Revision of the stiletto fly genera Acupalpa Kröber and Pipinnipons Winterton (Diptera, Therevidae, Agapophytinae) using cybertaxonomic methods, with a key to Australasian genera

Abstract Australian stiletto flies of the sister-genera Acupalpa Kröber, 1912 and Pipinnipons Winterton, 2001 (Diptera: Therevidae: Agapophytinae) are revised. Twelve new species of Acupalpa are described, while Acupalpa imitans (White, 1915), comb. n. is transferred from Pipinnipons and Acupalpa albimanis (Kröber, 1914), comb. n. is transferred from Ectinorhynchus Macquart as a senior synonym of Acupalpa pollinosa Mann. The total number of species of Acupalpa is therefore increased to 19: Acupalpa albimanis (Kröber), comb. n., Acupalpa albitarsa Mann, Acupalpa boharti sp. n., Acupalpa divisa (Walker), Acupalpa dolichorhyncha sp. n., Acupalpa glossa sp. n., Acupalpa imitans (White), comb. n., Acupalpa irwini Winterton, Acupalpa melanophaeos sp. n., Acupalpa miaboolya sp. n., Acupalpa minuta sp. n., Acupalpa minutoides sp. n., Acupalpa notomelas sp. n., Acupalpa novayamarna sp. n., Acupalpa rostrata Kröber, Acupalpa semirufa Mann, Acupalpa westralica sp. n., Acupalpa yalgoo sp. n. and Acupalpa yanchep sp. n. Three new species of Pipinnipons are described, increasing the total number of species to five: Pipinnipons chauncyvallis sp. n., Pipinnipons fascipennis (Kröber), Pipinnipons kampmeierae sp. n., Pipinnipons kroeberi Winterton, and P. sphecoda sp. n. Pipinnipons and Acupalpa are rediagnosed in light of the new species presented herein and revised keys to species are included. A dichotomous key to genera of Australasian Therevidae is included. As an empirical example of cybertaxonomy, taxonomic descriptions were composed using a character matrix developed in Lucid Builder (in Structured Descriptive Data (SDD) format) to generate natural language descriptions supplemented by online specimen and image databases. Web resources are provided throughout the document including: a) links to high resolution colour images of all species on Morphbank, b) registration of authors, publications, taxon names and other nomenclatural acts in Zoobank, with assignment of Life Science Identifiers (LSIDs) for each, c) links to Genbank accession records for DNA sequences, and d) assignment of LSIDs to specimen records with links to respective records in an online Therevidae specimen database.

The concept itself is not new, with single zoological registries (e.g. Brown 1961) and rapid descriptive processes (e.g. Erwin and Johnson 2000) espoused previously, but the actual empirical use of informatics tools to enhance the taxonomic descriptive process (i.e. online databases) is only now becoming a reality. Several authors have embraced modern cybertaxonomic methods through the incorporation of such digital, web-based, resources in taxonomic descriptions (e.g. Johnson et al. 2008;Deans and Kawada 2008;Miller et al. 2009;Penev et al. 2010;Blagoderov et al. 2010). The methods used by these authors, and this paper, are empirical examples of how digital tools can significantly speed the process of documenting biodiversity through rapid generation of natural language descriptions derived from matrix based character data in a standardized format suitable for multiple use (e.g. distributed morphological ontologies).

Materials and methods
Adult morphological terminology follows McAlpine (1981) as modified by  and . Hauser and Irwin (2003) provide a convincing argument for the use of the term pubescence (sensu Nichols 1989) instead of pruinescence (sensu ) to describe the various types of microtrichia covering the adult body and is used here throughout the text. The term velutum ) is retained to describe a particular type of very dense, unidirectional and reflective (i.e. velvet-like) microtrichia typically found on the male abdomen (silver coloured) or on the femora of agapophytine therevids (variously coloured but usually dark). Genitalia were macerated in 10% KOH to remove soft tissue, then rinsed in distilled water and dilute glacial acetic acid, and dissected in 80% ethanol. Genitalia preparations were placed in glycerine in a genitalia vial mounted on the pin beneath the specimen.
