Revision of the Psilota Meigen, 1822 flower flies (Diptera: Syrphidae) of Australia Author Young, Andrew D. Author Skevington, Jeffrey H. Author Steenis, Wouter Van text Zootaxa 2020 2020-02-17 4737 1 1 127 journal article 24026 10.11646/zootaxa.4737.1.1 91c2f25f-409c-4319-bc7c-feb9e4210de1 1175-5326 3669490 C38FB007-98AE-4E4C-8A18-B1C75588815E Psilota cuprea Macquart, 1850 (Species plate: Fig. 17 ; Map: Fig. 55B ) Chrysogaster cupreus Macquart, 1850: 317 . Type locality: Tasmania , Australia Diagnosis: Face black, pale pollinose, with pale pile. Frons and vertex black pilose. Thorax black. Thoracic pile white, or mixed black and white. Hind leg simple. Calypter variable in colour. Halter variable in colour. Abdominal tergites 2 to 4 mostly to entirely red. Tergite pile variable in colour. Body length: 6.8–11.4 mm . FIGURE 17. P . cuprea Macquart. A ) male dorsal habitus, B) male anterolateral face, C) male lateral habitus. Scale bar =1 mm. Description: MALE: Head : Head black, face white pollinose. Face white, frons and vertex black pilose. Antenna orange to dark brown. Postpedicel variable in shape. Eye densely brown pilose. Thorax : Thorax black. Legs entirely orange to entirely dark brown. Hind femur simple. Hind tibia simple. Pleuron lightly white pollinose or bare and shining. Pile pleuron entirely white or with mixed black and white pile. Pile on scutum and scutellum entirely white or with mixe black and white pile. Wing colourless. Wing microtrichia variable. Alula completely microtrichose. Halter variable in colour. Calypter variable in colour. Abdomen : Tergite 1 black. Tergite 2 entirely red or with a large, anterior black area. Tergite 3 entirely red or with a small medial black spot. Tergite 4 entirely red. Pilosity of tergites variable in colour. Sternites 2 to 4 reddish, with variably coloured pilosity. Genitalia : variable (see below). FEMALE: Identical except for normal sexual dimorphism. Specimens examined: AUSTRALIA . Australian Capital Territory : Black Mountain, -35.273675 149.097583 , 21.x.1966 ; 23.x.1969 , I.F.B. Common, light trap, ANIC29034958; ANIC29034939; CNC_ Diptera 253814 ( 3♀♀ , ANIC); Blundell’s [Cabin], -35.292886 149.141800 , 20.i.1935 ; 11.x.1950 ; 24.x.1950 , S.J. Paramonov; Fuller, ANIC29034941; ANIC29034951; ANIC29034953-29034954; CNC_ Diptera 253704; CNC_ Diptera 253804 ( 2♂♂ , 3♀♀ , ANIC); Blundells Ck., -35.220000 148.500000 , ii.1987 , D.H. Colless, Malaise trap, CNC_Dip- tera253794 ( 1♀ , ANIC); Canberra—CSIRO Botanical Garden, -35.278878 149.108917 , 26.x.1988 , F.C. Thomp- son, USNM_ENT_00034137; USNM_ENT_00034139-00034141 ( 4♀♀ , USNM); Canberra, Terr. 1, -35.282000 149.128686 , 20-30.xi.1953 , M. Hull, CNC_ Diptera 254972 ( 1♂ , CNC); Canberra, -35.282000 149.128686 , 2.xi.1947 , S.J. Paramonov, ANIC29034940 ( 1♂ , ANIC); Lees Creek Brindabella Rge., -35.338461 148.867925 , 1.i.1975 , G. Daniels, GDCBReg_19737 ( 1♀ , GDCB); Mount Ainslie, -35.266667 149.166667 , 28.xi.1996 , C.J. Burwell, QM_REG._NO._T223209 ( 1♂ , QM); New South Wales : ‘Lugen Park’ Nullo