, hence Cladomorphinae is most likely not a natural group in its present composition and recognition (→ 4). This is partly emphasized by the tree presented herein (Fig. 409), which is meant to show the possible relationships within present Cladomorphinae and to provide a basis and starting point for future studies. Cladomorphini + Cladoxerini + Cranidiini may be regarded as the Cladomorphinae sensu stricto but this clade might be paraphyletic, since further taxa currently attributed to other subfamilies (e.g. Diapheromerinae and Heteronemiinae) appear to be closely related (Hennemann & Conle, in preparation). Pterinoxylini n. trib. + Hesperophasmatini + Haplopodini form another clade that differs from Cladomorphinae sensu stricto by the lack of a median line in the eggs as well as the lack of gonoplacs and having the gonapophyses VIII of ♀♀ not conspicuously elongated (Fig. 409). However, extensive research, comprehensive phylogenetic studies and much more molecular data will be required for any broader discussion. Future phylogenetic or molecular studies on the New World Phasmatodea but Cladomorphinae in particular should in addition to African taxa of the subfamily Palophinae also include taxonomically and phylogenetically important “key genera” such as Cladomorphus, Cranidium, Haplopus, Cladoxerus or Hesperophama, since only examination of the type-genera of higher ordinate taxa will allow confirmed decisions on the systematic positions of these traditional subfamilies and tribes. As can be seen from the results and hypotheses presented herein, the higher classification of the New World anareolatae Phasmatodea is still far away from a natural classification based on phylogenetic and molecular analyses. It is therefore hoped, that future studies in these disciplines will also comprise the taxa here proposed in order to achieve a better understanding of the origins and relationships within the New World Phasmatodea.

8.1. Taxonomic checklist of Haplopodini Günther, 1953 Aploploides Rehn & Hebard, 1938 Type-species: Aploploides stenocephalum Rehn & Hebard, 1938: 49, by original designation of Rehn & Hebard, 1938: 49) 1. Aploploides stenocephalum Rehn & Hebard, 1938: 49, pl. 4: 18–21 (♂, ♀). Apteroplopus n. gen. Type-species: Dyme grosse-tuberculata Brunner v. Wattenwyl, 1907: 323, by present designation 1. Apteroplopus grosse-tuberculatus (Brunner v. Wattenwyl, 1907: 323) [Dyme].


