Corresponding author: Christine L. Lambkin (
Academic editor: T. Dikow
Bush Blitz is a three-year multimillion dollar program to document the plants and animals in hundreds of properties across Australia’s National Reserve System. The core focus is on nature discovery – identifying and describing new species of plants and animals. The Bush Blitz program has enabled the collection and description of beeflies (Diptera, Bombyliidae) from surveys in Western Australia and Queensland. Three new species of Australian beeflies belonging to the Exoprosopini are described;
While there are more than 140,000 published species in Australia, more than 40 per cent of continental Australia has never been comprehensively surveyed by scientists. This research was supported through funding from the Bush Blitz species discovery program, a partnership between the Australian Government, BHP Billiton and Earthwatch Australia. This innovative partnership harnesses the expertise of many of Australia’s top scientists from museums, herbaria, universities, and other institutions and organisations across the country. Bush Blitz is expected to uncover hundreds of new species and provide baseline scientific data that will help us protect our biodiversity for generations to come.
This paper describes three species of beeflies from the Exoprosopini (Diptera, Bombyliidae, Anthracinae); two captured during Bush Blitz surveys and a third species collected from south-western Queensland (Qld). All three species belong to genera recently described (
The beeflies belong to the Family Bombyliidae, a very large, cosmopolitan family of stoutly built flies, mostly with very characteristic venation. Almost 5000 species have been described worldwide (
Australian exoprosopines are large beeflies of diverse and striking appearance (
Adult Australian exoprosopines favour warm, sunny localities, especially in the more arid regions. Most have a strong, hovering flight, and are commonly taken from blossom, or sitting on patches of bare earth. Adults are pollen and nectar feeders, and many are important pollinators of native plants. Many species can be collected congregating on hilltops, demonstrating a landmark-based mating system (
This paper describes three new species of exoprosopine beeflies; two captured during Bush Blitz surveys and a third species collected from south-western Qld.
Four male specimens (
Previous phylogenetic analysis of the worldwide Exoprosopini showed that the Australian bombyliids that were previously placed in
Revised keys are provided for the genera of the Australian
We attempted to use cybertaxonomic tools to produce this paper as had been used to streamline taxonomic publication of new fly species by
The following collection acronyms are used in the text: Australian Museum, Sydney, New South Wales, Australia (AM); Queensland Museum, South Brisbane, Queensland (QM); Western Australian Museum, Perth, Western Australia (WAM). Numbers quoted with individual specimens are unique identifiers (e.g. WAM 82396, T152479 (QM), K 253702 (AM)) from the respective institutions database and are attached under each specimen on a white label. A single hind leg was removed from one specimen of each species and placed into absolute ethanol for frozen tissue storage at QM for future DNA extraction. Those samples were given a tissue number (e.g. A007534) that was entered into the QM Vernon database and attached under each specimen on a yellow label.
For explanation of morphological abbreviations, see Appendix 1.
The genitalia of each species were prepared by dissecting the terminal abdominal segments and then placing in cool 10% KOH overnight. Following maceration the specimen was washed, and then dissected in distilled water. Dissected genitalia were placed in alcohol for microscopic examination and into K–Y® Jelly for photography. All dissected parts from a specimen were placed in a genitalia vial containing glycerine which was pinned beneath the identification label.
