Zootaxa, Synopsis of Australian agathidine wasps...

Zootaxa 2480: 1–26 (2010) www.mapress.com / zootaxa/

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Synopsis of Australian agathidine wasps (Hymenoptera: Braconidae: Agathidinae) NICHOLAS B. STEVENS1, ANDREW D. AUSTIN2 & JOHN T. JENNINGS3 Australian Centre for Evolutionary Biology & Biodiversity (ACEBB), School of Earth & Environmental Sciences, The University of Adelaide, Adelaide, South Australia, 5005. E–mail: [email protected]; [email protected]; [email protected]

Abstract Agathidinae is a large subfamily of braconid wasps that are koinobiont endoparasitoids of lepidopteran larvae. Although the group is relatively well studied in the northern hemisphere, the Australian fauna is poorly known, with only 36 species recorded from the continent. This study presents a synopsis of the genera and species that occur in Australia, including a key to genera, information on their distribution and apparent species richness, and a list of species according to the latest generic concepts. Ten genera occur in Australia: Amputostypos Sharkey, Baeognatha Kokujev, Biroia Szépligeti, Braunsia Kriechbaumer, Coccygidium Saussure, Cremnops Foerster, Disophrys Foerster, Euagathis Szépligeti, Lytopylus Foerster and Therophilus Wesmael, with Lytopylus known only from a single introduced species. Baeognatha stat. rev. is taken out of synonymy with Therophilus and two genera are synonymised: Platyagathis Turner with Disophrys (syn. nov.) and Camptothlipsis with Baeognatha (syn. nov.). The genera Agathis Latreille and Bassus Fabricius are excluded from the Australian fauna, as the species described under these genera are here transferred to Therophilus, and no other species of either genus have been discovered from the continent. Further, the genera Baeognatha and Coccygidium are recorded from Australia for the first time, although not represented by any described species. The following new combinations are proposed: Amputostypos dilutum (Turner), comb. nov. (from Hypsostypos), A. exornatum (Turner), comb. nov. (from Hypsostypos), Disophrys leaena (Turner), comb. nov. (from Platyagathis), Lytopylus rufipes (Nees von Esenbeck) comb. nov. (from Bassus), Therophilus antipodus (Ashmead) comb. nov. (from Orgiloneura), T. festinatus (Turner), comb. nov. (from Bassus), T. latibalteatus (Cameron) comb. nov. (from Agathis), T. leucogaster (Holmgren) comb. nov. (from Agathis), T. malignus (Turner). comb. nov. (from Bassus), T. martialis (Turner) comb. nov. (from Agathis), T. meridionalis (Turner comb. nov. (from Bassus), T. minimus (Turner) comb. nov. (from Bassus), T. minor (Szépligeti) comb. nov. (from Bassus), T. pedunculatus (Szépligeti) comb. nov. (from Bassus), T. ruficeps (Szépligeti). comb. nov. (from Bassus), T. rufithorax (Turner) comb. nov. (from Agathis), T. rufobrunneus (Turner) comb. nov. (from Agathis), T. rugosus (Turner) comb. nov. (from Bassus), T. tricolor (Szépligeti) comb nov. (from Bassus), and T. xanthopsis (Turner) comb. nov. (from Agathis). In addition, Bassus tenuissimus (Turner) is synonomised with T. ruficeps (Szépligeti) syn. nov., and Agathis dimidiata (Brullé) is designated a nomen dubium. Key words: Agathidinae, Agathis, Amputostypos, Baeognatha, Bassus, Biroia, Braunsia, Camptothlipsis, Coccygidium, Cremnops, Disophrys, Euagathis, Hypsostypos, Orgiloneura, Platyagathis, Therophilus, mimicry

Introduction The braconid subfamily Agathidinae is a relatively speciose group of koinobiont endoparasitoids of lepidopteran larvae, with over 1,200 described species in 50 genera worldwide (Sharkey 1997, Yu et al. 2005, Sharkey et al. 2006). Most agathidines are solitary parasitoids, commonly of larvae that feed within concealed habitats such as silken galleries, although members of some genera (e.g., Disophrys Foerster and Coccygidium Saussure) are known to parasitise exposed larvae (Shaw and Huddleston 1991, Sharkey 1992, Sarmiento and Sharkey 2004). The subfamily is well defined and easily recognised by several wing venation characters including the presence of a fore wing fold between the prestigma and the medial vein, a narrow

Accepted by A. Aguiar: 31 Mar. 2010; published: 21 May 2010

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marginal cell terminating at the anterior wing margin well before the wing apex, and fore wing M+Cu not tubular in the basal third or more (Sharkey 1997). A recent phylogenetic study (Sharkey et al. 2006) also demonstrated the monophyly of the Agathidinae relative to other braconid subfamilies. The agathidine fauna of Australia has not been studied in any detail since the work of Turner (1918 a, b) who described 25 of the 36 recognised species for the continent. However, the wealth of material in collections indicates that the species diversity of agathidines is much greater than this, and that the fauna is dominated by one genus, Therophilus Wesmael. Our aim in this study was to provide a synopsis of the Australian Agathidinae at the generic level employing contemporary generic concepts that have recently been adopted for other regions of the world (Simbolotti and Achterberg 1999, Sharkey et al. 2006, 2009), and to assess levels of species richness. We provide a key to genera for the region, and a corrected list of species based on examination of primary types, along with relevant references. In addition, we present an overview of the fauna with notes on the distribution of genera, their biology, and occurrence of likely mimicry colour patterns.

Material and methods Abbreviations. Abbreviations used for institutions for where specimens are deposited are as follows: ANIC: BMNH: HNHM: MNHN: NHRS: SDEI: USNM:

Australian National Insect Collection, CSIRO, Canberra, Australia Natural History Museum, London, England Hungarian Natural History Museum, Budapest, Hungary Museum National d'Histoire Naturelle, Paris, France. Naturhistoriska riksBrulléémuseet, Stockholm, Sweden. Senckenberg Deutsches Entomologisches Institut, Müncheberg, Germany. National Museum of Natural History, Smithsonian Institution, Washington DC, USA

Imaging. Images of specimens were captured using both a Phillips XL30 field emission scanning electron microscope (FESEM) (Adelaide Microscopy and Microanalysis Research Facility, The University of Adelaide) and a Nikon DL1200 digital camera mounted on a Leica 16Z microscope (maximum magnification 115x with 10x objective lens) in conjunction with the software package Auto–montage (Syncroscopy: http:// www.syncroscopy.com) that allows a greater depth of focus to be achieved. Prior to image capture, non–type specimens were cleaned using absolute ethanol applied with a fine brush to remove obvious dirt and debris. For scanning electron microscopy, specimens were examined uncoated at 1.0 kV and a spot size of two or three.

Overview of the Australian fauna Generic and species diversity. Following examination of all available, relevant primary types, and material in major collections, we recognise ten genera as comprising the Australian fauna. These are Amputostypos Sharkey, Baeognatha Kokujev, Biroia Szépligeti, Braunsia Kriechbaumer, Coccygidium Saussure, Cremnops Foerster, Disophrys Foerster, Euagathis Szépligeti, Lytopylus Foerster, and Therophilus Wesmael. Lytopylus does not occur naturally on the continent and is known only by a single introduced species. Agathis Latreille and Bassus Fabricius are deemed to be absent from the Australian fauna, given that Agathis dimidiata Brullé is declared a nomen dubium, and all other Australian species previously considered to belong to Agathis and Bassus are here transferred to Therophilus. Also, following examination of the relevant type, Platyagathis is proposed as a new junior synonym for Disophrys, and Camptothlipsis is proposed as a new junior synonym for Baeognatha. Two genera, Baeognatha and Coccygidium, are recorded from the continent for the first time, both represented by undescribed species only.

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STEVENS ET AL.

The Australian agathidine fauna is dominated by one genus, Therophilus, with over 150 species present (Table 1). Coccygidium is the next most speciose genus with an estimated 20 to 25 species for the continent. Cremnops and Disophrys are the only other genera to be well represented in Australia, with the remaining genera apparently comprising relatively minor components of the fauna. TABLE 1. Number of described species and estimated species richness and distribution of Australian agathidine genera. Genus

No. described No. estimated Distribution in Australian biogeographic regions spp. spp.