Mountain Northeast of Rylstone, -32.649278 150.217650 , 22.x.1953 , T.G. Campbell, ANIC29034942 ( 1♂ , ANIC); 20 miles South of Mil- ton, -35.319178 150.439158 , 30.ix.1952 , S.J. Paramonov, ANIC29034943; ANIC29034959; CNC_ Diptera 253706; CNC_ Diptera 253815 ( 4♀♀ , ANIC); 3 km North of Lansdowne via Taree, -31.758753 152.534258 , on house steps, wet sclerophyll forest, rainforest margin, 25.xii.1988 , G. Williams, AustralianMuseumK410625 ( 1♀ , AM); 3 km South of Mount Wilson, Blue Mountains, -33.516011 150.378642 , 29.xi.1984 , D.K. McAlpine, B.J. Day, AustralianMuseumK407964 ( 1♂ , AM; 4 miles North of Bateman’s Bay, -35.658456 150.177744 , 14.x.1952 ; 12.x.1956 ; 22.x.1952 ,S.J. Paramonov,ANIC29034956;ANIC29034950; CNC_ Diptera 253705; CNC_ Diptera 253709 ( 1♂ 3♀♀ , ANIC); 4 mls W. of Amosfield, -28.658686 152.016650 , 17.x.1971 , E.M. Exley, UQIC_Reg94770 ( 1♀ , QM); 6 km Northeast Bilpin, Blue Mountains, -33.462750 150.572836 , 23.ix.1992 , N.W. Rodd, AustralianMuseumK410278 ( 1♀ , AM); Back Yamma State Forest, -33.339906 148.223703 , 11.xi.1964 , D.H. Colless, CNC_ Diptera 253708 ( 1♀ , ANIC); Bald Rock N.P., -28.8525 152.055556 ; -28.852500 152.055558 , on Leptospermum minutifolium blossom, 6-7.xi.1984 , D.K. Yeates, UQIC_Reg94630; UQIC_Reg94631-94633; UQIC_Reg94637-94638; UQIC_Reg94744- UQIC94745; UQIC_Reg94675 ( 4♂ m, # 5♀♀ , QM); Bawley Point, -35.500000 150.400000 , 21-23.ix.2000 , Rentz, CNC_ Diptera 253696 ( 1♂ , ANIC); Bilpin, Blue Mountains, -33.500259 150.532989 , 21.ix.1978 , N.W. Rodd, Aus- tralianMuseumK410429 ( 1♀ , AM); Blackheath, -33.635539 150.283625 , 24.xii.1921 , CNC_ Diptera 253806 ( 1♀ , ANIC); Boonoo Boonoo Falls, -28.801247 152.165061 , 12.xi.1981 , G. & A. Daniels, GDCBReg_19687 ( 1♀ , GDCB); Cherrybrook, electricity tower access track, end of Trevor’s Lane, -33.710139 151.059500 , 13.ix.2008 , S. Ginn, AustralianMuseumK410146 ( 1♂ , AM); Clyde Mountain, eastern foothills, -35.546072 149.950333 , 1219m , ex brackett fungus: Stereum semilugens , 15.viii.1967 , Z. Liepa, Emerged 25.VIII.1967 , ANIC29034945; ANIC29034952; CNC_ Diptera 253818 ( 3♂♂ , ANIC); Cowan, -33.590714 151.169561 , 7.xii.1931 , F.J. Gay, ANIC29034961 ( 1♀ , ANIC); Dobroyd Point, Sydney, -33.926375 151.125936 , 27.xi.1978 , N.W. Rodd, Australian- MuseumK410619 ( 1♀ , AM); Durras Bay, -35.645003 150.291439 , 12.xi.1953 , G. Daniels, CNC_ Diptera 253846 ( 1♀ , CNC); Durras, -35.646775 150.296939 , 29.ix.1959 , Z. Liepa, ANIC29034969 ( 1♀ , ANIC); Gibraltar Range National Park, Waratah track, -29.5 152.333333, on Leptospermum sp. Blossom, 10-12.xi.1984 , D.K. Yeates, UQIC_Reg94634 ( 1♀ , QM); Goondera Ridge, Royal National Park, -34.114342 151.046586 , 2.x.1978 , G. Daniels, GDCBReg_19533; GDCBReg_19534 ( 2♂♂ , GDCB); 22.viii.1976 , G. and A. Daniels, GDCBReg_19535 ( 1♂ , GDCB); Heathcote NP, Kingfisher Pool, -34.117042 150.952122 , dry eucalypt forest, 16.x.1994 , B.J.Sinclair, CNC_ Diptera 106594 ( 1♀ , CNC); Jervis Bay, -35.035431 150.668869 , 4.ix.1948 , E.F. Riek, ANIC29034949 ( 1♂ , ANIC); Killara, -33.768639 