Images were taken of the whole fly, external features, and dissected genitalia. A series of multiple-focal-depth digital images were taken using a Canon EOS 500D digital camera fitted, via a Leica 10446175 1x SLR Projection Lens, to a Leica MZ6 stereo dissecting microscope, and combined into a high resolution serial montage image using Helicon Focus v.5.2 Pro (
Distribution maps were produced using ArcView GIS version 3.1 (
We intended to use cybertaxonomic methods to document these newly discovered Australian beeflies, enabling descriptions of the three new species to be generated using web resources to populate electronic documents through links to Morphbank, Life Science Identifiers, and Zoobank as had been done by
Several initiatives around the world have been developing tools to bring revisionary taxonomy to the web. Recent examples include software produced through the CATE (Creating a taxonomic e-science,
The paper has been semantically tagged and enhanced using the Pensoft Mark Up Tool (PMT) which is based on the US National Library of Medicine’s DTD (Document Type Definitions) TaxPub extension
The nomenclatural and distributional information will be included in the Australian Faunal Directory (AFD), an open-access online catalogue of taxonomic and biological information on all animal species known to occur within Australia (ABRS, 2009), and the Australian Natural Heritage Assessment Tool (ANHAT), an open-access online map-supported database developed by the Australia Heritage Division of the Department of Sustainability, Environment, Water, Population and Communities that helps identify and prioritise areas for their natural heritage significance, focusing on biodiversity (
Phylogenetic analysis was based on 207 morphological characters from
Multistate characters used for phylogenetic analyses have been treated as unordered (non-additive
Phylogenetic analyses completed 100 random step-wise addition searches, with tree-bisection-reconnection (TBR) branch swapping, MULPARS, and branches having maximum length zero collapsed to yield polytomies in effect using PAUP*4.0b10 (
We used Bremer support (
Cladograms and character distribution were analysed in WinClada version 1.00.08 (
Cladistic analysis of the 42 taxa of 141 non-constant characters produced five most parsimonious trees (MPTs) of length =931, CI = 0.243, CI excluding uninformative characters = 0.231, and RI = 0.468. The five trees differ only in the placement of
Previous phylogenetic analysis of 207 morphological characters for the worldwide Exoprosopini showed that the Australian bombyliids that were previously placed in
In
In this phylogenetic analysis,
With the description of the three new species and the transferral of
Small black, rounded, dense, adpressed metallic scales dorsally on thorax and abdomen (
Included species:
Wing length 20.0 mm
Large dark flies with distinct triangular basal infuscation on the wings wings. Face and frons with transparent scales. Occiput with white scales broadly filling indentation. Collar whitish-cream. Broad laterothoracic stripe of dense white flattened scales. Scutum black with lime-green metallic scales except pink metallic scales anterolaterally to PR bristles and posterolaterally anterior to APA. Scutellum with lime-green metallic scales; very long, white, flattened-scale fringe on posterior margin. Widened base of costa with reddish-brown scales, white scales posteriorly. Wing pattern dimidiate (
Collection sites.
One of five most parsimonious cladograms (931 steps, CI = 0.24, RI =0.47). Part 1. Black circles = unique character changes, open circles = homoplasious changes. Bremer supports over branches.
One of five most parsimonious cladograms (931 steps, CI = 0.24, RI =0.47). Part 2. Black circles = unique character changes, open circles = homoplasious changes. Bremer supports over branches.
(
Due to extended storage in propylene glycol as retrieval of sample was prevented by extensive and prolonged flooding the specimen bears few setae, hairs or scales, therefore colour patterns referred to in the description are based on those remaining, usually at junctions of sclerites.
Wing length 9.0 mm.
Small dark flies with heavily infuscated wings, hyaline only apically and medial spot. Face orange with shiny reddish-brown scales, frons black with shiny black scales. Collar yellow. Narrow laterothoracic stripe of whitish scales. Scutum black with dull lime-green metallic scales except pinkish metallic scales anterolaterally and posteromedially. Scutellum dark brown, darker basally with royal-blue metallic scales, purple metallic scales laterally and posteriorly. Widened base of costa with shiny reddish-brown scales, no paler scales posteriorly. Wing pattern broadly dimidiate (
This species is named
(
Dimidiate wing pattern as in
Included species:
Paratypes.
Wing length 14 mm
Medium, dark, densely setose, flies with black, dimidiate wings with five indistinct yellowish spots (
This species is named as a noun in apposition after the three-year, multimillion dollar Bush Blitz program that organised and funded the survey on Charles Darwin Reserve, Karara, Lochada and Kadji Kadji Pastoral Leases in Western Australia on which this species was collected. The core focus of the Bush Blitz program is to document the plants and animals in hundreds of properties across Australia’s National Reserve System, and on nature discovery – identifying and describing new species of plants and animals. The Bush Blitz program also funded the survey in western New South Wales and Queensland on which
(
On collection, this species appeared similar to the two male specimens of
Wing with medial hyaline band not linear and narrowing apically, apical infuscated band not meeting posterior wing margin more broadly than medial hyaline band. Gonocoxae deeply narrowed medially, broadly indented basally, with tufts of thickened setae ventromedially, H projecting but not forming a finger-like extension; AE short; EP with medioventral process above AE; very long AAES reaching G margins; EJA racquet-shaped, very long. Sperm pump short with unpigmented papillae, apical endplate simple with thin processes; thick-walled round SR with distinct basal bulb.
Included species:
See reference to the rediagnosis of the genus
Wing with medial hyaline band linear and narrowing apically, apical infuscated band meeting posterior wing margin twice breadth of medial hyaline band (
The name for the genus
Included species:
See reference to the erection of the genus
Large dark flies (wing length 15-20 mm), wings as in
Map of distribution. Closed circle -
In
This species has been collected in the southern Australian Bassian region, from semi-arid and arid mallee areas.