Types are deposited in the following institutions and collections: Queensland Museum (Brisbane) (QM), Australian Museum (Sydney) (AMS), Australian National Insect Collection (Canberra) (ANIC), Senckenberg Deutsches Entomologisches Institut, Müncheberg, Germany (DEI), University of Queensland Insect Collection (Brisbane) (UQIC), University of California, Davis, Bohart Museum (UCDC), Western Australian Museum (Perth) (WAM), Michael E. Irwin private collection [to be ultimately housed in the California Academy of Sciences] (MEIC/CAS), Greg Daniels private collection [to be ultimately housed in the Australian Museum] (GDCB/AMS), Naturhistorisches Museum Wien, (NMW), Harvard University Museum of Comparative Zoology (MCZ), Oxford University Museum of Natural History (OUMNH), University of California, Riverside (UCR), Universität von Hamburg Zoologisches Institut und Zoologisches Museum (ZMUH). All types have been examined. Numbers quoted with individual specimens as MEI000000 are unique identifiers in the therevid database MANDALA and are attached to each specimen as a yellow or white label (Kampmeier & Irwin 2009). Links are provided in this document to Life Science Identifiers (LSIDs) to specimen records with links to respective records in an online MANADALA Therevidae specimen database and Discover Life (http://www.discoverlife.org). Note that some web browsers are not able to read and format RSS (Really Simple Syndication) feeds and/or XML without additional software extensions or plug-ins. Details of current issues with select web browsers and LSID resolvers can be found on the Biodiversity Information Standards (TDWG) LSID resolver website (http://lsid.tdwg.org/). Material examined lists were exported from MANDALA. Descriptions were constructed using Lucid Builder 3.5, using a matrix database of character states, which were then exported using a natural language function into XML and a text document. Links are provided to Genbank accession records for DNA sequences where available. Specimen images were taken using a digital camera with a series of images montaged using Helicon Focus (©HeliconSoft). Descriptions are aided by the provision of embedded URL links in the document to high-resolution digital images of all species in Morphbank. All nomenclatural acts are registered in Zoobank as per the recent proposed amendment to the International Code of Zoological Nomenclature for a universal register for animal names (Polaszek et al. 2005a,b;ICZN 2008).

Key to genera of Australasian Therevidae
The following key to genera supersedes those by  and  and includes all genera found throughout the region east of Wallace's Line, incorporating landmasses such as Australia, New Zealand, New Caledonia, Papua New Guinea, and eastern Indonesia. The enigmatic Taenogera genus group (sensu ) is herein included within an expanded concept of Agapophytinae . The subfamilies Xestomyzinae and Phycinae are absent from the Australasian region. Diagnosis. Antennal scape shorter than or equal to flagellum; antenna elongate, cylindrical, total length slightly longer than or equal to head length; upper part of frons flat or slightly concave above antenna; face either protruding anteriorly below antennal base, or broadly rounded, expansive, short dark setae often present; parafacial setae absent; palpus apically narrow or acute, not spatulate; mouthparts length variable, frequently elongate and forward projecting (Fig. 3H); male postocular ridge with single row of macrosetae immediately laterad of ocellar tubercle, female with more than one row; wing infuscate, usually strongly banded; setae absent on wing vein R 1 ; cell m 3 closed; velutum patches on fore and hind femora; femora without macrosetae; single type of setal pile on femora, setae not appressed; prosternal furrow without setae; post spiracular pile absent; pleuron orange to black, overlain with sparse silver pubescence; mid coxa without setae on posterior surface; gonocoxites with velutum patch on ventral surface (Fig. 3B); articulated gonocoxal process present; hypandrium present; ventral apodeme of parameral sheath forked; dorsal apodeme of parameral sheath 'T'-shaped (Fig. 3F); three spermathecae in female; spermathecal sac present, sac simple or with smaller additional lobes basally, often with outer elongate lobes; spermathecal ducts joining common duct before bursa (Fig. 3G), female     Comments. Acupalpa is a genus with some distinctive wasp mimicking species (Figs 1-2), often strikingly coloured with black and orange. The male terminalia are relatively conserved throughout both Acupalpa and Pipinnipons, and species identification is more easily and reliably accomplished using external characters of either sex. Closely related to Pipinnipons and Agapophytus, Acupalpa can be distinguished by the elongate, cylindrical antennae, scape not longer than flagellum, face usually expansive and protruding, and palpi that are acuminate or narrowly cylindrical. The latter two characters specifically differentiate Acupalpa from Pipinnipons, as the face is always narrow and the palpi spatulate in Pipinnipons. Agapophytus is separated from Pipinnipons and Acupalpa by the length of the scape ranging from relatively equal length, to significantly longer than the flagellum.