Amputostypos Sharkey

2

5

Torresian

Baeognatha Kokujev

0

~6

Timorian and Torresian

Biroia Szépligeti

1

4

Torresian and northern Kosciuskan

Braunsia Kriechbaumer

2

>5


Coccygidium Saussure

0

>20

Timorian and Torresian mostly; rarely Eyrean and Southwestern

Cremnops Foerster

4

>10


Disophrys Foerster

7

>15

Timorian and Torresian; rarely Southwestern

Euagathis Szépligeti

1

~5


Lyptopylus Foerster

1

1

Introduced; southern Kosciuskan

Therophilus Wesmael

17

>150

Widespread throughout continent

The Bassus problem. Sharkey et al. (2006) undertook a preliminary phylogenetic study using both morphology and sequence data from the 28S rDNA gene to explore agathidine relationships. One of the major findings was that Bassus (as it was then circumscribed) was polyphyletic and, even though only 10 exemplar species were included for the genus, they fell out in four separate lineages. Based on these results, Sharkey et al. (2009) began a process of dividing Bassus s.l. into a number of smaller genera, and reinstated the names Camptothlipis, Lytopylus and Therophilus. Therophilus is poorly defined and is difficult to identify reliably, a situation that existed for Bassus s.l. The changes made to the morphological definition of Bassus by Sharkey et al. (2009) mean that no Bassus s.s. species were represented in the phylogenetic analysis of Sharkey et al. (2006). Therefore, it is unknown where Bassus s.s. (based on the type species B. calculator Fabricius) fits within the evolution of the Agathidinae. Sharkey et al. (2009) concede that Therophilus is still polyphyletic, so many species end up being transferred from one polyphyletic genus (Bassus) to another (Therophilus), and so further changes will undoubtedly need to be made in the future. In regard to the Australian fauna, Therophilus is by far the most speciose genus in Australia and comprises about two–thirds of all agathidine species on the continent. Only a single Australian Therophilus species (as Bassus sp. 4) was included in Sharkey et al. (2006) so the nature of the evolution of the group in the region is not known. Given the morphological diversity evident among the large number of Australian Therophilus species examined, it is likely that they represent several independent lineages, possibly some separate to those identified by Sharkey et al. (2006). Distribution of Genera. The northern regions of Australia (viz. Timorian, Torresian and northern Kosciuskan) contain the greatest generic diversity with all agathidine genera represented (Fig. 1, Table 1). Therophilus is the only widespread genus that occurs throughout all regions on the mainland and Tasmania in a range of environments from tropical and subtropical, to temperate, arid and semi–arid. Species of all other genera are mostly found only in the northern regions of Australia that are largely characterised by tropical to subtropical environmental conditions. There are a few exceptions to this pattern such as an undescribed species of Coccygidium recorded from the Great Victorian Desert in the arid Eyrean region of central Australia, and a second undescribed species from the temperate Southwestern region (Fig. 1). Disophrys leaena (Turner), comb. nov., represents the most southerly record for the genus, which is also recorded from the temperate Southwestern region. Australian agathidine genera

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FIGURE 1. Australian biogeographic regions. Nomenclature adapted from Heatwole (1987).

Biology. The Agathidinae are mostly solitary koinobiont endoparasitoids of Lepidoptera larvae, although Sarmiento and Sharkey (2004) recently recorded a gregarious species. Most commonly, concealed larvae (e.g., within leaf curls, seeds, silken galleries, mines or retreats) are attacked (Shaw and Huddleston 1991). However, members of the tribe Disophrini, having short ovipositors, are the exception and parasitise exposed larvae (Sharkey 1992). A limited number of biological studies indicate a range of oviposition strategies occur within the group. For instance the eggs of the Nearctic species, Agathis gibbosa (Say) and A. unicolor (Schrottky) are inserted into the ganglion of their host’s ventral nerve cord (Odebiyi and Oatman 1972, 1977). The placement of eggs into the nerve fibre is postulated to circumvent the host’s immune system, thereby preventing encapsulation by host haemocytes and death of the parasitoid egg (Godfray 1994, Quicke 1997). This is known to occur in several parasitoid taxa (Flanders 1973) but is not considered widespread within the Braconidae (Shaw and Quicke 2000). Dondale (1954) and Quednau (1970) observed in the Holarctic species Bassus dimidiator (Nees von Esenbeck) and Agathis pumila (Ratzeburg), respectively, that eggs were not inserted into the nerve fibre, but attached to the outside wall of the host nerve tissue instead. In contrast to the ganglion associated oviposition strategy, Simmonds (1947) reported that the Nearctic species Cremnops vulgaris (Cresson) (as Bracon) deposits its egg directly into the host’s haemocoel. Regardless of where the egg is initially deposited, the resultant wasp larva develops within the host’s body cavity until the third instar emerges from the final host larval stage, often after the penultimate host instar has spun its cocoon (Bhat and Gupta 1977, Shaw and Huddleston 1991). The parasitoid may then feed externally, devouring the host caterpillar prior to pupating. The final wasp instar larva usually spins its cocoon within the cocoon of the deceased host or on the host remains. However, some species construct their cocoon on leaves a little distance away from the remains of the host and/or cocoon (Bhat and Gupta 1977, Shaw and Huddleston 1991, NBS pers. obs.).

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In general, agathidines do not appear to be very host specific. A wide range of different lepidopteran host groups (over 25 families spanning 12 superfamilies) are attacked, with numerous agathidine species known to parasitise 10 or more host species from five or more families each (Thompson 1953, Bhat and Gupta 1977, Nixon 1986, Sharkey 1996). One Palaearctic species, Lytopylus rufipes (as Bassus) (Nees von Esenbeck), which has been introduced into Australia, has been recorded from 19 Palaearctic host species representing six families (Sharkey 1996). However, other species show host associations which are specific enough to allow for deployment outside their native range as biological control agents (Sharkey 1997). In Australia, the hosts of very few species have been recorded, and those that have been documented are associated with native lepidopteran pests. Undescribed Therophilus species are known to parasitise the native legume pest, Etiella behrii Zeller (Pyralidae) (Austin et al. 1993), and the significant polyphagous pest Epiphyas postvittana (Walker) (Tortricidae) (Paull and Austin 2006), which feeds on a large range of native and horticultural species. Both pest species have been accidentally introduced into other countries, E. behrii into Indonesia (Wakamura et al. 1999) and E. postvittana into New Zealand, New Caledonia, Hawaii, the western USA and the United Kingdom (Bradley 1956, Geier and Briese 1980, Danthanarayana 1983, Varela et al. 2008), thus raising the prospect of employing Therophilus spp. as potential control agents. Mimicry colour patterns. An interesting aspect of the Australian agathidine fauna is the presence of two major colour forms, each believed to be aposematic and part of different mimicry complexes. One form comprises contrasting yellow–brown and black colour that exhibits a variety of patterns on the body and wings of species in the mostly tropical genera Braunsia, Coccygidium, Cremnops and Disophrys (Figs 4B, C, 8B, C). Central and South American species in the genera Alabagrus Sharkey and Sesioctonus Viereck, also display similar colour patterns and are believed to represent a large putative mimicry complex containing over 2,000 ichneumonoid species, as well as an unknown number of species from other pterygote orders, including several hundred reduviid species (Briceńo 2003, Leathers and Sharkey 2003). The second major pattern comprises distinctive and contrasting black, red–orange and white, and is referred to herein as the BROW colour pattern (Figs 7B, C, 9B). An earlier depiction of members of this putative mimicry complex are illustrated in Plates 3, 5 and 6 of Insects of Australia (Naumann 1991). The BROW pattern is widespread across numerous Australian braconid subfamilies (e.g., Helconinae, Braconinae and Doryctinae) (Quicke et al. 1992, Belokobylskij et al. 2004, Iqbal et al. 2006) and has also been observed in numerous lepidopteran, dipteran, reduviid and mirid species (Naumann 1991, authors pers. obs.). Within the Australian Agathidinae, the BROW pattern is confined mostly to Therophilus species, although it is also known in two species of Disophrys (see Comments under Disophrys). At this stage it is not clear whether agathidine members of both mimicry complexes represent Mullerian or Batesian mimics, or a combination of both. Leathers and Sharkey (2003) proposed that species of Alabagrus in the tropical South American mimicry complex were Batesian mimics, on the basis that an offensive odour detectable to humans is not exuded nor is a painful sting known, unlike other braconid and reduviid members of the complex. Numerous braconines have been observed to deliver a painful sting and exude pungent odours when handled that could act as deterrents to predators. On this basis, Quicke et al. (1992) hypothesised that the BROW pattern acted as an aposematic signal and that most braconines in the complex were Mullerian mimics. It is not known whether Cremnops, Coccygidium, Disophrys or Therophilus species sting potential predators or are capable of releasing pungent odours. It is interesting to note that if braconid members of both putative complexes were indeed Mullerian mimics, solely on the basis that the female wasp could use her ovipositor defensively, then male wasps would be considered Batesian mimics, thereby weakening the signal established by the female model.