The finger-like projection on the H (
1 | Metallic scales on body, black reflecting blue, bluish-black, green or maroon, no white or yellow scales on T2–7 or on S2–7 ( |
|
– | No metallic reflecting black scales; white or yellow scales present T2–7, usually distinct bands or lateral triangles on T3 ( |
2 |
2 (1) | Wing dimidiate and at least apical half of anal cell margin hyaline, at most short narrow lobe following m-m into m2, no medial hyaline band ( |
|
– | Wing not dimidiate or anal cell fully infuscated; distinct medial hyaline band usually present ( |
3 |
3 (2) | Female spermathecal reservoir a long cylinder; T2 with black scales; T6–7 with white scales; proboscis extending beyond oral cavity, not longer than head | |
– | Female spermathecal reservoir round to subquadrate never a long cylinder; T2 with some yellow scales unless T6 or T7 with black scales or proboscis longer than head .. | 4 |
4 (3) | Male with no medioventral process on epiphallus above aedeagus, anterior arms of aedeagal sheath long, reaching gonocoxal margins; quadrate sub-epandrial sclerites in epandrium. EITHER Deeply infuscated wings, only apex hyaline; paler yellowish spots at base of R2+3, at base of CuA1, join of R1 and Rs, r-m and base of m-cu; T6 black scales; OR medial hyaline band a narrow line; black scales forming median circle apex of T2 and base of T3, yellow scales anteriorly and laterally on T2, medially and laterally on T3 | |
– | Male with medioventral process on epiphallus above aedeagus ( |
5 |
5 (4) | Male with anterior arms of aedeagal sheath long, reaching gonocoxal margins ( |
6 |
– | Male with anterior arms of aedeagal sheath short, not reaching gonocoxal margins; if ventral ridge on gonocoxae very small or absent then hypandrium not projecting. Abdominal yellow scales at least anteriorly T2, S2–3 with dense, white scales; hemispherical tufts of macrochaetae laterally on T1 white or yellow, not dark reddish-brown or black; wing infuscation not distinctly variegated, IF yellow scales only anteromedially T2 then scales on S5–7 not black, at most reddish-brown | 7 |
6 (5) | Medial hyaline band linear, narrowing anteriorly, with apex of anal cell and cup hyaline ( |
|
– | Medial hyaline band not linear, not narrowing anteriorly; male gonocoxae with no finger-like projection from hypandrium | |
7 (5) | Male epandrium with strongly grouped setae on anterolateral flange; ventral ridge on gonocoxae large, distinctly projecting basally, hypandrium not projecting, epiphallus with rounded projecting lateral lobes, ejaculatory apodeme short, extending beyond gonocoxae by less than length of gonostylus, hind-tibial scales not protruding, dark flies, T2 and T4 mostly black scales | |
– | Male epandrium with loose setae, without an anterolateral flange; hind-tibial scales protruding, pale yellowish flies with striped abdominal vestiture, T2 and T4 mostly yellow scales |
1 | No pre-apical infuscated band on wing ( |
2 |
– | Pre-apical infuscated band on wing present | 3 |
2 (1) | Infuscation of wing blade almost complete except for hyaline apical area and isolated spot over dc ( |
|
– | Infuscation of wing blade only extending over half wing area, indistinct extension along R4+5 and isolated mottled area along m-m ( |
|
3 (1) | Anal and posterior cells with notched hyaline area, infuscation extending along CuA2; apically-directed spur-vein on i-r1 cross vein |
|
– | Anal and posterior cells without extension along CuA2, rarely spur-vein on i-r1 cross vein | 4 |
4 (3) | Hyaline medial band continues anteriorly through entire r5; dark thorax; abdomen: males dark prussian-blue, almost black; females bluish-green |
|
– | Hyaline band usually through dc anteriorly, not entirely through r5, or absent; thorax and abdomen not as above | 5 |