151.163469 , 13.x.1934 , D.F.W., ANIC29034955 ( 1♀ , ANIC); Lorien Wildlife Refuge, 3 km North of Lansdowne, -31.758753 152.534258 , 7.xi.1993 , G. & T. Williams, AustralianMuseumK411046 ( 1♂ , AM); Mooney Mooney Creek, near Gosford, -33.437894 151.246467 , 1.xii.1989 , D.K. McAlpine, B.J. Day, AustralianMuseumK408009 ( 1♀ , AM); Mount Canobolas State Recreation Area, summit Mount Canobolas, 12.8 km Southwest of Orange, -33.343889 148.982500 , 1395m , 18.x.2002 , C. Lambkin, N. Starick, hand collected, ANIC29035762 ( 1♂ , ANIC); Mount Gibralter, Comboyne Plateau, -31.644328 152.422989 , shrub complex, 21.i.1985 , G. Williams, AustralianMuseumK410621 ( 1♀ , AM); Mount Tomah, Blue Mountains, -33.549614 150.416033 , 26.ix.1978 ; 26.x.1978 ; 28.xii.1985 , N.W. Rodd, N.W. Rodd Collection, AustralianMuseumK410430; AustralianMuseumK410252; AustralianMuseumK410154; AustralianMuseumK410163; AustralianMuseumK410175; AustralianMuseumK410182 ( 2♂♂ , 3♀♀ , AM); Mt Canobolas State Recreation Area, summit Mt Canobolas 12.8 km SW Orange, -33.343889 148.982500 , 1395m , 18.x.2002 , C.L. Lambkin, N.T. Starick, hand net, CNC_ Diptera 253847 ( 1♀ , ANIC); Nadgee NR, near Eden, -37.399378 149.908592 , wet sclerophyll forest, 21.xii.1985 ; 14-15.ii.1986 , E.A. Sugden; D.J. Bickel, AustralianMuseumK410515 ( 1♀ , AM); Nadgee Reserve 4.7km S Merrica Hut, -37.399378 149.908592 , 21.xii.1985 ; 21.xii.1986 , E.A. Sugden, CNC_ Diptera 254983; Jeff_ Skevington_Specimen43807; Jeff_Skevington_Specimen43810-43811 ( 1♂ , 3♀♀ , CNC; QM); New England Nat. Pk. Via Ebor, -30.476022 152.660183 , 22–23.i.1966 , B. Cantrell, UQIC_Reg94705 UQIC_Reg94636; UQIC_ Reg94635 ( 3♀♀ , QM); New England Nat. Pk. near Wright’s Lookout, -30.476022 152.660183 , 14.x.1961 ; 29– 31.x.2002 , C.W. Frazier; G.,M.Wood, CNC_ Diptera 253803; CNC_ Diptera 106427 ( 2♀ , ANIC; CNC); Pigeon House Mtn., Ulladulla, -35.349175 150.265064 , 13.xi.1976 , G. Daniels, GDCBReg_19767 ( 1♀ , GDCB); Royal National Park, Flat Rock Creek, near Sydney, -33.926375 151.125936 , From the rotten caudex of Xanthorrhoea resinosa , 10.viii.1973 , D.K. McAlpine, S.P. Kim, emergence on 4.IX.1973 , AustralianMuseumK410255 ( 1♀ , AM); Rutherford Ck Brown Mtn. nr. Nimmitabel, -36.601300 149.381311 , 26.xii.1974 , G. Daniels, GDCBReg_19536 ( 1♂ , GDCB); Rutherford Creek, Brown Mtn. nr Nimmitabel, -36.581667 149.551111 , 14.xi.1976 , G. Daniels, GDCBReg_19684 ( 1♀ , GDCB); Siding Springs Observatory, Summit, -31.255558 149.061111 , 31°15’20’’S 149°3’40’’E , 17.x.1997 , J. Skevington, C. Lambkin, S. Winterton, UQIC_Reg94532–94536 ( 5♂♂ , QM); Stanwell Tops, -34.219903 150.991808 , 9.x.1977 , G. Daniels, GDCBReg_19528–19531 ( 4♂ , GDCB); Tindery, -35.754172 149.280764 , 1.i.1938 , E.H. Kipps, ANIC29034968 ( 1♀ , ANIC); Wahroonga, Cook Trig, trail off Grosvenor Street, -33.699450 151.126500 , 28.ix.2007 , D. Britton, S. Ginn, L. Holme, LHP002/01, AustralianMuseumK410478 ( 1♂ , AM); Warrumbungle National Park Area, -31.272222 149.061111 ; -31.279689 149.046797 , observatory hilltop; Observatory