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Cephaloplopus n. gen. Type-species: Cephaloplopus pulchellus n. sp., by present designation 1. Cephaloplopus alope n. sp. 2. Cephaloplopus euchlorus n. sp. 3. Cephaloplopus laetus n. sp. 4. Cephaloplopus pulchellus n. sp. Diapherodes Gray, 1835 Type-species: Mantis gigantea Gmélin, 1789: 2055, by subsequent designation of Kirby, 1904a: 362. 1. Diapherodes achalus (Rehn, 1904: 68) [Aplopus]. n. comb. = Diapherodes longiscapha Redtenbacher, 1908: 435. n. syn. 2. Diapherodes angulata (Fabricius, 1793: 13) [Mantis]. rev. stat. = Haplopus grayi Kaup, 1871: 36. n. syn. 3. Diapherodes dominicae (Rehn & Hebard, 1938: 53). n. comb. 4. Diapherodes gigantea gigantea (Gmélin, 1789: 2055) [Mantis]. = Mantis gigas Drury, 1773: 89, pl. 50 (♀). = Cyphocrana cornuta St. Fargeau & Audinet-Serville, 1825: 445. 5. Diapherodes gigantea saintluciae n. ssp. 6. Diapherodes jamaicensis (Drury, 1773: 88, pl. 44: 1 (♂)) [Mantis]. n. comb. = Mantis bispinosa Fabricius, 1775: 274. = Haplopus christopheri Westwood, 1859: 84, pl. 33: 4, 4a (♀). n. syn. = Diapherodes glabricollis Gray, 1835: 33. n. syn. = Haplopus murinus Redtenbacher, 1908: 429. n. syn. = Dipaherodes pulverulentus Gray, 1835: 34. n. syn. 7. Diapherodes laevicollis Redtenbacher, 1908: 434. 8. Diapherodes martinicensis Lelong & Langlois, 2005: 264. Haplopus Burmeister, 1838 Type-species: Cyphocrana micropterus St. Fargeau & Audinet-Serville, 1825: 445, by monotypy. = Aplopus Gray, 1835: 13, 34. 1. Haplopus bicuspidatus de Haan, 1842: 128. 2. Haplopus brachypterus n. sp. 3. Haplopus intermedius n. sp. 4. Haplopus micropterus (St. Fargeau & Audinet-Serville, 1828: 445) [Cyphocrana]. = Phasma angulata Stoll, 1813: 61, pl. 21: 77 (♀). = Haplopus bituberculatum de Haan, 1842: 128. n. syn. = Haplopus cythereus Westwood, 1859: 86, pl. 18: 5, 5a & b (♂). n. syn. = Haplopus ligia Westwood, 1859: 89, pl. 1: 1, 1a, 1b (♂) & 2, 2a (♀). n. syn. = Haplopus ligiolus Redtenbacher, 1908: 430. n. syn. = Haplopus obtusus Redtenbacher, 1908: 431. n. syn. = Diapherodes spinipes Gray, 1835: 34. n. syn. 5. Haplopus scabricollis (Gray, 1835: 34) [Diapherodes]. = Aplopus mayeri Caudell, 1905: 83. n. syn. = Aplopus similis (Rehn, 1904: 65). n. syn. 6. Haplopus sobrinus n. sp. 7. Haplopus woodruffi n. sp. Paracranidium Brock, 1998 Type-species: Diapherodes (Cranidium) pumilio Westwoood, 1843: 50, pl. 61: 2, by original designation of Brock, 1998: 28. 1. Paracranidium pumilio (Westwood, 1843: 50, pl. 61: 2 (♀)) [Diapherodes (Cranidium)].



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Parhaplopus n. gen. Type-species: Haplopus cubensis Saussure, 1868: 68, by present designation) 1. Parhaplopus cubensis (Saussure, 1868: 68) [Haplopus]. n. comb. 2. Parhaplopus evadne (Westwood, 1859: 85, pl. 18: 6, 6a & b (♂)) [Haplopus]. n. comb. 3. Parhaplopus navarroi n. sp. Venupherodes n. gen. Type-species: Platycrana venustula Audinet-Serville, 1838: 242, by present designation 1. Venupherodes venustula (Audinet-Serville, 1838: 242) [Platycrana]. n. comb. = Phasma havaniense Westwood, 1859: 34, pl. 22: 7, 7a, 7b (♀). = Haplopus juvenis Redtenbacher, 1908: 430, pl. 20: 4 (♂). n. syn.

8.2. List of new taxa Pterinoxylini n. trib. Apteroplopus n. gen. Cephaloplopus n. gen. Parhaplopus n. gen. Venupherodes n. gen. Cephaloplopus alope n. sp. Cephaloplopus euchlorus n. sp. Cephaloplopus laetus n. sp. Cephaloplopus pulchellus n. sp. Haplopus brachypterus n. sp. Haplopus intermedius n. sp. Haplopus sobrinus n. sp. Haplopus woodruffi n. sp. Parhaplopus navarroi n. sp. Diapherodes gigantea saintluciae n. ssp.