5 (4) | Collar white with contrasting tuft of black lateral Ma at base of pronotal lobe, bright green thorax, anterolaterally dark maroon; bright, dark blue abdomen; anal and cup fully infuscated |
|
– | Collar entirely white or yellow, at most 4 reddish Ma above postpronotal lobe; thorax, abdomen, anal and cup not as above | 6 |
6 (5) | Face yellow, most facial setae shiny gold; blue-green thorax and abdomen |
|
– | Face orange-red to reddish-brown; if yellow, most facial setae black; thorax and abdomen not as above | 7 |
7 (6) | Thorax green | 8 |
– | Thorax dark, not green | 11 |
8 (7) | Thorax anterolaterally with dark maroon scales; anal and cup infuscation various | 9 |
– | Thorax entirely green; anal and cup hyaline apically | 10 |
9 (8) | Abdomen bluish-green; brown wing infuscation, anal and cup hyaline apically; blue face scales |
|
– | Abdomen entirely purple; black wing infuscation, anal and cup fully infuscated; purple face scales |
|
10 (8) | Thorax bright yellowish-green; abdomen blue to bluish-green metallic scales; infuscated wing band short, much narrower than hyaline band |
|
– | Thorax dark bluish-green; abdomen T2 blue-green, T3–7 blue, T4–6 admixed maroon at least laterally; infuscated wing band broader than hyaline band |
|
11 (7) | Abdomen purple with blue scales on T4–6, Queensland |
|
– | Abdomen entirely purple, no blue scales on T4–6, Western Australia |
|
1 | No yellow scales on T2–7 ( |
2 |
– | Abdominal yellow scales at least anteriorly T2; Ma on T1 white, not black |
|
2 (1) | Wing without small paler yellowish spots in infuscation, m-m without infuscation |
|
– | Wing with small paler yellowish spots in infuscation; m-m with infuscation ( |
3 |
3 (2) | Wing without short narrow lobe following m-m into m2 ( |
|
– | Wing with short narrow lobe following m-m in m1 into m2 |
|
Firstly, we thank the Bush Blitz program partners and the program managers from the Australian Biological Resources Study (ABRS), a section within the Parks Division of the Department of Sustainability, Environment, Water, Population and Communities (especially Brooke Glasser, Annabel Wheeler, and Kate Gillespie) for organising and funding the Bush Blitz field work on Charles Darwin Reserve, Karara, Lochada and Kadji Kadji Pastoral Leases in WA, funding the Bush Blitz survey of Culgoa Floodplains NP Qld, Culgoa NP and Ledknapper NR NSW (
CLL and Noel Starick thank the Mackenzie family of Plevna Downs Station for the enthusiasm and interest they always show in all forms of natural history, the welcome and hospitality they have given to us and all staff from the Queensland Museum, and the encouragement they provide to the local Natural History Society, regional property owners and community members on development of a knowledge base of biodiversity of arid areas. For providing collection permits to collect and take samples from National Reserves, we thank Jacqui Brock (Scientific Permit W1TK05498008: Queensland Parks and Wildlife Service, DERM, QLD) and Brendon Neilly (Scientific Research Licence S10016: NPWS, OEH, NSW). We thank David Britton and Jacqui Recsei (AM, Sydney) for supplying specimens of
We thank Neal Evenhuis and an anonymous reviewer for their positive comments on the manuscript. Lastly, but not least, we gratefully acknowledge the work of Geoff Thompson (QM) for photographing specimens and guidance in the use of the imaging systems, Paul Avern (QM) for help with writing the occurrence data in Darwin Core format, Federica Turco (QM) for production of the distribution map and help with the GBIF and Dryad uploads, Vladimir Blagoderov, (Natural History Museum, London) for help with Scratchpads, Debbie Paul (School of Computational Science, Florida State University Tallahassee) for assistance with Morphbank, and Lyubo Penev and Teodor Georgiev (Pensoft Publishers, Sofia, Bulgaria) for their support and assistance with data publication through GBIF and Dryad.