Comments. Ectinorhynchus albimanis is herein transferred to Acupalpa with A. pollinosa becoming a junior synonym of Acupalpa albimanis comb. n. Mann (1928) redescribed E. albimanis based on a series of specimens, but clearly did not examine the type, as his redescription does not match that in Kröber (1914) nor reflect characteristics of the type. Acupalpa albimanis is morphological similar to A. albitarsa and A. yanchep sp. n. The colouration of abdomen and tarsi is diagnostic for this species. Diagnosis. Frons profile rounded above antenna; antenna black, scape sometimes brown; wing irregularly banded; pleuron black; tarsi white with brown to black basitarsus; abdomen with sparse silver velutum on anterior segments (denser in male).
Comments. The distinctive tarsal colouration of A. albitarsa and the closely related A. yanchep sp. n., separates these species from all other Acupalpa. Acupalpa albitarsa is an eastern species while A. yanchep sp. n. is western. Females are difficult to distinguish but males differ in the shape of the frons and in general body shape and size. The white patterning of the scutum is less pronounced in this species. Diagnosis. Body size relatively small; frons rounded above antenna; scutum glossy black with bronze pubescence; tibia yellow with dark apices, fore tibia white-cream; abdomen black, without velutum.
Comments. Acupalpa boharti sp. n. is a small, dark species similar to A. miaboolya sp. n. This species is known only from the type series collected in southwestern Australia. The small body size and leg and body colouration are diagnostic for this species.
Etymology. This species is named in honour of the collector, R. M. Bohart. Diagnosis. Frons profile concave above antenna; antenna black; pleuron black; wing dark banded; femora and tibia black; abdomen black, segments 1-3 yellow at least laterally; abdominal velutum present in male.
Comments. The male of A. divisa has long been unknown, and herein described for the first time. Hardy (1939) proposed that this species was a synonym of A. pollinosa, but has been subsequently proved incorrect as corresponding sexes of both species are now known. Diagnosis. Mouthparts elongate; frons profile rounded above antenna; antenna black; scutum glossy black; pleuron black; wing dark banded; abdominal segments 1-3 orange, rest black; abdominal velutum absent.
Comments. Acupalpa dolichorhyncha sp. n. is a distinctive species with elongate mouthparts and orange-banded abdominal segments 1-3. This western species is morphologically similar to A. melanophaeos sp. n., also from Western Australia, and A. glossa sp. n. from Victoria. Etymology. The specific epithet is derived from Gr. dolichos, long; rhynchus, snout, referring to the elongate mouthparts.
Comments. Acupalpa glossa sp. n. is similar to A. dolichorhyncha sp. n. in colour pattern and elongated mouthparts, but is easily differentiated based on leg colour. This species is known only from the type series collected in Victoria.
Comments. The type of Phycus imitans was kept in the G.H. Hardy collection, which was moved from Brisbane to Katoomba, but is now apparently lost or destroyed by pests. While not stating whether he actually examined the type, Mann (1929) redescribed this species in Agapophytus based on an unjustified allotype designation and associated female from southeast Queensland.  transferred the species to Pipinnipons based on the description of  and the specimens referred to by Mann (1929). The two specimens examined by Mann (1929) as putative A. imitans were located in the QM collection and no further material has been collected from Tasmania. Based on the discovery of new material from southeast Queensland more closely matching the original description by  than any other material examined (including the allotype designated by Mann (1929)), a male specimen (MEI165187) is herein designated as a Neotype to stabilise the concept of the species. This problematic species has characteristics that indicate a closer relationship to species of Acupalpa (particularly palpi shape) and is herein transferred from Pipinnipons.
Comments. Acupalpa irwini is a relatively large species differentiated by the characteristic leg and abdomen colouration. This species is known only from the type series collected in Western Australia. Diagnosis. Frons profile rounded above antenna; scape yellow, flagellum dark; scutum grey to black; pleuron black (reddish posteriorly in female); wing banded infuscate; coxae orange; legs orange to yellow, tarsi dark distally and fore-basitarsus white; abdomen black, segments 1-3 orange to yellow; abdominal velutum absent.
Comments. Acupalpa melanophaeos sp. n. is a relatively large species from Western Australia with distinctive leg and abdomen colouration. It is morphologically similar to A. novayamarna sp. n. and A. notomelas sp. n. The coxae are pale in this species along with a rounded face, rather than protruding anteriorly in similar species (e.g. A. irwini).
Etymology. The specific epithet is derived from the Gr. melanos, black; phaeos, light, shiny, referring to the scutum colouration. Diagnosis. Frons profile rounded above antenna; antenna brown to black; scutum glossy black with pubescent stripes of grey and brown; pleuron black; wing faintly infuscate; femora and tibia dark, fore tibia pale distally; abdomen black, without silver velutum.