8.3. List of taxonomic changes 1. Aploploides Rehn & Hebard, 1938: 49 (Type-species: Aploploides stenocephalum Rehn & Hebard, 1938: 49), removed from Cranidiini Günther 1953 and transferred to the tribe Haplopodini rev. stat. [→ 5] 2. Aplopus similis (Rehn, 1904: 65), synonymised with Haplopus scabricollis (Gray, 1835) (n. syn.) [Valid name: Haplopus scabricollis (Gray, 1835) → 5.5] 3. Bacteria grossetuberculata (Brunner v. Wattenwyl, 1907: 318), removed from Bacteria Berthold, 1827 and transferred to Apteroplopus n. gen. [Valid name: Apteroplopus grossetuberculatus (Brunner v. Wattenwyl, 1907) n. comb. → 5.2] 4. Baculini Günther, 1953: 555 (Type-genus: Baculum Saussure, 1861: 127), synonymised with Cladoxerini Karny, 1923: 237 (n. syn.) [→ 4.2.1] 5. Baculum Saussure, 1861: 127 (Type-species: Bacillus (Baculum) ramosum Saussure, 1861: 127), synonymised with Cladoxerus St. Fargeau & Audinet-Serville, 1827: 445 (n. syn.) [→ 4.2.1] 6. Diapherodes Gray, 1835: 33 (Type-species: Mantis gigantea Gmélin, 1789: 2055), removed from Cranidiini Günther 1953 and transferred to the tribe Haplopodini rev. stat. [→ 5] 7. Diapherodes christopheri (Westwood, 1859: 84), synonymised with Diapherodes jamaicensis (Drury, 1773) (n. syn.) [Valid name: Diapherodes jamaicensis (Drury, 1773) → 5.4.2] 8. Diapherodes gigantea dominicae Rehn & Hebard, 1938: 53, removed from synonymy with Diapherodes gigantea (Gmélin, 1789) and established as a valid species of [Valid name: Diapherodes dominicae (Rehn & Hebard, 1838) n. stat. → 5.4.1]