For a full description of these characters see Appendix 2 in
Morphological terminology follows that of
1. Head scales: (0) absent; (1) present
2. Face scale density: (0) absent; (1) sparse; (2) overlapping; (3) carpet
3. Frons scale density: (0) absent; (1) sparse; (2) overlapping; (3) carpet; (4) distinct dense medial patch
4. Frons with vertical groove: (0) absent; (1) depression; (2) groove
5. Frons with horizontal depression: (0) absent; (1) shallow; (2) distinct; (3) deep
6. W head/W thorax: (0) <; (1) =; (2) >
7. L antennae to compound eye/L scape: (0) <; (1) ≥; (2) ≥ 2x; (3) ≥ 4x
8. L antennal separation/L scape: (0) <; (1) ≥; (2) ≥ 2x; (3) ≥ 3x
9. Male compound eye separation /W OT: (0) meet; (1) <; (2) =; (3) ≤ 2x; (4) ≤ 3x; (5) > 3x
10. Female compound eye separation /W OT: (0) ≤ 2x; (1) ≤ 3x; (2) > 3x
11. OT to posterior margin of compound eye/ L OT: (0) ≤ OT; (1) > OT; (2) ≥ 2x; (3) ≥ 3x
12. L occiput/L OT: (0) ≤ 2x; (1) < 3x; (2) occiput long, well developed ≥ 3x
13. L vertex; i.e. L OT to occipital groove/ L OT: (0) ≤ L OT; (1) ≤ 2x; (2) wide > 2x
14. Depth occipital foveal depression: (0) no vertex; (1) not depressed; (2) shallow; (3) deep, slopes posteriorly at 45° to a short OG
15. W occipital foveal depression: (0) no vertex; (1) not depressed; (2) narrower than compound eye separation; (3) wider than compound eye separation
16. Apical occipital groove: (0) narrow; (1) wide rounded
17. Shape of head laterally: (0) round; (1) protruding but rounded, blunt; (2) conical
18. L proboscis: (0) < L oral cavity; (1) > L oral cavity; (2) > L head; (3) > 1.5x L head; (4) > 2x L head
19. L palps/ L proboscis: (0) rudimentary; (1) < 0.25x, short; (2) < 1, long
20. Genae setae surrounding oral cavity: (0) not grouped; (1) grouped laterally; (2) grouped apically, small tuft
21. L setae below antennae/L scape: (0) >; (1) ≤; (2) ≤ 0.5
22. Horizontal depression between antennae: (0) absent or shallow; (1) distinct; (2) deep
23. W of face projection/W compound eye including indentation posterior margin of eye: (0) < 1/4; (1) ≤ 1/2; (2) ≤ 1
24. W of the indentation on the posterior margin of the compound eye/L OT: (0) ≤ L OT; (1) > L OT; (2) ≥ 2x L OT
25. L of the line from the posterior margin of compound eye bisecting the compound eye facets/L OT: (0) absent; (1) <; (2) ≥; (3) ≥ 2x
26. L scape /L pedicel: (0) ≤; (1) ≤ 3x; (2) > 3x
27. L PP/ L pedicel: (0) ≤ 3x; (1) ≤ 4x; (2) > 4x
28. PP base shape: (0) broad not round base; (1) onion-like, round base abruptly narrowed; (2) medially divided, laterally; (3) conical, broad base, gradually narrowed
29. L PP rod/L base: (0) long > 2x; (1) short < 2x; (2) no thin rod
30. Distinct joint between PP and BSM: (0) absent; (1) present
31. L BSM/L pedicel: (0) absent; (1) ≤; (2) ≤ 2x; (3) ≤ 3x; (4) > 3x
32. BSM apical hairs: (0) absent; (1) present
33. L ASM/W BSM: (0) < W BSM, minute spine; (1) > W BSM; (2) > 2x; (3) conical twisted hat
34. Collar Ma: (0) pointed; (1) midstyle; (2) pectinate
35. Scm reflective vestiture: (0) bright; (1) very dull dark; (2) not reflective
36. Ma AN: (0) pointed; (1) midstyle; (2) pectinate
37. L PR bristles/ L PN: (0) > 2x; (1) >; (2) <; (3) absent
38. LT vestiture: (0) bare; (1) some hair; (2) dense hair
39. Ma LT: (0) absent; (1) pointed; (2) midstyle; (3) pectinate
40. MT vestiture: (0) bare; (1) some hair; (2) dense hair
41. L PA bristles/Scu: (0) absent; (1) < 0.5; (2) ≤; (3) >
42. L thorax/Scu: (0) ≤ 2x; (1) ≤ 3x; (2) > 3x
43. Scu vestiture reflective: (0) bright; (1) very dull dark; (2) absent
44. C1 very long setae: (0) absent; (1) some; (2) dense
45. L forefemur/ L coxa: (0) ≤ 1.5x; (1) ≤ 2x; (2) ≤ 2.5x; (3) > 2.5x
46. Forefemoral spines: (0) absent; (1) short < W femur; (2) long
47. Forefemoral long hairs: (0) absent; (1) some; (2) dense
48. Foretibial spicules: (0) absent; (1) some; (2) dense
49. L foretarsus/ L foretibia: (0) ≥ 1; (1) ≥ 0.75; (2) > 0.5