Description. Body length= 5.0-6.5 mm. Head. Frons wider than ocellar tubercle, profile rounded above antenna, pubescence sparse silver-grey, without setae, surface texture smooth; face broadly rounded, glabrous; gena with pale setae (female) or dark setae (male); parafacial glabrous; mouthparts relatively short (approximately equal to head length), or elongate and projecting anteriorly; palpus brown-black, acuminate; occiput glabrous, glossy black; antennal base flat; frons roughly level with eye in profile; antennal length approximately equal to head; scape light brown to black, length approximately equal to flagellum, scape with sparse black setae; flagellum black, base of flagellum with short, dark setae. Thorax. Scutum black, overlain with stripes of grey and brown pubescence; scutellum overlain with sparse grey pubescence; pleuron black, overlain with sparse silver-grey pubescence; wing largely hyaline, faint band midway (male) or infuscate with pale band midway, hyaline ocellations basally (female); haltere knob white; coxae black, overlain with silver pubescence; femora brown to black; tibia brown; fore tibia pale distally; tarsi black, mid and hind basitarsi pale basally. Scutal chaetotaxy: np, 3; sa, 1; pa, 1; dc, 3; sc, 1. Abdomen. Black, overlain with bronze pubescence, silver velutum absent; terminalia dark. Comments. Acupalpa miaboolya sp. n. is a relatively small, dark species from Western Australia very similar to A. boharti sp. n. This species can be differentiated from the latter based on scutal pattern; the scutum has grey and brown stripes in A. miaboolya sp. n. while the scutum of A. boharti sp. n. has more uniform brown-bronze pubescence.
Etymology. This species is named after the region in which the specimens were collected, Miaboolya beach, on the north-central coast of Western Australia.
Comments. Acupalpa minuta sp. n. is closely related to A. minutoides sp. n., sharing characteristics such as very small size, two notopleural setae and an antenna with a short scape and a greatly elongate flagellum. It can be differentiated based on the colour of the setae on the coxae and by the width of the frons. This species is known only from a single male individual from Western Australia.
Etymology. The specific epithet is derived from the L. minutus, small, little, referring to the diminutive body size.
Comments. See comments under A. minuta sp. n. This species is known only from three male specimens from Western Australia.

Acupalpa notomelas
Comments. Acupalpa notomelas sp. n. is similar to A. melanophaeos sp. n. and A. novayamarna sp. n., but can be differentiated based on abdomen colouration. The palpi are very slender and elongate.
Comments. This species is very similar to A. melanophaeos sp. n., but can be differentiated by the pale genal setae, larger proportion of the pleuron being orange, and male frons slightly wider. Only the male is known for this western species.
Comments. The type of A. rostrata is apparently destroyed.  redescribed this distinctive species, without designating a neotype as the species was still identifiable based on the original description alone. With the description of the new species herein a neotype is designated to stabilise the taxon and remove any possibility of confusion in the future. Acupalpa rostrata is differentiated from other Acupalpa species by the unique leg and antennal colouration. Acupalpa semirufa Mann urn:lsid:zoobank.org:act:CF947CED-BCB0-4A62-AFDF-433DDBF3E1D3 http://species-id.net/wiki/Acupalpa_semirufa Fig. 19 Acupalpa semirufa Mann 1929: 27;Irwin and Lyneborg 1989: 354  Diagnosis. Frons profile concave above antenna; antenna dark; scutum dark; pleuron black; wing dark banded; femora orange to yellow; tibia yellow, darker distally; abdomen black with segments 2-3 yellow-orange, overlain with silver velutum in male.
Comments. Acupalpa semirufa is a common species in south-eastern Queensland and northern New South Wales. It is similar to A. divisa and A. yalgoo sp. n., but can be differentiated easily from the former by the orange leg colour (black in A. divisa) and from the latter by the projecting face (rounded in A. yalgoo sp. n.) and two wing bands (single band in A. yalgoo sp. n.). Diagnosis. Frons profile rounded above antenna; scape yellow-brown, flagellum black; scutum grey-black with dark and pale stripes; pleuron black; wing dark banded; femora yellow with extensive dark suffusion dorsally [hind femur dark]; tibia brown; abdomen black with brown pubescence, silver velutum absent.
Comments. Acupalpa westralica sp. n. is known only from the holotype female from southern Western Australia. This species superficially resembles A. rostrata in colouration, but the head shape suggests a close relationship to A. notomelas sp. n.
Etymology. The specific epithet is derived from the western distribution of this species.