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9. Diapherodes longiscapha Redtenbacher, 1908: 435, synonymised with Diapherodes achalus (Rehn, 1904) (n. syn.) [→ 5.4.2] 10. Diapherodes venustula (Audinet-Serville, 1838: 242), removed from Diapherodes Gray, 1835 and transferred to the genus Venupherodes n. gen. [Valid name: Venupherodes venustula (Audinet-Serville, 1838) n. comb. → 5.8] 11. Haplopodini Günther, 1953: 557 (Type-genus: Haplopus Burmeister, 1838: 576), re-established as a valid tribe (rev. stat.) [→ 5] 12. Haplopus Burmeister, 1838: 560, 576 (Type-species: Cyphocrana micropterus St. Fargeau & Audinet-Serville, 1825: 445), removed from Cranidiini Günther 1953 and transferred to the tribe Haplopodini rev. stat. [→ 5] 13. Haplopus bituberculatum de Haan, 1842: 128 (Phasma (Haplopus)), synonymised with Haplopus micropterus (St. Fargeau & Audinet-Serville, 1825) (n. syn.) [Valid name: Haplopus micropterus (St. Fargeau & Audinet Serville, 1825) → 5.5] 14. Haplopus cubensis Saussure, 1868: 68, removed from Haplopus Burmeister, 1838 and transferred to the genus Parhaplopus n. gen. [Valid name: Parhaplopus cubensis (Saussure, 1868) n. comb. → 5.7] 15. Haplopus cytherea Westwood, 1859: 86, synonymised with Haplopus micropterus (St. Fargeau & AudinetServille, 1828) (n. syn.) [Valid name: Haplopus micropterus (St. Fargeau & Audinet-Serville, 1828) → 5.5] 16. Haplopus dubius (Gray, 1835: 34), removed from Haplopus Burmeister, 1838. The generic placement remains uncertain, but here provisionally transferred to Phanocloidea Zompro, 2001 [Valid name: Phanocloidea dubia (Gray, 1835) n. comb. → 5.9] 17. Haplopus evadne Westwood, 1859: 85, removed from Haplopus Burmeister, 1838 and transferred to the genus Parhaplopus n. gen. [Valid name: Parhaplopus evadne (Westwood, 1859) n. comb. → 5.7] 18. Haplopus glabricollis (Gray, 1835: 33), synonymised with Diapherodes jamaicensis (Drury, 1773) (n. syn.) [Valid name: Diapherodes jamaicensis (Drury, 1773) → 5.4.2] 19. Haplopus grayi Kaup, 1871: 36, removed from synonymy with Diapherodes gigantea (Gmélin, 1789) and synonymised with Diapherodes angulata angulata (Fabricius, 1793) (n. syn.) [Valid name: Diapherodes angulata angulata (Fabricius, 1793) → 5.4.1] 20. Haplopus jamaicensis (Drury, 1773: 88), removed from Haplopus Burmeister, 1838 and transferred to the genus Diapherodes Gray, 1835 (n. comb.) [Valid name: Diapherodes jamaicensis (Drury, 1773) → 5.4.2] 21. Haplopus juvenis Redtenbacher, 1908: 430, synonymised with Venupherodes venustula (Audinet-Serville, 1838) n. comb. (n. syn.) [Valid name: Venupherodes venustula (Audinet-Serville, 1838) n. comb. → 5.8] 22. Haplopus ligia Westwood, 1859: 89, synonymised with Haplopus micropterus (St. Fargeau & AudinetServille, 1828) (n. syn.) [Valid name: Haplopus micropterus (St. Fargeau & Audinet-Serville, 1828) → 5.5] 23. Haplopus ligiolus Redtenbacher, 1908: 432, synonymised with Haplopus micropterus (St. Fargeau & AudinetServille, 1828) (n. syn.) [Valid name: Haplopus micropterus (St. Fargeau & Audinet-Serville, 1828) → 5.5] 24. Haplopus mayeri (Caudell, 1905: 83), synonymised with Haplopus scabricollis (Gray, 1835) (n. syn.) [Valid name: Haplopus scabricollis (Gray, 1835) → 5.5] 25. Haplopus murinus Redtenbacher, 1908: 430, synonymised with Diapherodes jamaicensis (Drury, 1773) (n. syn.) [Valid name: Diapherodes jamaicensis (Drury, 1773) → 5.4.2] 26. Haplopus obtusus Redtenbacher, 1908: 431, synonymised with Haplopus micropterus (St. Fargeau & AudinetServille, 1828) (n. syn.) [Valid name: Haplopus micropterus (St. Fargeau & Audinet-Serville, 1828) → 5.5] 27. Haplopus pulverulentus (Gray, 1835: 34), synonymised with Diapherodes jamaicensis (Drury, 1773) (n. syn.) [Valid name: Diapherodes jamaicensis (Drury, 1773) → 5.4.2] 28. Haplopus spinipes (Gray, 1835: 34), a replacement name for Phasma angulata Palisot de Beauvois, 1805: 166 (not Fabricius, 1793, not Stoll, 1813), hence a synonym of Haplopus micropterus (St. Fargeau & AudinetServille, 1828) (n. syn.) [Valid name: Haplopus micropterus (St. Fargeau & Audinet-Serville, 1828) → 5.5] 29. Laciphorus Redtenbacher, 1908: 351 (Type-species: Laciphorus lobulatus Redtenbacher, 1908: 351, by monotypy), here removed from Cladomorphinae: Hesperophasmatini and transferred to Diapheromeridae: Diapheromerinae: Diapheromerini [→ 4.2.4] 30. Mantis angulata Fabricius, 1793: 13, removed from synonymy with Diapherodes gigantea (Gmélin, 1789) and re-established as a valid species [Valid name: Diapherodes angulata angulata (Fabricius, 1793) rev. stat. → 5.4.1] 31. Paracranidium Brock, 1998b: 28 ( Type-species: Diapherodes (Cranidium) pumilio Westwood, 1843: 50),



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removed from Cranidiini Günther 1953 and transferred to the tribe Haplopodini rev. stat. [→ 5] 32. Pterinoxylus Audinet-Serville, 1838: 226 (Type-species: Pterinoxylus difformipes Audinet-Serville, 1838: 227 (= Haplopus eucnemis Burmeister, 1838: 577)), removed from Hesperophasmatini Bradley & Galil, 1977: 188 and transferred to Pterinoxylini n. trib. [→ 4.2.5] 33. Tersomia Kirby, 1904b: 431 (Type-species: Tersomia brasiliensis Kirby, 1904a: 432), here removed from Cladomorphinae: Hesperophasmatini and transferred to Heteromenimiidae: Heteronemiinae: Paraleptyniini [→ 4.2.4] 34. Wattenwylia Toledo Piza, 1938: 54. (Type-species: Wattenwylia foliata Toledo-Piza, 1938: 6, by original designation), here removed from Pachymorphinae: Gratidiini and transferred to Cladomorphinae: Cladoxerini [→ 4.2.1]