50. Foretarsal microchaetae: (0) absent; (1) very few < 10; (2) present
51. Foretarsal microchaetae: (0) absent; (1) ends bulbous; (2) ends slightly bent; (3) ends distinctly bent
52. L foreclaw/ L midclaw: (0) < claw; (1) ≤ half; (2) ≤ third
53. Midfemoral spines: (0) absent; (1) short < W femur; (2) some long
54. Midfemoral long hairs: (0) absent; (1) some; (2) dense
55. Midtibial spicules: (0) absent; (1) some; (2) dense
56. L midpulvilli/ L claw: (0) ≥; (1) <; (2) ≤ half; (3) < 0.2
57. Midpulvilli: (0) large, flattened, membranous; (1) small rounded setose; (2) chisel-conical
58. Hindfemoral spines: (0) absent; (1) short < 0.5 W femur; (2) some long
59. Hindfemur long hairs: (0) absent; (1) some; (2) dense
60. Long hindtibial scales: (0) no long scales; (1) some long scales; (2) fluffy - protruding; (3) feathery; (4) very long, dense, feathered fringes
61. Hindtibial spicules: (0) absent; (1) some spicules; (2) apical patch; (3) dense spicules
62. Hindtibial spicules L: (0) absent; (1) same; (2) inner row longer
63. L hindpulvilli/L claw: (0) ≥; (1) <; (2) ≤ half; (3) ≤ 0.2
64. Hindpulvilli: (0) large, flattened, membranous; (1) small rounded setose; (2) chisel -conical
65. Patagium: (0) absent hairs only; (1) present scales
66. Basicosta: (0) absent; (1) blunt; (2) sharp
67. Crossvein forming an extra anterior apical submarginal cell: (0) absent; (1) extra apical submarginal cell
68. R2+3 join R4+5: (0) basal to r-m > L r-m; (1) at r-m
69. R2+3 rises from R4+5: (0) acutely; (1) at right angles
70. 2nd r-m crossvein: (0) absent; (1) present
71. Spurvein base R2+3: (0) absent; (1) bump or present
72. R2+3 apical loop: (0) long apical loop; (1) loop > 180°; (2) at least 90° bend; (3) absent
73. i-r1, R2+3 to R4: (0) absent; (1) present
74. i-r1 crossvein: (0) absent; (1) straight; (2) slightly sinuous; (3) distinctly sinuous
75. Spurvein i-r: (0) absent; (1) bump or present
76. L i-r1/L r-m: (0) absent; (1) ≤; (2) ≤ 2x; (3) < 3x; (4) ≥ 3x
77. Spur-vein base R4+5: (0) absent; (1) bump or present
78. Spur-vein R4: (0) absent; (1) bump or present
79. R5/R1 meet wing: (0) R5 distal to R1; (1) equal; (2) R5 basal to R1
80. i-r2, R4 to R5: (0) absent; (1) present
81. M1: (0) straight; (1) slightly sinuous; (2) sinuous
82. L m-m/r-m: (0) ≤ 2x; (1) ≤ 3x; (2) > 3x
83. m-m: (0) straight; (1) slightly sinuous; (2) sinuous
84. m-m spurvein: (0) absent; (1) into m2; (2) into discal; (3) crossvein form basal cell
85. m-m to hind wing margin: (0) oblique; (1) parallel; (2) horizontal
86. M2: (0) straight; (1) slightly sinuous; (2) sinuous
87. r5 open: (0) open; (1) narrow < r-m; (2) just closed; (3) closed and stalked - acute; (4) closed and stalked -obtuse
88. m1 open: (0) open; (1) closed and stalked
89. m2 open: (0) open; (1) closed and stalked - acute
90. L mcu/r-m: (0) > 3x; (1) < 3x; (2) ≤ 2x; (3) ≤
91. m-cu: (0) straight; (1) slightly sinuous; (2) 90° basally; (3) sinuous
92. Spur-vein m-cu: (0) absent; (1) spur into discal cell
93. Spur-vein into M2: (0) absent; (1) spur-vein into m2; (2) cross-vein to CuA1
94. W anal/ W posterior cubital: (0) ≤; (1) ≤ 1.5x; (2) ≤ 2x; (3) > 2x
95. cup open: (0) open; (1) narrow < r-m; (2) closed at wing margin (
96. Anal lobe margin: (0) hairs; (1) some scales; (2) scales dense
97. Anal cell: (0) broad rounded; (1) rounded; (2) thin linear; (3) very reduced
98. Alula reduced: (0) not reduced; (1) reduced, L < 4x W
99. Alula margin: (0) hairs; (1) some scales; (2) scales dense
100. Squamal margin: (0) hairs; (1) some scales; (2) scales dense
101. Squama reduced: (0) not reduced; (1) reduced, L < 4x W
102. wing L: (0) ≤ 10; (1) 10-15; (2) 16-20; (3) 21-25; (4) > 25
103. wing L/W: (0) ≤ 3x; (1) long > 3x
104. L wing/ L abdomen: (0) ≥ 3x; (1) ≥ 2 x; (2) > 1.5 x; (3) >
105. Abdomen apically: (0) rounded; (1) narrowed; (2) truncate, parallel sided