Comments. Acupalpa yalgoo sp. n. is a western species similar to A. semirufa and A. glossa sp. n., but differs from both in leg colouration and wing patterning.
Etymology. The specific epithet is derived from the type locality of this species, near the Western Australian township of Yalgoo. Diagnosis. Frons profile rounded above antenna; antenna dark; scutum black with irregular brown and white pubescent markings; pleuron black; wing irregularly banded; legs dark, tibia pale basally; abdomen black [sometimes orange apically in female], silver velutum present in male.
Comments. Acupalpa yanchep sp. n. is morphologically similar to A. albitarsa, with females difficult to separate except for more pronounced white scutal patterning in many individuals. This species is found in western Australia while A. albitarsa is found in eastern and southern regions. There is a pronounced size difference in the sexes of A. yanchep sp. n. with males considerably smaller than females. The leg colouration and scutal patterning is distinctive for this species.
Etymology. This species is named after the type locality, the township of Yanchep, Western Australia. Diagnosis. Antenna elongate, cylindrical, total length slightly longer than or equal to head length; scape shorter than flagellum; frons flat, smooth; face as narrow strip below antenna, glabrous; palpus spatulate apically; mouthparts short; occiput with single row of postocular setae immediately laterad of ocellar tubercle in male, multiple rows in female; wing banded infuscate or hyaline; setae absent on wing vein R 1 ; cell m 3 closed; elongate velutum patches on fore and hind femora; femora without macrosetae; single type of setal pile on femora, setae not appressed; prosternal furrow without setae; mid coxa without setae on posterior surface; post spiracular pile absent; gonocoxites with velutum patch on ventral surface (Fig. 24); articulated gonocoxal process present; hypandrium present; ventral apodeme of parameral sheath forked; dorsal apodeme of parameral sheath 'T'-shaped; three spermathecae in female; spermathecal sac present, usually with two smaller, additional lobes and/or outer reticulated lobes along length; spermathecal ducts joining common duct before bursa; female with A1 and A2 acanthophorite spines well developed; female sternite 8 emarginate along posterior margin. Comments. Pipinnipons is a distinctive genus of wasp mimicking therevids, often with metallic pubescence, yellow and black marking and banded wings (Fig. 23). It can be distinguished among related genera by the elongate, cylindrical antennae, scape not longer than flagellum, narrow face and palpi spatulate. The latter two characters specifically differentiate Pipinnipons from Acupalpa, as the face is broadly rounded, often produced, and the palpi are acuminate or narrowly cylindrical in Acupalpa. While the mouthparts are of variable length in Acupalpa (and often elongate and forward projecting), the mouthparts of Pipinnipons are always relatively short. As stated in the comments under Acupalpa, Agapophytus is separated from Pipinnipons and Acupalpa by the length of the scape ranging from relatively equal length, to significantly longer than the flagellum. The modified setae patch on abdominal tergite 2 mentioned by  as a characteristic of Pipinnipons is not present in all the new species described here, and is no longer considered diagnostic for the genus as it is also found sporadically in other, unrelated genera such as Neodialineura Mann, 1928 andBonjeania Irwin &Lyneborg, 1989. The male terminalia are relatively conserved throughout the genus, and species identification is more easily done using external characters of both sexes. Pipinnipons is distributed along coastal eastern Australia from northern Queensland to Tasmania.
Comments. Pipinnipons chauncyvallis sp. n. is known only from a small conservation area near Bagdad, Tasmania. This species differs from all other Pipinnipons by the body colouration and the numerous pale setae on the frons.
Etymology. This species is named after the type locality, Chauncyvale Wildlife Sanctuary, owned by the Chauncy family who established and maintain the sanctuary. Diagnosis. Wing dark banded; pleuron orange to maroon; legs orange to maroon, tarsi lighter; abdomen dark red, tergite 2-3 red-brown, gold-bronze velutum on segments 4-7.
Comments. The gold-bronze abdominal velutum covering in both sexes and the dark orange pleuron and leg colouration make P. fascipennis easily recognisable. This species is found in closed forest areas, including rainforest. Diagnosis. Wing mostly hyaline; pleuron black; coxae, femora and tibia orange; abdomen black, segments 2-3 orange with dark medial patch.
Comments. Pipinnipons sphecoda sp. n. is a relatively large, apparently wasp-mimicking species known only from female specimens collected from various sites in Tasmania. The dramatic colouration of species makes it quite unlike any other stiletto fly species.
Etymology. The species epithet is derived from the Gr. sphekodos, wasp-like.