Key to Australian genera of Agathidinae 1 Fore tarsal claws cleft (Figs 2A and C); preapical spines not present on anterior surface of mid tibia .................... 2 Fore tarsal claws not cleft (Fig. 2B); preapical spines present on anterior surface of mid tibia (Fig. 2D)................ 7 2 (1) Base of fore tarsal claw with spines forming comb (= pectinate) (Fig. 2C); ovipositor sheaths longer than 0.5 length of metasoma (Fig. 9C); notauli always present (e.g., Fig. 3B) ...................................................... Cremnops Foerster

Australian agathidine genera


5

-

Base of fore tarsal claw without spines forming comb (= not pectinate) (Fig. 2A); ovipositor sheaths often shorter than 0.5 length of metasoma (Fig. 9D); if ovipositor sheaths longer than 0.5 length of metasoma then notauli absent (Fig. 3A) ..................................................................................................................................................................... 3 3 (2) Frons lacking lateral carinae ...................................................................................................................................... 4 Frons with lateral carinae present (Fig. 3B)............................................................................................................... 5 4 (3) Hind trochantellus lacking pair of longitudinal carinae on ventral surface ................................ Euagathis Szépligeti Hind trochantellus with pair of longitudinal carinae on ventral surface (e.g., Fig. 2G) .......... Amputostypos Sharkey 5 (3) Fore wing cell 1–RS triangular, posterior margin longer than anterior margin (Fig. 4A); fore wing vein 2–RS2 absent (Fig. 4A); fore tibial spur long, equal to or greater than 0.8 basitarsus length, aristate and sinuate (Fig. 2E) .. .................................................................................................................................................. Coccygidium Saussure Fore wing cell 1–RS quadrate, both posterior and anterior margins of similar length (Fig. 4B); fore wing vein 2– RS2 often present (Fig. 4B); fore tibial spur length less than 0.7 basitarsus length, not aristate or sinuate (Fig. 2F) .. ..................................................................................................................................................................................... 6 6 (5) Fore wing vein 2–RS2 absent; notauli absent (e.g., Fig. 3A); ovipositor sheaths longer than 0.5 length of metasoma (e.g., Fig. 9C) .................................................................................................................................... Biroia Szépligeti Fore wing vein 2–RS2 present (Fig. 4B); notauli present (Fig. 3B); ovipositor sheaths less than 0.5 length of metasoma (e.g., Fig. 9D) ....................................................................................................................... Disophrys Foerster 7 (1) Fore wing cell 1–RS absent (Fig. 4C) ....................................................................................... Baeognatha Kokujev Fore wing cell 1–RS present (e.g., Fig. 4D) ............................................................................................................. 8 8 (7) Fore wing vein 2–RS2 present (e.g., Fig. 4B), may be reduced to short stub basally .......... Braunsia Kriechbaumer Fore wing vein 2–RS2 absent (Fig. 4D) .................................................................................................................... 9 9 (8) Hind tarsal claw lacking basal lobe............................................................................................................... Bassus F. Hind tarsal claw with basal lobe present (Fig. 2B) ................................................................................................. 10 10 (9) T2+3 entirely covered with prominent uniform striate sculpturing (Fig. 3C); fused plate of T2+3 dominates dorsal surface of metasoma (Fig. 3C); metasomal foramen ventral margin ends dorsal to dorsal margin of hind coxal cavities (Fig. 5A); prominent and straight transverse carinae present between metasomal foramen and hind coxal cavities (Fig. 5A) ................................................................................................................................... Lytopylus Foerster T2+3 entirely smooth, or if sculpturing present then faint, often non–striate, and mostly restricted to T2 only; fused plate of T2+3 does not dominate dorsal surface of metasoma (Fig. 3D); metasomal foramen ventral margin extends below level of dorsal margin of hind coxal cavities (Fig. 5B); prominent and straight transverse carinae not present between metasomal foramen and hind coxal cavities (Fig. 5B) ............................................... Therophilus Wesmael

Amputostypos Sharkey, 2009 (Figs 2F, 10D) Amputostypos Sharkey, 2009: 30. Type species: Disophrys concolor Szépligeti, 1908: 229, by original designation.

Diagnosis. Length: 4–7 mm; colour: commonly yellow and dark brown (Fig. 10D), sometimes white on anterior ventral metasomal sclerites; frons never with lateral carinae; notauli present and scrobiculate (e.g., Fig. 3B); tarsal claws cleft, but base not pectinate (e.g., Fig. 2A); mid tibial preapical spines absent; fore tibial spur not elongate, sinuate or aristate, length less than 0.75 length of basitarsus (Fig. 2F); pair of longitudinal carinae present on ventral surface of hind trochantellus (e.g., Fig. 2G); fore wing cell 1–RS present and sessile, vein 2–RS2 absent; ovipositor short, length less than 0.5 metasoma length (Fig. 10D). Comments. Sharkey et al. (2006) reinstated Hypsostypos as a valid taxon, redefined its limits, and showed the genus to be closely related to Coccygidium and Euagathis. Sharkey et al. (2009) erected Amputostypos to include species previously belonging to Hypsostypos but which lacked high ridges surrounding the antennal insertions. Members of Amputostypos can be distinguished from Coccygidium by the lack of lateral carinae on the frons and the fore tibial spur being considerably shorter than the basitarsal segment, and from Euagathis by the presence of longitudinal carinae on the hind trochantellus. Species richness and distribution. Amputostypos is not well represented in Australia, with only five species identified in collections, two of which are described. They appear to be restricted to the Torresian and northern Kosciuskan regions.

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FIGURE 2. Agathidinae leg characters: A, Biroia sp., fore tarsal claw cleft (arrowed), base non–pectinate; B, Braunsia sp., fore tarsal claw simple, non–cleft, basal lobe quadrate, non–pectinate; C, Cremnops sp., fore tarsal claw cleft, base pectinate (arrowed); D, Braunsia sp., mid tibial preapical spines (circled); E, Coccygidium sp., fore tibial spur long, sinuate and aristate (arrowed); F, Amputostypos sp., fore tibial spur short, non–sinuate and non–aristate (arrowed); G, Disophrys sp., posterior view of metasoma, showing paired longitudinal carinae on ventral surface of hind trochantellus (arrowed). Scale lines, A, B and C = 0.05 mm; D, E and F = 0.1 mm; G = 1 mm.

List of Australian species Amputostypos dilutum (Turner, 1918b), comb. nov. Disophrys diluta Turner, 1918b: 228. Holotype BMNH ♀; Type locality: Mackay, Queensland; Parrot, 1953: 200 [catalogue]; Shenefelt, 1970b: 393 [catalogue]; Yu et al., 2005 [catalogue]. Hypsostypos dilutum (Turner); Sharkey et al., 2006: 561 [generic transfer].



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FIGURE 3. Agathidinae, dorsal mesoscutum and head: A, Therophilus sp., notauli absent; B, Disophrys sp., notauli present and scrobiculate (small arrow), lateral carinae of frons present (large arrow). Agathidinae lateral metasoma: C, Lytopylus rufipes; D, Therophilus sp. Scale lines = 0.5 mm.

Amputostypos exornatum (Turner, 1918b), comb. nov. Disophrys exornata Turner, 1918b: 227–28. Holotype BMNH ♀; Type locality: Kuranda, Queensland. Parrot, 1953: 200 [catalogue]; Shenefelt, 1970b: 394 [catalogue]; Yu et al., 2005 [catalogue]. Sharkey et al., 2006: 561 [generic transfer]. Hypsostypos exornatum (Turner); Sharkey et al., 2006: 561 [generic transfer].

Baeognatha Kokujev, 1903, stat. rev. (Figs 4C, 6A, 8A) Baeognatha Kokujev, 1903: 243; Type species: Baeognatha turanica Kokujev, 1903, by monotypy; Shenefelt, 1970b: 367 [catalogue]; Bhat & Gupta, 1977: 86 [key, descriptions]; Nixon, 1986: 229 [key, descriptions]; Sharkey, 1992: 441 [in tribe Microdini]; Yu et al., 2005 [catalogue]; Sharkey et al., 2009: 47 [as synonym of Therophilus]. Camptothlipsis Enderlein, 1920: 166; Type species: Camptothlipsis costalis Enderlein, 1920, by original designation; Shenefelt, 1970b: 378 [catalogue]; Bhat & Gupta, 1977: 78 [key, descriptions]; Tobias, 1976: 214 [synonym of Baeognatha Kokujev, 1903]; Nixon, 1986: 229 [synonym of Baeognatha]; Simbolotti & Achterberg, 1992: 6 [synonym of Bassus]; Sharkey et al. 2006: 556 [synonym of Bassus]; Sharkey et al., 2009 [reinstated]. syn. nov.