8.4. List of designated neotypes and lectotypes Neotypes: 1. Mantis angulata Fabricius, 1793: 13. [♀ in MNCN] 2. Platycrana venustula Audinet-Serville, 1838: 242. [♀ in MNHU] Lectotypes: 1. Diapherodes longiscapha Redtenbacher, 1908: 435. 2. Diapherodes scabricollis Gray, 1835: 34. 3. Haplopus christopheri Westwood, 1859: 84, pl. 33: 4, 4a (♀). 4. Haplopus cytherea Westwood, 1859: 86, pl. 18: 5, 5a, 5b (♂). 5. Haplopus juvenis Redtenbacher, 1908: 430, pl. 20: 4 (♂). 6. Haplopus ligia Westwood, 1859: 89, pl. 11: 1, 1 a, 1 b (♂) & 2, 2a (♀). 7. Haplopus ligiolus Redtenbacherk, 1908: 432. 8. Haplopus murinus Redtenbacher, 1908: 429.

9. ACKNOWLEDGEMENTS We would like to express our thanks to the curators and staff of the following museums and institutions for access to their collections, providing useful information and the loan of specimens: Dr. Tony Robillard, Simon Poulain & Emmanuel Delfosse (MNHN, Paris); Dr. Susanne Randolf & Dr. Ulrike Aspöck (NHMW, Vienna), Dr. M. Brancucci & Dr. D. Burckhardt (NHMB, Basel), Dr. J. Cools (ISNB, Bruxelles), Dr. Michael Ohl & Ulrike Dorandt (ZMHB, Berlin), Dr. P. Schwendinger (MHNG, Geneva), Dr. George Beccaloni & Mrs. Judith Marshall (NHMUK, London), Dr. George McGavin & Dr. Darren J. Mann (OXUM, Oxford), Dr. Wolfgang Schneider (HLDH, Darmstadt), Prof. Klaus Schönitzer & Tanja Kothe (ZSMC, Munich), Prof. Herbert Strümpel & Dr. Kai Schütte (ZMUH, Hamburg), Dr. Caroline Pepermans (RMNH, Leiden), Jason Weintraub, Dr. John Gelhaus & Dr. Daniel Otte (ANSP, Philadelphia) and Ruth H. Bastardo (IIBZ, Santo Domingo). Dr. R. E. Woodruff and Dr. Michael C. Thomas (FSCA, Gainesville) kindly arranged loans of material from FSCA. Judith Marshall (NHMUK) is also thanked for her efforts in assisting with the search for Drury’s types in the collection of the Natural History Museum, London (NHMUK) and measuring the specimens. Furthermore, we thank biologists Ruth Bastardo (IIBZ, Santo Domingo) and Brígido Hierro (Departamento de Vida Silvestre, Ministerio de Medio Ambienty y Recursos Naturales, Santo Domingo) for their enthusiastic participation as field assistants in the Hispaniolan Orthopteroids Project which greatly benefited from their great expertise with logistics and fieldwork. Collecting and export permits were graciously provided by the Departamento de Vida Silvestre, Ministerio de Medio Ambiente y Recursos Naturales, Santo Domingo. We also acknowledge National Science Foundation grant DEB-0103042 to Dr. Daniel Otte (ANSP) and Dr. Daniel PerezGelabert (USNM) to survey the Hispaniolan fauna of orthopteroid insects. Biology students América Sánchez and Candy Ramírez kindly supplied photos and data of specimens in the collection of the IIBZ (Santo Domingo). Paul D. Brock (NHMUK) shall be thanked for help in many ways, e.g. providing photos of Gray's types in LSUK and the types of Fabricius in HMUG as well as information on Haplopus micropterus from the British


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Virgin Islands. Yannick Bellanger (Trédias, France) and Philippe Lelong (Sainte-Foy d´Aigrefeuille, France) are thanked for data and photos of the specimens of Diapherodes collected by ASPER on Dominica and Saint Lucia as well as culture-stock of Diapherodes martinicensis. Bruno Kneubühler (Luzern, Switzerland) kindly supplied eggs of Diapherodes martinicensis. Kristien Rabaey & Rob Simoens (Veurne, Belgium) generously provided the authors with culture-stock of Diapherodes gigantea and Venupherodes venustula. Mieke Duytschaever (Belgium) donated a captive reared gynandromorph of D. gigantea and provided pictures. Carlos de Soto Molinari (Santo Domingo) is thanked for his contribution of a photo of a live female Haplopus micropterus taken in the San Cristobál Province and Eladio Fernández (Santo Domingo) for his contribution of a photo of a male Parhoplopus evadne taken in Haiti. Mel Rívera (Puerto Rico) kindly provided pictures of live Diapherodes achalus taken at Maricao, Puerto Rico.