106. Abdomen L/ W T2: (0) > 2.5x; (1) < 2.5x; (2) < 2x; (3) ≤ 1.5x; (4) ≤
107. Abdominal vestiture reflective: (0) bright; (1) very dull dark; (2) not reflective
108. Abdominal bristles/hair: (0) dorsally and laterally; (1) lateral and apically T7; (2) apically; (3) absent
109. Long lateral hairs > T1: (0) dense tufts; (1) some; (2) absent
110. Ma T1: (0) pointed; (1) midstyle; (2) pectinate
111. Scales: (0) absent; (1) adpressed scales; (2) upstanding scales; (3) long upstanding scale tufts
112. Male genitalia twisted: (0) no twisting, gonocoxae ventral; (1) 90°; (2) 180°, gonocoxae dorsal
113. E setae grouping: (0) not grouped; (1) medioapically; (2) lateroapically; (3) laterally
114. E setae group: (0) not grouped; (1) loose; (2) strong; (3) dense tufts
115. Epandrial spines: (0) absent; (1) short and broad; (2) long
116. E apically: (0) deeply indented; (1) concave-indented; (2) truncate; (3) convex rounded; (4) convex pointed
117. E apical flange: (0) absent; (1) < quarter base; (2) < third base; (3) < half base; (4) > half rest base
118. E medial flange: (0) absent; (1) slight; (2) distinct < quarter base
119. L E basal flange: (0) absent; (1) < quarter base; (2) < third base (
120. Mid W/L basal flange: (0) absent; (1) < quarter length base; (2) < half length; (3) > half length (4) > length
121. E posterolateral flange: (0) absent; (1) < quarter length base
122. E basal flange recurved: (0) absent (
123. E basally extended: (0) absent; (1) present
124. SES: (0) absent; (1) linear; (2) triangular; (3) quadrate; (4) single (
125. L SES/G W: (0) absent; (1) < eighth; (2) < quarter; (3) > quarter
126. G setae: (0) some; (1) dense; (2) tufts
127. G setae group: (0) absent; (1) not grouped; (2) apically; (3) medially; (4) laterally; (5) basally
128. Thick G setae number: (0) absent; (1) no thick setae; (2) some; (3) many; (4) 6-8 long
129. Thick G setae position: (0) absent; (1) no thick setae; (2) apically; (3) medially; (4) laterally; (5) basally
130. G subapical indentation: (0) absent; (1) slight < third; (2) narrowed apically > third
131. W G medial indentation: (0) absent; (1) < third; (2) > third; (3) > half
132. G medial weakness: (0) absent; (1) desclerotised line; (2) lines of weakness; (3) division medially
133. G ventral division: (0) line fusion basally; (1) line fusion entire; (2) fused medially; (3) fused basally; (4) fused
134. G medioventrally: (0) deeply indented; (1) indented medially; (2) flat; (3) convex shell
135. G ventral ridge: (0) absent; (1) slight; (2) distinct
136. G basal projection of the ventral ridge: (0) absent; (1) slight; (2) distinct; (3) recurved hook
137. G basomedial margin: (0) deeply indented; (1) indented, concave; (2) smooth, linear; (3) convex
138. H: (0) absent; (1) present
139. H laterally: (0) absent; (1) indented between G, smooth; (2) projecting; (3) with spur; (4) with finger (
140. RM: (0) small; (1) > L GS; (2) large recurved
141. L G A: (0) absent; (1) < GS; (2) > GS
142. G plates dorsoapically: (0) medially parallel; (1) angled basomedial plates diverge dorsally
143. G dorsoapical plates extension apically: (0) not extended apically; (1) small apical extension; (2) long apical extension beyond the base of the gonostyli
144. GS: (0) large base, long pointed flange; (1) laterally bifid; (2) simple curved hook; (3) medial hook; (4) lateral hook
145. GS basal projection: (0) absent; (1) small; (2) < GS
146. L AE: (0) < GS; (1) = GS; (2) > GS
147. AE EP separate: (0) absent; (1) present
148. EP: (0) absent; (1) present
149. EP deep ventral notch: (0) no EP; (1) absent; (2) medial
150. EP expanded apically: (0) no EP; (1) not expanded (
151. EP apical plate: (0) no EP; (1) absent; (2) apical plate
152. EP medioventral projection: (0) no EP; (1) absent; (2) above AE
153. EP lateroapical lobes: (0) no E; (1) absent; (2) rounded dorsally; (3) pointed dorsally
154. EP lateral projection laterally: (0) no EP; (1) absent (2) lateral