Diagnosis. Length: 3–5 mm; colour: commonly mostly yellow with variable black or dark brown markings (Figs 8A); lateral carinae on frons always absent; notauli present and distinctly scrobiculate (e.g., Fig. 3B); claws non–cleft with rounded basal lobes present (e.g., Fig. 2C); mid tibial preapical spines always present (e.g., Fig. 2D); fore wing cell 1-RS absent in Australian species (Fig. 4C), however, does vary from absent to small cell present in type species (Simbolotti and Achterberg 1992); ovipositor length greater than 0.5 metasoma length (e.g., Fig. 8B). Comments. Historically, species characterised by the absence of fore wing cell 1–RS were classified as either Camptothlipsis or Baeognatha. Camptothlipsis was treated as a synonym of Baeognatha (e.g., Tobias

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(1976) and Nixon (1986)) prior to Camptothlipsis being synonymised with Bassus by Simbolotti and Achterberg (1992). In Sharkey et al. (2006), two undescribed species (treated as Bassus), characterised by the absence of fore wing cell 1–RS and granulate sculpturing on T1, were resolved as sister taxa to one another. This result prompted Sharkey et al. (2009) to reinstate the concept of Camptothlipsis as defined by Bhat and Gupta (1977) who used scupturing of the propodeum and anterior metasomal tergites (viz. granulate versus non–granulate, usually striate) to differentiate Camptothlipsis from Baeognatha. In addition, Sharkey et al. (2009) synonymised Baeognatha with Therophilus. However, examination of Australian material, suggests that the granulate sculpturing patterns of the propodeum and anterior metasomal tergites are not reliable generic level characters because they also occur in a number of Therophilus species. Therefore, Baeognatha as interpreted by Nixon (1986), is reinstated. Species richness and distribution. No Australian species of Baeognatha have been described. The genus represents a relatively small component of the Australian fauna, comprising approximately six species that occur in the Timorian and Torresian regions of the continent.

FIGURE 4. Agathidinae fore wing venation characters: A, Coccygidium sp., fore wing cell 1–RS triangular, vein 2–RS2 absent; B, Disophrys sp., fore wing cell 1–RS quadrate, vein 2–RS2 present; C, Baeognatha sp., fore wing cell 1–RS absent; D, Therophilus unimaculatus, fore wing cell 1–RS small and petiolate, vein 2–RS2 absent. Scale lines A, B and D = 1 mm; C = 0.5 mm.

Biroia Szépligeti, 1900 (Figs 2A, 6B, 9A) Biroia Szépligeti, 1900: 62. Type species: Biroia elegans Szépligeti, 1900, by monotypy; Turner, 1918b: 229 [description]; Parrott, 1953: 199 [catalogue]; Shenefelt, 1970b: 368 [catalogue]; Bhat & Gupta, 1977: 265 [description, catalogue]; Sharkey, 1992: 437 [in tribe Cremnoptini]; Yu et al., 2005 [catalogue]; Sharkey et al., 2006: 557 [phylogeny, in tribe Cremnoptini]; Sharkey et al., 2009: 33 [key, description]. Isoptronotum Enderlein, 1920: 193. Type species Isoptronotum taeiocauda Enderlein, 1920, by original designation; Shenefelt, 1970b: 418 [catalogue]; Bhat & Gupta, 1977: 159 [key, description]; Sharkey et al., 2006: 557 [synonym of Biroia].

Diagnosis. Length: 7–10 mm; colour: body varying from red–orange to yellow (Fig. 9A); wings infuscate; inter–antennal region with paired carinae or protuberances (Fig. 6B); frons with incomplete lateral carinae; notauli always absent (e.g., Fig. 3A); claws cleft, base of claws not pectinate (Fig. 2A); mid tibial preapical spines absent; pair of longitudinal carinae not present on ventral surface of hind trochantellus; fore wing cell



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1–RS present, quadrate and sessile (e.g., Fig. 4B), vein 2–RS2 absent; ovipositor long, greater than 0.5 metasomal length (e.g., Fig. 9C). Comments. Biroia is closely related to Cremnops (Sharkey et al. 2006) from which it can be distinguished by the basal claws being non-pectinate and notauli absent. It is not well represented in Australia and is more diverse in the Oriental region (Bhat and Gupta 1977), with species occurring in Papua New Guinea to which the Australian representatives appear to be most closely allied. Species richness and distribution. Only four species are known of which one is described. All are from the northeast part of the continent in the Torresian and northern Kosciuskan biogeographic regions.

FIGURE 5. Agathidinae posterior mesosoma: A, Lytopylus rufipes, prominent and straight transverse carina (arrowed) present between metasomal foramen and hind coxal cavities; B, Therophilus sp., ventral margin of metasomal foramen ventral to hind coxal cavity dorsal margin. Scale lines = 0.1 mm.

FIGURE 6. Agathidinae anterior head view: A, Baeognatha sp.; B, Biroia sp., paired carinae present on inter–antennal region. Scale lines = 0.5 mm.

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List of Australian species Biroia solitaria Turner, 1918b Biroia solitaria Turner, 1918b: 229. Holotype BMNH ♀ ; Type locality: Mackay, Queensland. Parrot, 1953: 199 [catalogue]; Shenefelt, 1970b: 369 [catalogue]; Yu et al., 2005 [catalogue].

FIGURE 7. Agathidinae anterior and lateral views of elongate heads: A, Cremnops commutator Turner; B, Disophrys sp., paired carinae present on inter–antennal region. Scale lines = 0.5 mm.

Braunsia Kriechbaumer, 1894 (Figs 2B, 9B) Braunsia Kriechbaumer, 1894: Type species: Braunsia bicolor Kriechbaumer, 1894 [= Braunsia bilunata Enderlein, 1904, new name for bicolor Kriechbaumer because junior secondary homonym of Brullé, 1846], by subsequent designation, Viereck, 1914: 24; Turner, 1918b: 224 [description]; Parrott, 1953: 199 [catalogue]; Shenefelt, 1970b: 370 [catalogue]; Bhat & Gupta, 1977: 50 [key, description]; Chou & Sharkey, 1989: 175 [key, description]; Sharkey, 1992: 443 [in tribe Microdini]; Sharkey, 1996: 58 [in tribe Eumicrodini, amendment to incorrectly applied Microdini]; Yu et al., 2005 [catalogue]; Sharkey et al., 2006: 557 [notes, phylogeny, in tribe Agathidini s.l.]; Sharkey et al., 2009: 34 [key, description]. Metriosoma Szépligeti, 1902: 74. Type species: Metriosoma munda Szépligeti, 1902, by monotypy; Shenefelt, 1970b: 313 [synonym of Agathis]; Sharkey et al., 2006: 557 [synonym of Braunsia]. Lissagathis Cameron, 1911: 245. Type species: Lissagathis bicarinata Cameron, 1911, by monotypy; Shenefelt, 1970b: 312 [synonym of Agathis]; Sharkey et al., 2006: 557 [synonym of Braunsia]. Laccagathis Watanabe, 1934: 121. Type species: Laccagathis formosana Watanabe, 1934, by original designation; Shenefelt, 1970b: 419 [catalogue]; Bhat & Gupta, 1977 [key, description]: 47; Chou & Sharkey, 1989: 187 [description]; Sharkey, 1996: 62 [description]; Sharkey et al., 2006: 557 [synonym of Braunsia]. Pholeocephala Achterberg, 1988: 48. Type species: Pholeocephala lieftincki Achterberg, 1988, by monotypy; Sharkey et al., 2006: 557 [synonym of Braunsia].

Diagnosis. Length: 8–14 mm; colour: body variable with different combinations of yellow or orange in conjunction with light or dark brown and black; wings with dark and light shaded pattern (Fig. 9B); interAustralian agathidine genera


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antennal region broad, often largely flat, may possess medial elevated mound (e.g., Fig. 6A); lateral carinae on frons absent; notauli range from totally absent to present, deep and heavily scrobiculate (e.g., Figs 3A and B); claws simple with basal lobe, are non-pectinate (Fig. 2B); mid tibial preapical spines present (Fig. 2D); fore wing cell 1–RS mostly quadrate; vein 2–RS2 always present (e.g., Fig. 4B), may be reduced to basal stub only; ovipositor long, always greater than 0.5 metasoma length (Fig. 9B). Comments. The Australian fauna contains species that would have previously been placed in Laccagathis, Metriosoma and Pholeocephala prior to these genera being synonymised with Braunsia by Sharkey et al. (2006). The molecular data indicate that Braunsia is closely related to Lytopylus (referred to as the Bassus macadamia group or incorrectly as Bassus s.str. in (Sharkey et al. 2006). Braunsia species can be distinguished from Lytopylus by the presence of fore wing vein 2–RS2. Species richness and distribution. Braunsia is not well represented in Australia with only 5–10 species present, two of which are described. All occur in northern Australia in the Timorian and Torresian biogeographic regions.

FIGURE 8. Lateral habitus of Agathidinae spp.: A, Baeognatha sp.; B, Therophilus unimaculatus, exhibiting BROW (see text) colour pattern; C, Therophilus nr latibalteatus, exhibiting BROW (see text) colour pattern; D, Therophilus sp. with dense propodeal setal field. Scale lines = 1 mm.

List of Australian species Braunsia bicolor (Brullé, 1846) Agathis bicolor Brullé, 1846: 483. Holotype unknown, ♀; Type locality: Australia. Braunsia bicolor (Brullé); Parrot, 1953: 199 [catalogue, generic transfer]; Shenefelt, 1970b: 370 [catalogue]; Yu et al., 2005 [catalogue].

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FIGURE 9. Lateral habitus of Agathidinae spp.: A, Biroia sp.; B, Braunsia sp., exhibiting tropical mimicry complex colour pattern; C, Cremnops sp., exhibiting tropical mimicry complex colour pattern; D, Coccygidium sp. Scale lines = 1 mm.