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FIGURES 334–341. Ventral view of abdominal sternum VII of ♀♀ showing the praeopercular organ. 334. Aploploides stenocephalum Rehn & Rehn, 1938 ♀ PT: NW-Cuba, Pinar del Rio [ANSP]; 335. Cephaloplopus alope n. gen., n. sp. ♀ HT: Cuba, Camagüey [ANSP]; 336. Cephaloplopus euchlorus n. gen., n. sp., ♀ PT: Dominican Republic, Santo Domingo Province, Sierra Prieta [USNM]; 337. Cephaloplopus pulchellus n. gen., n. sp., ♀ PT: Dominican Republic, Pedernales Province, Boca de la Cañada [USNM]; 338. Haplopus bicuspidatus de Haan, 1842 ♀: captive reared from Dominican Republic [coll. FH, No. 0342-6] 339. Haplopus brachypterus n. sp.. ♀ PT: Dominican Republic, Santiago Province [USNM]; 340. Haplopus micropterus (St. Fargeau & Audinet-Serville, 1828) ♀: Dominican Republic, Pedernales Province, Boca de la Cañada [USNM]; 341. Haplopus scabricollis Gray, 1835 ♀: Loggerhead Key [ANSP].


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FIGURES 342–348. Ventral view of abdominal sternum VII of ♀♀ showing the praeopercular organ. 342. Haplopus sobrinus n. sp. ♀ HT: Cuba, Camagüey [ANSP]; 343. Parhaplopus navarroi n. gen., n. sp. ♀ PT: Dominican Republic, Santo Domingo Province, Santo Domingo [USNM]; 344. Parhaplopus cubensis (Saussure, 1868) n. gen., n. comb. ♀: Cuba, Havana [MNHN]; 345. Diapherodes venustula (Audinet-Serville, 1838) n. gen., n. comb. ♀: captive reared from NW-Cuba, Pinar del Rio [coll. FH, No. 0599-3]; 346. Diapherodes angulata (Fabricius, 1793) ♀ NT: Guadeloupe [MNHN]; 347. Diapherodes dominicae Rehn & Hebard, 1938 ♀ HT: Dominica [ANSP]; 348. Diapherodes gigantea gigantea (Gmélin, 1789) ♀: captive reared from Grenada [coll. FH, No. 0359-1].



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FIGURES 349–356. Ventral view of abdominal sternum VII of ♀♀ showing the praeopercular organ and anal segment of ♂♂ showing the vomer. 349. Diapherodes martinicensis Lelong & Langlois, 2005 ♀: captive reared from Martinique [coll. FH, No. 0750-1]; 350. Diapherodes achalus (Rehn, 1904) ♀: Puerto Rico, Mayaguez [ANSP]; 351. Diapherodes jamaicensis (Drury, 1773) ♀: captive reared from Jamaica [coll. FH, No. 0428-17]; 352. Aploploides stenocephalum Rehn & Rehn, 1938 ♂ PT: NW-Cuba, Pinar del Rio [ANSP]; 353. Cephaloplopus euchlorus n. gen., n. sp., ♂ PT: Dominican Republic, Santo Domingo Province, Sierra Prieta [USNM]; 354. Cephaloplopus pulchellus n. gen., n. sp., ♂ HT: Dominican Republic, Pedernales Province, Boca de la Cañada [USNM]; 355. Parhaplopus evadne (Westwood, 1859) n. gen., n. comb. ♂ HT: Dominican Republic, Santo Domingo [NHMUK]; 356. Parhaplopus navarroi n. gen., n. sp. ♂ HT: Dominican Republic, Monseñor Nouel Province, Bonao [USNM].