155. EP medial projection laterally: (0) no EP; (1) absent; (2) medial
156. EP pair ventral projections: (0) no EP; (1) absent; (2) medial below AE
157. L EP: (0) no EP; (1) < GS base; (2) > GS base; (3) > G margin
158. EP recurved apicomedial projection: (0) no EP; (1) absent; (2) present dorsally
159. EP recurved apically: (0) no E; (1) absent; (2) apex recurved lateral view
160. EP apical setae: (0) no EP; (1) absent; (2) present
161. BP expanded: (0) not expanded; (1) round; (2) swollen spherical; (3) bilobed
162. LAEA: (0) spoon convex up; (1) spoon concave; (2) linear; (3) absent
163. L lateral AE A: (0) < L GS; (1) < G margin; (2) = G margin; (3) absent
164. AAES: (0) spoon convex; (1) spoon concave; (2) narrow wedge; (3) linear
165. AAES: (0) < B; (1) = L GS; (2) < G margin; (3) to G margin
166. EJA: (0) racquet - round; (1) linear
167. L EJA: (0) within G; (1) = G; (2) > G < L GS; (3) > G > L GS
168. AC spines: (0) 3 prs; (1) 4 prs; (2) 5 prs; (3) > 5 prs; (4) > 10 prs
169. AC spines apically: (0) thin tapering; (1) broader spoon shaped
170. T9+10 sclerites: (0) 1 sclerite; (1) 3 sclerites
171. T9 dorsal medioapical unsclerotised lacuna: (0) absent; (1) present
172. T8 dorsal medioapical unsclerotised lacuna: (0) absent; (1) present
173. T8 hair: (0) apical half; (1) apical half bare medially; (2) apical edge
174. T8 laterally: (0) not indented; (1) indented arms
175. T8 A divided medially: (0) not divided; (1) slightly; (2) distinctly ≥ half width
176. T8 A lateral projections: (0) absent; (1) slight; (2) not linear; (3) linear
177. T8 A L/medial W: (0) margin thickened, sclerotised; (1) ≤ quarter; (2) ≤ half; (3) <; (4) ≥; (5) > 2x; (6) ≥ 3x
178. T8 A internal structure: (0) absent; (1) linear; (2) quadrate plate
179. S8 sclerites: (0) 1 linear 2 round; (1) 1 linear 2 round 1 medial; (2) U-shaped 2 triangular; (3) 2 round; (4) sheet
180. Furca: (0) U-shaped; (1) 3 separate sclerites; (2) 2 rods
181. ST tube: (0) short < third L pump; (1) present; (2) long > × 8 L pump
182. Basal endplate: (0) absent; (1) small; (2) present (
183. Basal endplate: (0) absent; (1) simple-thin processes; (2) thick processes; (3) funnel
184. Sperm pump: (0) short pump; (1) very long pump
185. Long pump papillae: (0) no long papillae; (1) unpigmented; (2) pigmented
186. Pump processes basally: (0) no processes; (1) short processes; (2) long papillae
187. Pump processes medially: (0) no processes; (1) short processes; (2) long papillae
188. Pump processes apically: (0) no processes; (1) short processes
189. Apical endplate: (0) large; (1) present; (2) small; (3) absent
190. Apical endplate: (0) absent; (1) simple- thin processes; (2) thick processes; (3) funnel; (4) double
191. SR: (0) L ≤ W; (1) L > W; (2) L > 2 W; (3) L > 4 W; (4) L > 6 W; (5) L > 8 W; (6) L > 30 W
192. ST basal sclerotised plate: (0) absent; (1) basal sclerotised plate
193. Tube ST to pump: (0) absent; (1) present; (2) long
194. SR shape: (0) round square; (1) oval; (2) pear, expanded apically; (3) long; (4) expanded basally
195. SR apically: (0) rounded blunt; (1) nipple; (2) narrowed; (3) knob
196. ST round basal bulb: (0) no round BB; (1) round BB
197. SR pigmented: (0) unpigmented; (1) pigmented; (2) basally unpigmented
198. ST long medial tube: (0) no medial tube; (1) tube medially
199. Long membranous base: (0) no long base; (1) long MB
200. ST long MB basally swollen: (0) no long base; (1) symmetrically; (2) asymmetrically
201. ST clear rings: (0) no rings; (1) clear ring; (2) long striated collar
202. SR to pump: (0) symmetrical; (1) asymmetrical
203. SR medially bent: (0) absent; (1) bent reservoir
204. SR apically bent: (0) absent; (1) tip only
205. Tubules: (0) absent; (1) present
206. SR walls: (0) thin unsclerotised; (1) thick sclerotised
207. SR walls: (0) no dimples; (1) with dimples thin unsclerotised