Braunsia diversipennis Turner, 1918b Braunsia diversipennis Turner, 1918b: 224 [examined]. Holotype BMNH ♂; Type locality: Mackay, Queensland. Parrot, 1953: 199 [catalogue]; Shenefelt, 1970b: 371 [catalogue]; Yu et al., 2005 [catalogue].

Coccygidium Saussure, 1892 (Figs 2E, 4A, 9D) Coccygidium Saussure, 1892: 15. Type species: Coccygidium luteum, Saussure, 1892, by monotypy; Shenefelt, 1970b: 392 [synonym of Disophrys]; Chou & Sharkey, 1989: 178 [key, description]; Sharkey, 1992: 441 [in tribe Disophrini]; Sharkey, 1996: 16 [key, description]; Yu et al., 2005 [catalogue]; Sarmiento and Sharkey, 2005: 66 [type descriptions]; Sharkey et al., 2006: 557 [notes, phylogeny, in tribe Disophrini]; Sharkey et al., 2009: 36 [key, description]. Brachyrhopalum Kriechbaumer, 1894: 312. Type species: Brachyrhopalum pallidum Kriechbaumer, 1894, by monotypy; Shenefelt, 1970b: 392 [synonym of Disophrys]; Chou & Sharkey, 1989: 178 [synonym of Coccygidium]. Neophylax Ashmead, 1900: 119. Type species: Neophylax snyderi Ashmead, 1900, by monotypy; Shenefelt, 1970b: 425 [synonym of Zelomorpha]; Chou & Sharkey, 1989: 178 [synonym of Coccygidium]. Ahngeria Kokujev, 1902: 6. Type species: Ahngeria transcaspica Kokujev, 1902, by monotypy; Shenefelt, 1970b: 366 [catalogue]; Sharkey, 1992: 441 [synonym of Coccygidium]. Dichelosus Szépligeti, 1902: 71. Type species: Dichelosus fuscipennis Szépligeti, 1902, by monotypy; Shenefelt, 1970b: 391 [catalogue]; Sarmiento & Sharkey, 2005: 66 [synonym of Coccygidium]. Lisitheria Cameron, 1904: 306. Type species: Lisitheria nigricornis Cameron, 1904, by monotypy; Shenefelt, 1970b: 425 [synonym of Zelomorpha]; Chou & Sharkey, 1989: 178 [synonym of Coccygidium]. Spilomicrodus Cameron, 1904: 323. Type species: Spilomicrodus nigriceps Cameron, 1904, by monotypy; Shenefelt, 1970b: 425 [synonym of Zelomorpha]; Chou & Sharkey, 1989: 178 [synonym of Coccygidium]. Xanthomicrodus Cameron, 1904: 157. Type species: Xanthomicrodus iridipennis Cameron, 1904, by monotypy; Shenefelt, 1970b: 425 [synonym of Zelomorpha]; Chou & Sharkey, 1989: 178 [synonym of Coccygidium].



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Caenophylax Schulz, 1911: 88. Type species: Neophylax snyderi Ashmead, 1900, replacement name for Neophylax Ashmead; Shenefelt, 1970b: 425 [synonym of Zelomorpha]; Chou & Sharkey, 1989: 178 [synonym of Coccygidium].

Diagnosis. Length: 5–7 mm; colour: commonly entirely yellow with dark brown antennae and wing patterning, more rarely with body yellow-brown and black colour pattern; inter-antennal region with medial elevated mound that may be rounded (e.g., Fig. 6A) or carinate, or with paired carinae or protuberances present (e.g., 7B); frons with lateral carinae (e.g., Fig. 3B); notauli present, complete and scrobiculate (e.g., Fig. 3B); tarsal claws cleft, but base not pectinate (e.g., Fig. 2A); mid tibial preapical spines absent; fore tibial spur long, exceeding 0.8 length of basitarsus, sinuate and aristate (Fig. 2E); pair of longitudinal carinae present on ventral surface of hind trochantellus (e.g., Fig. 2G); fore wing cell 1–RS always present, sessile and triangular (Fig. 4A), vein 2–RS2 absent (Fig. 4A); ovipositor short, length much less than 0.5 metasoma length, curved ventrally (Fig. 9D). Comments. Coccygidium has not been recorded from Australia previously, so there are no described species from the region. The genus is closely related to Euagathis and Hypsostypos (Sharkey et al. 2006) but can be distinguished from them by the presence of an elongate fore tibial spur that is nearly as long as or longer than the basitarsal segment and is characteristically sinuate and aristate. Generally, Coccygidium species are mostly yellow and relatively small. Only a few species from northern Australia display the yellowbrown and black colour pattern which is thought to be part of a mimicry complex (see above). Species richness and distribution. The genus is well represented in Australia with more than 20 species present. Most occur in the Timorian, Torresian and northern Kosciuskan regions of the continent. However, several species exist in the more temperate Southwestern and arid Eyrean regions (see under Generic Distribution above).

Cremnops Foerster, 1862 (Figs 2C, 7A, 9C) Cremnops Foerster, 1862: 246. Type species: Agathis deflagrator Nees von Esenbeck, 1834 (= Ichneumon desertor Linnaeus, 1758), by monotypy; Turner, 1918b: 221 [key, description]; Shenefelt, 1970b: 382 [catalogue]; Bhat & Gupta, 1977: 266 [description, catalogue]; Nixon, 1986: 191 [key, description]; Chou & Sharkey, 1989: 182 [key, description]; Sharkey, 1992: 439 [in tribe Cremnoptini]; Sharkey, 1996: 13 [key, description]; Yu et al., 2005 [catalogue]; Sharkey et al. 2006: 558 [notes, phylogeny, in tribe Cremnoptini]; Sharkey et al., 2009: 38 [key, description].

Diagnosis. Length: 5–10 mm; colour: yellow and dark brown or black; head often elongate (Fig. 7A); inter– antennal region with medial elevated mound that may be broadly rounded (e.g., Fig. 6A) to carinate, paired carinae or protuberances not present; frons sometimes with lateral carinae but usually without; notauli present, complete and scrobiculate (e.g., Fig. 3B); tarsal claws cleft, base pectinate (Fig. 2C); mid tibial preapical spines absent; fore tibial spur not elongate (e.g., Fig. 2F); pair of longitudinal carinae not present on ventral surface of hind trochantellus; fore wing cell 1–RS always present and sessile, vein 2–RS2 either absent or basal sector present; ovipositor long, length exceeding 0.5 metasoma length (Fig. 9C). Comments. Cremnops is closely related to Biroia (Sharkey et al. 2006) but can be distinguished from this genus, indeed all agathidine genera, by the presence of comb-like (pectinate) spines on the base of the fore and mid tarsal claws. All Australian species display variations in the yellow and dark brown colour pattern thought to be associated with a putative mimicry complex. Species richness and distribution. Cremnops is well represented in Australia with over 20 species, only four of which are described. The genus appears to be restricted to the northern parts of the continent in the Timorian, Torresian and northern Kosciuskan biogeographic regions.

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List of Australian species Cremnops commutator Turner, 1918b Cremnops commutator Turner, 1918b: 223. Holotype BMNH ♀; Type locality: Mackay, Queensland. Parrot, 1953: 199 [catalogue]; Shenefelt, 1970b: 383 [catalogue]; Yu et al., 2005 [catalogue].

Cremnops dissimilis Turner, 1918b Cremnops dissimilis Turner, 1918b: 222. Holotype BMNH ♀; Type locality: Mackay, Queensland. Parrot, 1953: 199 [catalogue]; Shenefelt, 1970b: 383 [catalogue]; Yu et al., 2005 [catalogue].

Cremnops marginipennis Turner, 1918b Cremnops marginipennis Turner, 1918b: 222. Holotype BMNH ♀; Type locality: Kuranda, Queensland. Parrot, 1953: 199–200 [catalogue]; Shenefelt, 1970b: 384 [catalogue]; Yu et al., 2005 [catalogue].

Cremnops xanthostigma Szépligeti, 1900 Cremnops xanthostigma Szépligeti, 1900: 61. Holotype NHM ♀; Type locality: Papua New Guinea. Turner, 1918b: 221 & 222 [key, notes]; Parrot, 1953: 200 [catalogue]; Shenefelt, 1970b: 384 [catalogue]; Yu et al., 2005 [catalogue].