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FIGURES 357–364. Ventral view of anal segment of ♂♂ showing the vomer. 357. Haplopus bicuspidatus deHaan, 1842 ♂: captive reared from Dominican Republic [coll. FH, No. 0342-8]; 358. Haplopus brachypterus n. sp. ♂ HT: Dominican Republic, Santiago Province [USNM]; 359. Haplopus intermedius n. sp., ♂ HT: Dominican Republic, Valverde Province, nr. Maizal [USNM]; 360. Haplopus micropterus (St. Fargeau & Audinet-Serville, 1828) ♂: captive reared from Dominican Republic [coll. FH, No. 0243-90]; 361. Haplopus scabricollis Gray, 1835 ♂: Loggerhead Key [ANSP]; 362. Haplopus scabricollis Gray, 1835 ♂: Vanassa Island [USNM]; 363. Haplopus woodruffi n. sp. ♂ HT: Cayman Islands, Cayman Brac [FSCA]; 364. Venupherodes venustula (Audinet-Serville, 1838) n. gen., n. comb. ♂: captive reared from NW-Cuba, Pinar del Rio [coll. FH, No. 0599-1].



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FIGURES 365–372. Ventral view of anal segment of ♂♂ showing the vomer. 365. Diapherodes angulata (Fabricius, 1793) ♂: Guadeloupe [MNHN]; 366. Diapherodes dominicae Rehn & Hebard, 1938 V: Guadeloupe, Picard River track [coll. ASPER]; 367. Diapherodes gigantea saintluciae n. ssp. ♂ PT: Saint Lucia, Barre de l’Isle Trail [coll. ASPER]; 368. Diapherodes gigantea gigantea (Gmélin, 1789) ♂: captive reared from Grenada [coll. FH, No. 0359-34]; 369. Diapherodes martinicensis Lelong & Langlois, 2005 ♂: captive reared from Martinique [coll. FH, No. 0750-5]; 370. Diapherodes achalus (Rehn, 1904) ♂: Puerto Rico, Parador, Casa Grande [FSCA]; 371. Diapherodes jamaicensis (Drury, 1773) ♂: Jamaica, Portland [FSCA]; 372. Diapherodes laevicollis Redtenbacher, 1908, ♂: Jamaica, Saint Andrew [FSCA].


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FIGURES 373–374. Maps. 373. Map of the West Indies showing the principle islands; 374. Known distribution of Aploploides Rehn & Rehn, 1938 [green] and Venupherodes n. gen. [red].



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FIGURES 375–376. Maps. 375. Known distribution of Diapherodes Gray, 1835 and the two species-groups here defined: Gigantea-group [green] and Jamaicensis-group [red]; 376. Distribution of Haplopus Gray, 1835. Although there have not yet been definite records from the southern islands of the Bahamas Caicos and Turks Islands, Haplopus is most certainly also present on these islands. Hence, they are here included in the distribution range.


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FIGURES 377–378. Maps. 377. Known distribution of Cephaloplopus n. gen. [green] and Apteroplopus n. gen. [red]; 378. Known distribution of Parhaplopus n. gen. [green] and Paracranidium Brock, 1998 [red].



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FIGURES 379–380. Maps. 379. Known records of the species of Cephaloplopus n. gen. and Parhaplopus n. gen. on Hispaniola; 380. Known records of the species of Haplopus Gray, 1835 on Hispaniola.


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FIGURE 381. Maps. Distribution of the species of the gigantea-group of Diapherodes Gray, 1835 in the Lesser Antilles.



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FIGURES 382–384. Live insects. 382. Diapherodes achalus (Rehn, 1904) ♀: Puerto Rico, Maricao [Ó Mel J. Rívera]; 383. Diapherodes achalus (Rehn, 1904) ♂: Puerto Rico, Maricao [Ó Mel J. Rivera]; 384. Venupherodes venustula (Audinet-Serville, 1838) n. gen., n. comb. ♀: captive reared from NW-Cuba, Pinar del Rio.