Disophrys Foerster, 1862 (Figs 3B, 4B, 7B, 10A & B) Disophrys Foerster, 1862: 246. Type species: Agathis caesa Klug, by ICZN designation 1989: 76; Turner, 1918b: 225 [key, description]; Parrott, 1953: 200 [catalogue]; Shenefelt, 1970b: 391 [catalogue]; Bhat & Gupta, 1977: 267 [notes, catalogue]; Nixon, 1986: 190 [key, descriptions]; Chou & Sharkey, 1989 [key, description]: 183; Sharkey, 1992: 441 [in tribe Disophrini]; Yu et al., 2005 [catalogue]; Sharkey et al., 2006: 558 [notes, phylogeny, in tribe Disophrini]; Sharkey et al., 2009: 40 [key, description]. Pseudagathis Kriechbaumer, 1894: 65. Type species: Pseudagathis calabarica Kriechbaumer, 1894, by monotypy; Shenefelt, 1970b: 392 [synonym of Disophrys]; Chou & Sharkey, 1989: 183 [synonym of Disophrys]. Pseudocremnops Szépligeti, 1915: 152. Type species: Pseudocremnops atripennis Szépligeti, 1915, by monotypy; Shenefelt, 1970b: 423 [catalogue]; Sharkey et al., 2006: 558 [synonym of Disophrys]. Platyagathis Turner, 1918a: 113. Type species: Platyagathis leaena Turner, 1918a, by monotypy; Parrott, 1953: 200; Shenefelt, 1970b: 423. syn. nov.

Diagnosis. Length: 5–12 mm; colour: commonly yellow and dark brown (Fig. 10A), rarely with BROW colour pattern (Fig. 10B); inter-antennal region with medial elevated mound that may be broadly rounded to carinate, paired carinae or protuberances may be present (Fig. 7B); frons with lateral carinae (Fig. 3B); notauli present, deeply impressed and scrobiculate (Fig. 3B); tarsal claws cleft but base not pectinate (e.g., Fig. 2A); mid tibial preapical spines absent; fore tibial spur not elongate (e.g., Fig. 2F); pair of longitudinal carinae present on ventral surface of hind trochantellus (Fig. 2G), although can be lacking in non-Australian members of the genus (e.g. Sharkey et al. 2009); fore wing cell 1–RS always present and sessile, distinctly quadrate, vein 2–2RS present (Fig. 4B); ovipositor short, length less than 0.5 metasoma length (e.g., Fig. 9D). Comments. Most members of this genus exhibit variation in the yellow and dark brown colour pattern associated with a putative mimicry complex. However, an interesting aspect of this genus in Australia is the presence of two species exhibiting the BROW colour pattern. One species, D. leaena (Turner) (Fig. 10B), represents the most southerly occurrence for Disophrys, occurring in the Southwestern region of the



15

continent. It is nearly identical in colour and pattern to Therophilus latibalteatus (Cameron) (e.g., Fig. 8C), including possessing a distinct propodeal setal field. The other Disophrys species exhibiting the BROW pattern occur on the opposite side of the continent in the Torresian region. Species richness and distribution. Disophrys is well represented in Australia with over 20 species recognised, seven of which are described. Most of the species diversity occurs in the northern parts of the continent in the Timorian, Torresian and northern Kosciuskan regions. However, one species occurs in the temperate Southwestern region (see under Generic distribution and Comments above).

List of Australian species Disophrys diversipes Turner, 1918b Disophrys diversipes Turner, 1918b: 227. Holotype BMNH ♀; Type locality: Kuranda, Queensland. Parrot, 1953: 200 [catalogue]; Shenefelt, 1970b: 393 [catalogue]; Yu et al., 2005 [catalogue].

Disophrys leaena (Turner, 1918a), comb. nov. Platyagathis leaena Turner, 1918a: 114 [examined]. Holotype BMNH ♀; Type locality: Yallingup, Western Australia. Parrot, 1953: 200 [catalogue]; Shenefelt, 1970b: 423 [catalogue]; Yu et al., 2005 [catalogue].

Disophrys nigropectus Turner, 1918b Disophrys nigropectus Turner, 1918b: 229. Holotype BMNH ♀; Type locality: Kuranda, Queensland. Parrot, 1953: 200 [catalogue]; Shenefelt, 1970b: 399 [catalogue]; Yu et al., 2005 [catalogue].

Disophrys ruberrima Turner, 1918b Disophrys ruberrima Turner, 1918b:226. Holotype BMNH ♀; Type locality: Mackay, Queensland. Parrot, 1953: 200 [catalogue]; Shenefelt, 1970b: 400 [catalogue]; Yu et al., 2005 [catalogue].

Disophrys rufifrons Turner, 1918b Disophrys rufifrons Turner, 1918b: 226. Holotype BMNH ♀; Type locality: Port Essington, Northern Territory. Parrot, 1953: 200 [catalogue]; Shenefelt, 1970b: 400 [catalogue]; Yu et al., 2005 [catalogue].

Disophrys signatipennis Turner, 1918b Disophrys signatipennis Turner, 1918b: 225. Holotype BMNH ♀; Type locality: Kuranda, Queensland. Parrot, 1953: 200 [catalogue]; Shenefelt, 1970b: 401 [catalogue]; Yu et al., 2005 [catalogue].

Disophrys similipicta Turner, 1918b Disophrys similipicta Turner, 1918b: 228. Holotype BMNH ♀; Type locality: Mackay, Queensland. Parrot, 1953: 200 [catalogue]; Shenefelt, 1970b: 402 [catalogue]; Yu et al., 2005 [catalogue].

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Euagathis Szépligeti,1900 (Fig. 10C) Euagathis Szépligeti,1900: 62. Type species: Euagathis bifasciata Szépligeti, 1900, by subsequent designation Viereck, 1914: 32; Shenefelt, 1970b: 408 [catalogue]; Bhat & Gupta, 1977: 183 [key, description]; Chou & Sharkey, 1989: 186 [key, description]; Simbolotti & Achterberg, 1990: 3 [key, description]; Sharkey, 1992: 441 [in tribe Disophrini]; Simbolotti & Achterberg, 1995: 6 [key, description]; Sharkey, 1996:21 [key, description]; Achterberg & Chen, 2002: 311 [key, description]; Achterberg, 2004: 2 [key, description]; Yu et al., 2005 [catalogue]; Sharkey et al., 2006: 558 [notes, phylogeny, in tribe Disophrini]; Sharkey et al., 2009: 42 [key, description]. Chromomicrodus Ashmead, 1900: 129. Type species: Chromomicrodus abbotti Ashmead, 1900, by monotypy; Baltazar, 1961: 391 [synonym of Euagathis]; Shenefelt, 1970b: 409 [synonym of Euagathis]. Holcotroticus Cameron, 1902: 41. Type species: Holcotroticus ruficollis Cameron, 1902, by monotypy; Shenefelt, 1970b: 417 [catalogue]; Sharkey, 1992: 441 [in tribe Disophrini]; Simbolotti & Achterberg, 1995: 6 [synonym of Euagathis]. Balcemena Cameron, 1903: 130. Type species: Balcemena longicollis Cameron, 1903, by monotypy; Shenefelt, 1970b: 368 [catalogue]: Bhat & Gupta, 1977: 173 [catalogue]; Sharkey, 1992: 441 [in tribe Disophrini]; Achterberg & Chen, 2002: 313 [synonym of Euagathis].

FIGURE 10. Lateral and dorsal habitus of Agathidinae spp.: A, Disophrys sp.; B, D. leaena (Turner), exhibiting BROW (see text) mimicry complex colour pattern; C, Euagathis sp.; D, Amputostypos sp. Scale lines = 1 mm.

Diagnosis. Length: 5–12 mm; colour: commonly yellow and dark brown; inter–antennal region with paired carinae present (e.g., Figs 6B and 7B); frons without lateral carinae; notauli present, with (e.g., Fig. 3B) or without sculpturing, often complete or only present anteriorly; tarsal claws cleft, but base not pectinate (e.g., Fig. 2A); mid tibial preapical spines absent; fore tibial spur short, not exceeding 0.75 length of basitarsus (e.g., Fig. 2F); hind trochantellus without longitudinal carinae; fore wing cell 1–RS always present and sessile, vein 2–RS2 absent or basal sector only present; ovipositor short, length less than 0.5 metasoma length (e.g., Fig. 10D). Comments. Euagathis species display the characteristic yellow and dark brown colour pattern associated with a putative mimicry complex that also includes species of Cremnops and Disophrys. Euagathis is closely related to Coccygidium and Hypsostypos (Sharkey et al. 2006) but can be distinguished from both genera by the absence of longitudinal carinae on the hind trochantellus. Species richness and distribution. Euagathis is represented in Australia with about five species, only one of which is described. The genus occurs in the Torresian and northern Kosciuskan regions of the continent.



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List of Australian species Euagathis brevitibialis Achterberg, 2004 Euagthis brevitibialis Achterberg, 2004: 13. Holotype ANIC ♀; Type locality: Mount Webb National Park, Queensland. Yu et al., 2005 [catalogue].