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FIGURES 385–387. Live insects. 385. Venupherodes venustula (Audinet-Serville, 1838) n. gen., n. comb. ♂: captive reared from NW-Cuba, Pinar del Rio; 386. Venupherodes venustula (Audinet-Serville, 1838) n. gen., n. comb. ♀ and ♂: captive reared from NW-Cuba, Pinar del Rio; 387. Venupherodes venustula (Audinet-Serville, 1838) n. gen., n. comb. ♀♀ usual green colour-form and specimen with brown mottling: captive reared from NW-Cuba, Pinar del Rio.



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FIGURES 388–390. Live insects. 388. Diapherodes jamaicensis (Drury, 1773) ♂: captive reared from Jamaica; 389. Diapherodes jamaicensis (Drury, 1773) ♂: captive reared from Jamaica; 390. Diapherodes jamaicensis (Drury, 1773) ♀: captive reared from Jamaica.


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FIGURES 391–393. Live insects. 391. Diapherodes jamaicensis (Drury, 1773) ♀ green specimen with brown markings: captive reared from Jamaica [Ó Mieke Duytschaever]; 392. Diapherodes gigantea gigantea (Gmélin, 1789) ♀: captive reared from Grenada; 393. Diapherodes gigantea gigantea (Gmélin, 1789) ♀: captive reared from Grenada.



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FIGURES 394–395. Live insects. 384. Diapherodes martinicensis Lelong & Langlois, 2005 ♀: captive reared from Martinique; 385. Diapherodes martinicensis Lelong & Langlois, 2005 ♂: captive reared from Martinique.


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FIGURES 396–398. Live insects. 396. Diapherodes gigantea gigantea (Gmélin, 1789) ♂: captive reared from Grenada. 397. Parhaplopus evadne (Westwood, 1859) n. gen., n. comb. defensive reaction of ♂: Hispaniola, SW-Haiti, Macaya National Park [Ó Eladio Fernández]; 398. Parhaplopus evadne (Westwood, 1859) ♂: Haiti, Department du Sud, Formon [Ó Ruth Bastardo].



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FIGURES 399–401. Live insects. 399. Haplopus micropterus (St. Fargeau & Audinet-Serville, 1828) ♀: Dominican Republic, Pedernales Province, Boca de la Cañada 400. Haplopus micropterus (St. Fargeau & Audinet-Serville, 1828) ♀: Haplopus micropterus (St. Fargeau & Audinet-Serville, 1828) ♀: Dominican Republic, Santo Domingo Province, Sierra Prieta; 401. Haplopus micropterus (St. Fargeau & Audinet-Serville, 1828) ♂: Dominican Republic, Baní Peravia Province, Pueblo Nuevo.


HENNEMANN ET AL.

FIGURES 402–405. Live insects. 405. Haplopus micropterus (St. Fargeau & Audinet-Serville, 1828) ♀: Dominican Republic, San Cristóbal Prov. [Ó Carlos de Soto Molinari]; 403. Haplopus micropterus (St. Fargeau & Audinet-Serville, 1828) ♂: Dominican Republic, Santo Domingo Prov., ~5 km N of Pedregal; 404. Haplopus micropterus (St. Fargeau & Audinet-Serville, 1828) ♂: captive reared from unknown locality in the Dominican Republic; 405. Haplopus micropterus (St. Fargeau & Audinet-Serville, 1828) ♀: Dominican Republic, La Altagracia Prov., ~2 km N Bayahibe.



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FIGURES 406–408. Live insects. Gynandromorph specimen of Diapherodes gigantea gigantea (Gmélin, 1789) captive reared from Grenada [Ó Mieke Duytschaever]. 406. Habitus; 407. Closeup of head and thorax showing the rudimentary dextral wings; 408. Closeup of apex of abdomen (dorsal view).


HENNEMANN ET AL.

FIGURES 409. Possible relationships within the subfamily Cladomorphinae sensu lato. The tree shown above is merely meant to visualise the possible relationships here suggested. It does not restrict to defining apomorphies for each clade or taxon, but also uses key-features that are helpful for distinction. Hence, it should not be interpreted as a strict phylogenetic cladogramm.



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