Lytopylus Foerster, 1862 (Fig. 3C, 5A) Lytopylus Foerster, 1862. Type species: Lytopylus azygos, Viereck, 1905, by subsequent designation; Shenefelt, 1970b: 313 [catalogue]; Chou & Sharkey, 1989: 151 [synonym of Bassus]; Sharkey et al., 2009: 45 [reinstated]. Aerophilopsis Viereck, 1913: 129; Type species: Bassus erythrogaster Viereck, 1913, by monotypy; Shenefelt, 1970b: 312 [synonym of Agathis]; Marsh, 1979: 191 [synonym of Agathis]; Chou & Sharkey, 1989: 151 [synonym of Bassus]; Sharkey et al., 2009: 45 [synonym of Lytopylus]. Aerophilina Enderlein, 1920: 205; Type species: Aerophilina bicristata Enderlein, 1920, by monotypy; Shenefelt, 1970b: 312 [synonym of Agathis]; Chou & Sharkey, 1989: 151 [synonym of Bassus]; Sharkey et al., 2009: 45 [synonym of Lytopylus]. Ioxia Enderlein, 1920: 119; Type species: Ioxia faceta Enderlein, 1920, by monotypy; Shenefelt, 1970b: 312 [synonym of Agathis]; Chou & Sharkey, 1989: 151 [synonym of Bassus]; Sharkey et al., 2009: 45 [synonym of Lytopylus]. Hormagathis Brues 1926: 287. Type species: Hormagathis mellea Brues, 1926, by monotypy; Shenefelt, 1970b: 309 [synonym of Aerophilus]; Simbolotti & Achterberg, 1992: 6 [synonym of Bassus]; Sharkey et al., 2009: 45 [synonym of Lytopylus]. Obesomicrodus Papp, 1971: 338. Type species: Obesomicrodus nigra Papp, 1971, by monotypy; Simbolotti & Achterberg, 1992: 6 [synonym of Bassus]; Sharkey et al., 2009: 45 [synonym of Lytopylus]. Facilagathis van Achterberg & Chen, 2004: 78. Type species: Facilagathis spinulata Achterberg & Chen, 2004, by original designation; Sharkey et al., 2006: 557 [synonym of Bassus]; Sharkey et al., 2009: 45 [synonym of Lytopylus].

Diagnosis. Length: 4.5–5 mm; colour: variable, can be mostly dark brown to black or with mesosoma entirely orange; lateral carinae on frons always absent (e.g., Fig. 3A); notauli distinct, deeply impressed and scrobiculate (e.g., Fig. 3B); claws always non-cleft with triangular basal lobes present; mid tibial preapical spines always present (e.g., Fig. 2D); fore wing cell 1–RS petiolate and triangular cell 1–RS present (e.g., Fig. 4D); fore wing vein 2–RS2 always absent (e.g., Fig. 4D); possess strong posterior transverse carina between hind coxal cavities and metasomal foramen (Fig. 5A); metasomal tergites T1–T3 with prominent striate sculpturing throughout (Fig. 3C); ovipositor length greater than 0.5 metasoma length (e.g., Fig. 8B), can exceed body length. Comments. One of the four separate lineages of Bassus s.l. in the Sharkey et al. (2006) agathidine phylogeny, containing B. macadamia Briceño and Sharkey, was considered to be congeneric with the type species of the genus, B. calculator. Sharkey (2006) referred to this lineage as Bassus s.str., because he assumed that B. macadamia and B. calculator both possessed a strong transverse carina between the hind coxal cavities and the metasomal foramen (Fig. 5A), and striate sculpturing on metasomal tergites one to three Sharkey et al. (2006). However, later examination of the type revealed that B. calculator does not possess these characters and therefore there is no basis for postulating a congeneric relationship between these two species (Sharkey et al. 2009; pers. obs). Because of this, Sharkey et al. (2009) reinstated the oldest available name for species congeneric with B. macadamia, which is Lytopylus Foerster. Species richness and distribution. Lytopylus does not occur naturally in Australia, and is represented only by the introduced species L. rufipes, which is known from the southeast of the continent. This species now occurs widely outside of Europe and has been reared from more than 20 host species, many of which are tortricids (Shenefelt 1970b; Simbollotti and Achterberg 1992; Sharkey 1996; Yu et al. 2005). However, the nature of its introduction and hosts in Australia are unknown.

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List of Australian species Lytopylus rufipes (Nees von Esenbeck, 1812) comb. nov., introduced Microdus rufipes Nees von Esenbeck, 1812: 189. Holotype destroyed, syntypes lost, neotype MNHN ♀; Type locality: Germany. Braunsia rufipes (Nees von Esenbeck); Telenga, 1955: 277 [generic transfer]; Shenefelt, 1970b: 375 [catalogue]. Bassus rufipes (Nees von Esenbeck); Simbolotti & Achterberg, 1992: 35 [key, description, neotype designation, generic transfer]; Sharkey, 1996: 48 [key, description]; Yu et al., 2005 [catalogue]. Bassus diversus Muesebeck, 1933: 48 [examined]. Holotype USNM ♀; Type locality: Mitsuoka, Nagano, Japan. Sharkey, 1996: 48 [synonymy]; Yu et al., 2005 [catalogue]. Braunsia germanica Enderlein, 1904: 436. Holotype DEI ♀; Type locality: Rügen, Germany. Enderlein 1908: 128 [synonymy]; Yu et al., 2005 [catalogue].

Therophilus Wesmael, 1837 (Fig. 3A & D, 4D, 5B, 8B, C & D) Therophilus Wesmael, 1837: 15; Type species: Microdus (Therophilus) conspicuous Wesmael, 1837, by subsequent designation Viereck, 1914; Shenefelt, 1970b: 313 [catalogue]; Chou and Sharkey, 1989: 151 [synonym of Bassus]; Sharkey et al., 2009: 47 [elevated generic status]. Orgiloneura Ashmead, 1900: 355; Type species: Orgiloneura antipoda Ashmead, 1900, by monotypy [examined]; Parrott, 1953: 198 [catalogue]; Shenefelt, 1970a: 251[catalogue]; Sharkey et al., 2009: 47 [synonym]. Agathiella Szépligeti, 1902: 73; Type species: Agathiella pedunculata Szépligeti, 1902, by monotypy [examined]; Turner, 1918a, 108 [key, descriptions]; Parrott, 1953: 198 [catalogue]; Shenefelt, 1970b: 312 [synonym of Agathis]; Chou & Sharkey, 1989: 151 [synonym of Bassus]; Sharkey et al., 2009: 47 [synonym]. Aerophiliodes Strand, 1911: Type species: Aerophiliodes testaceator Strand, 1911, by monotypy; Shenefelt, 1970b: 308 [catalogue]; Sharkey, 1992: 441 [in tribe Microdini]; Sharkey et al., 2009: 47 [synonym].

Diagnosis. Length: 3.5–12.5 mm; colour: highly variable, can be entirely yellow, black or dark brown, or with varying combinations of yellow or orange with black or dark brown, or display BROW colour pattern (Figs 8B–D); lateral carinae on frons always absent; notauli present (e.g., Fig. 3B) or absent (Fig. 3A); claws always non-cleft, small to large quadrate or rounded basal lobes present (e.g., Fig. 2B); mid tibial preapical spines always present (e.g., Fig. 2D); fore wing cell 1-RS present (Fig. 4D); fore wing vein 2–RS2 always absent (Fig. 4D); ovipositor length greater than 0.5 metasoma length (Fig. 8B), can exceed body length. Comments. Therophilus was originally erected as a subgenus of Microdus, and treated as a synonym of Bassus prior to Sharkey et al. (2009) who resurrected the genus, along with Camptothlipis and Lytopylus, in an attempt to deal with the polyphyletic nature of Bassus (Sharkey et al. 2006). However, Therophilus, as defined above, is still polyphyletic (see comments above under the Bassus problem). To date, no distinguishing morphological features have been identified that enable the reliable identification of the various Therophilus groups, thus precluding them from being recognised as separate genera. The taxonomic changes below mostly came about from the tranfer of species from Bassus s.l., following the generic changes invoked by Sharkey et al. (2009), and recognition that some species were misplaced generically (e.g., in Agathis) following examination of the relevant types. Species richness and distribution. Therophilus is by far the largest and most widespread Agathidine genus in Australia. Examination of material from major collections indicates that there are at least 150 species in Australia, of which only 17 have been described. Members of the genus are found in a wide variety of environments, and are recorded from all Australian biogeographic regions. Interestingly, with only a few exceptions (see above), Therophilus is the only genus to naturally occur in the southern, more temperate regions of Australia including Tasmania.



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List of Australian species Therophilus antipodus (Ashmead, 1900), comb. nov. Orgiloneura antipoda Ashmead, 1900: 355 [examined]. Holotype USNM, ♀; Type locality: Australia. Parrott, 1953: 198 [catalogue]; Shenefelt, 1970a: 251 [catalogue]; Yu et al., 2005 [catalogue]; Sharkey et al., 2009: 47 [generic transfer].

Therophilus festinatus (Turner, 1918a), comb. nov. Agathiella festinata Turner, 1918a: 111 [examined]. Holotype BMNH [catalogue]. Agathis festinata (Turner); Shenefelt, 1970b: 333 [generic transfer]. Bassus festinatus (Turner); Yu et al., 2005 [generic transfer].

♀; Type locality: Queensland. Parrott, 1953: 198

Therophilus latibalteatus (Cameron, 1906), comb. nov. Agathis latibalteata Cameron, 1906: 26 [examined]. lectotype BMNH ♂; Type locality: Australia. Shenefelt 1970b: 340 [lectotype designation]; Yu et al., 2005 [catalogue]. Agathiella latibalteata (Cameron); Turner, 1918a: 110 [key, notes, generic transfer]; Parrott, 1953: 198 [catalogue].

Therophilus leucogaster (Holmgren, 1868), comb. nov. Ischius leucogaster Holmgren, 1868: 427 [examined]. Holotype NHRS ♂; Type locality: Sydney, New South Wales. Orgilus leucogaster (Holmgren); Parrott, 1953: 198 [catalogue, generic transfer]. Agathis leucogaster (Holmgren); Shenefelt, 1970b: 340 [catalogue, generic transfer]; Yu et al., 2005 [catalogue].

Therophilus malignus (Turner, 1918a), comb. nov. Agathiella maligna Turner, 1918a: 112 [examined]. Holotype BMNH ♀; Type locality: Yallingup, Western Australia. Parrott, 1953: 198 [catalogue]. Agathis maligna (Turner); Shenefelt, 1970b: 342 [catalogue, generic transfer]. Bassus malignus (Turner); Yu et al., 2005 [catalogue, generic transfer].

Therophilus martialis (Turner, 1918a), comb. nov. Microdus martialis Turner, 1918a: 108 [examined]. Holotype BMNH ♀; Type locality: Kuranda, Queensland. Agathis martialis (Turner); Parrott, 1953: 198 [catalogue, generic transfer]; Shenefelt, 1970b: 343 [catalogue]; Yu et al., 2005 [catalogue].

Therophilus meridionalis (Turner, 1918a), comb. nov. Agathiella meridionalis Turner, (1918a: 110) [examined]. Holotype BMNH ♀; Type locality: Mount Wellington, Tasmania. Parrott, 1953: 198 [catalogue]. Agathis meridionalis (Turner); Shenefelt, 1970b: 343 [catalogue, generic transfer]. Bassus meridionalis (Turner); Yu et al., 2005 [catalogue, generic transfer].

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Therophilus minimus (Turner, 1918a), comb. nov. Agathiella minima Turner, (1918a: 113) [examined]. Holotype BMNH ♀; Type locality: Kuranda, Queensland. Parrott, 1953: 198 [catalogue]. Agathis minima (Turner); Shenefelt, 1970b: 344 [catalogue, generic transfer]. Bassus minimus (Turner); Yu et al., 2005 [catalogue, generic transfer].

Therophilus minor (Szépligeti, 1905), comb. nov. Agathiella minor Szépligeti, (1905: 53) [examined]. Holotype HNHM ♀; Type locality: Sydney, New South Wales. Agathis minor (Szépligeti); Shenefelt, 1970b: 344 [catalogue, generic transfer]. Bassus minor (Szépligeti); Yu et al., 2005 [catalogue, generic transfer].

Therophilus pedunculatus (Szépligeti, 1905), comb. nov. Microdus pedunculatus Szépligeti, 1905: 51 [examined]. Holotype HNHM ♂; Type locality: Sydney, New South Wales. Turner, 1918a: 106 [notes]. Agathis pedunculatus (Szépligeti); Parrott, 1953: 198 [catalogue, generic transfer]; Shenefelt, 1970b: 348 [catalogue]. Bassus pedunculatus (Szépligeti); Papp, 2004: 175 [catalogue, generic transfer]; Yu et al., 2005 noted that Bassus pedunculatus (Szépligeti, 1905) was preocupied by B. pedunculatus (Szépligeti, 1902) from New Guinea.

Therophilus ruficeps (Szépligeti, 1905), comb. nov. Agathiella ruficeps Szépligeti, 1905: 52 [examined]. Holotype HNHM ♂; Type locality: Sydney, New South Wales. Turner, 1918a: 110 [notes]; Parrott, 1953: 198 [catalogue]. Agathis ruficeps (Szépligeti); Shenefelt, 1970b: 352 [catalogue, generic transfer]. Bassus ruficeps (Szépligeti); Papp, 2004: 175 [catalogue, generic transfer]; Yu et al., 2005 [catalogue]. Agathiella tenuissima Turner, 1918a: 111 [examined]. Holotype BMNH ♀; Type locality: Australia, Victoria. Parrott, 1953: 198 [catalogue]. Agathis tenuissima (Turner); Shenefelt, 1970b: 360 [catalogue, generic transfer]. Bassus tenuissimus (Turner); Yu et al., 2005 [catalogue, generic transfer] syn. nov.

Therophilus rufithorax (Turner, 1918a), comb. nov. Microdus rufithorax Turner, 1918a: 107 [examined]. lectotype BMNH ♀; Type locality: Kalamunda, Western Australia. Agathis rufithorax (Turner); Parrot, 1953: 199 [catalogue, generic transfer]; Shenefelt, 1970b: 353 [catalogue]; Yu et al., 2005 [catalogue].

Therophilus rufobrunneus (Turner, 1918a), comb. nov. Microdus rufobrunneus Turner, 1918a:106 [examined]. Holotype BMNH ♀; Type locality: Townsville, Queensland. Agathis rufobrunneus (Turner); Parrot, 1953: 199 [catalogue, generic transfer]; Shenefelt, 1970b: 353 [catalogue]. Agathis rufobrunnea (Turner); Yu et al., 2005 [catalogue].

Therophilus rugosus (Turner, 1918a), comb. nov. Agathiella rugosa Turner, 1918a: 112 [examined]. Holotype BMNH ♀; Type locality: Eaglehawk Neck, Tasmania. Agathiella rugosa (Turner); Parrot, 1953: 198 [catalogue]. Agathis rugosa (Turner); Shenefelt, 1970b: 353 [catalogue, generic transfer]. Bassus rugosus (Turner); Yu et al., 2005 [catalogue, generic transfer].



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Therophilus tricolor (Szépligeti, 1905), comb nov. Agathiella tricolor Szépligeti, 1905: 52 [examined]. Holotype HNHM ♀; Type locality: Sydney, New South Wales. Turner, 1918a: 109 & 110 [key, notes]; Parrot, 1953: 199 [catalogue]. Agathis tricoloralis (Szépligeti): Shenefelt, 1970b: 363 [catalogue, generic transfer; replacement name for Agathiella tricolor which is preoccupied by Agathis tricolor Gerstaecker,1858]. Bassus tricolor (Szépligeti); Papp 2004: 186 [catalogue, generic transfer]; Yu et al., 2005 [catalogue, noted Bassus tricolor (Szépligeti) was a senior secondary homonym of B. tricolor (Granger, 1949)].

Therophilus unimaculatus (Turner, 1918a), comb nov. Agathiella unimaculata Turner, 1918a: 111 [examined]. Holotype BMNH ♀; Type locality: Kuranda, Queensland. Parrot, 1953: 199 [catalogue]. Agathis unimaculata (Turner); Shenefelt, 1970b: 364 [catalogue, generic transfer]. Bassus unimaculatus (Turner); Yu et al., 2005 [catalogue, generic transfer].

Therophilus xanthopsis (Turner, 1918a), comb. nov. Microdus xanthopsis Turner, 1918a: 107 [examined]. Holotype BMNH ♀; Type locality: Yallingup, Western Australia. Agathis xanthopsis (Turner); Parrot, 1953: 199 [catalogue, generic transfer]; Shenefelt, 1970b: 365 [catalogue]; Yu et al., 2005 [catalogue].

Species nomen dubium Agathis dimidiata Brullé, 1846 Agathis dimidiata Brullé, 1846: 487. Holotype unknown, ♂; Type locality: Hobart, Tasmania. Parrott, 1953: 198 [catalogue]; Shenefelt, 1970b: 330 [catalogue]; Yu et al., 2005 [catalogue].

Comments. The holotype of Agathis dimidiata Brullé is unknown and presumably lost. During this study, no material examined could be matched with the original description which defines colour and the dorsal surface sculpturing relatively well. However, the original description is insufficient, with no detail regarding mouth parts, tarsal claws or wing venation provided, so we are unable to confirm its generic placement. In addition, the holotype was collected from Hobart over a 150 years ago. The habitat of this area has undergone a dramatic transformation since European settlement and if A. dimidiata’s natural range was confined to the Hobart region then there is the real possiblity that the species could now be extinct. For these reasons we hereby designate A. dimidiata as a nomen dubium.

Acknowledgments We wish to thank Kees van Achterberg, Cate Paull, Donald Quicke, Michael Sharkey and Bob Wharton for their help and advice at various stages of this project; the curators of the various collections listed above for the loan of material, and the staff at Adelaide Microscopy (The University of Adelaide) for assistance with the SEM. This project was funded in part by grants from the Australian Biological Resources Study, The University of Adelaide, and The Nature Foundation of South Australia.

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