hilltop, 17.x.1997 , J. Skevington, UQIC_Reg94543–94545 ( 3♂ , QM); Waste Point, Kosciusko Na- tional Park, -36.427114 148.309022 , 24.i.1984 , David W. Inouye, AustralianMuseumK410600 ( 1♀ , AM); West of Mount Mackenzie, 9 km West of Tenterfield, -29.066667 151.966667 , paddock with bracken fern, 9.xii.2003 , G. Daniels, AustralianMuseumK411060 ( 1♀ , AM); Woolgoolga, -30.113036 153.183792 , 28.i.1923 , Health Dept., CNC_ Diptera 253805 ( 1♀ , ANIC); Woy woy, -33.489839 151.319942 , 3.x.1925 , Mackerras, CNC_ Diptera 253809 ( 1♀ , ANIC); Queensland : 7 km North of Maryborough, -25.470267 152.695200 , 23.viii.1977 , K.J. Lamkin, QM_ REG._NO._T223216 ( 1♀ , QM); Agnes Water, -33.710139 151.059500 , 3.ix.1987 , N.W. Rodd, AustralianMu- seumK410174 ( 1♂ , AM); Bald Rock National Park, Boundary Swamp Creek, -28.816667 152.066667 , Veg. over water, 11.x.1998 , J. Skevington, sweeping, UQIC_Reg94746 ( 1♂ , QM); Bluff Range, West Slope, Biggenden, - 25.600000 152.050000 , 21.viii.1975 , H. Frauca, ANIC29034946; CNC_ Diptera 253816 ( 2♀ , ANIC); Brisbane, - 27.526975 153.040322 , 1911, H. Hacker, QM_REG._NO._T223208 ( 1♂ , QM); x.1925 , G.H. Hardy, Australian- MuseumK407091 ( 1♀ , AM); Brown Lake, N Stradbroke Is., S.E.Q., -27.490105 153.432379 , on Leptospermum flavescens blossom, 21–24.ix.1984 , D.K. Yeates, UQIC_Reg94674 ( 1♂ , QM); Bundaberg, -24.866972 152.350972 , viii-ix.1971, H. Frauca, ANIC29034970 ( 1♀ , ANIC); Collaroy, Sydney, -33.926375 151.125936 , 1921, Health Dept. Date is ambiguous reads ”2.9.21”, ANIC29034947 ( 1♂ , ANIC); Dayboro, 0.5 km South, -27.200000 152.816667 , 18–19.ix.1999 , S.G. Evans, QM_REG._NO._T223227 ( 1♀ , QM); Flinders Peak nr. Boonah, - 27.800000 152.816667 , summit, 13.ix.1995 , D.K. Yeates, UQIC_Reg94538-94539 ( 2♂♂ , QM); Flinders Peak, via Peak Crossing, -27.816667 152.816667 , 13.ix.1995 , C.J. Burwell, D.K. Yeates, QM_REG._NO._T223211-223215 ( 5♂♂ , QM); Girraween National Park, Hilltop 1.2km NE of Bald Rock Campground, -28.821942 151.937850 , 31.xii.2014 ; 1.i.2015 , J.H. Skevington, CNC384663-384670; CNC384926-384927; CNC384624-38430; CNC451705–451706; ( 20♂♂ , CNC); Kenniff’s Lookout, Mt. Moffatt sect., Carnarvon Nat. Pk., -25.165167 147.840419 , 25.ix.1986 , D.K. Yeates, UQIC_Reg94624 ( 1♂ , QM); Lamington N.P., -28.045967 153.127983 , 29.x.1955 , F.A. Perkins, UQIC_Reg94905 ( 1♀ , QM); Miriam Vale, -24.330397 151.557592 , on Angrophora costata , 5.xii.1977 , T. Sears, UQIC_Reg94623 ( 1♂ , QM); Mount Marlay Summit, Stanthorpe, -28.666667 151.933333 , 20.x.1997 ; 22.x.2005 , J. Skevington, C. Lambkin, S. Winterton; D. K. Yeates, UQIC_Reg94539–94541; ANIC29035763–29035764 ( 5♂♂ , QM); -28.653583 151.946906 ; -28.653583 151.946908 , on Leptospermum , 22.ix.1985 , D.K. Yeates, UQIC_Reg94626;, UQIC_Reg94639; UQIC_Reg94668; UQIC_Reg94735 ( 4♂♂ , QM); Mt. Beerburrum, Glasshouse Mts., -26.950000 152.933333 , 280m , 29.x.1991 , R. Eastwood, GDCBReg_19735 ( 1♂ , QM); Mt. Norman Area, Via Wallangarra, -28.873267 151.973822 , 2.x.1971 , G.B. Monteith, UQIC_Reg94769 ( 1♀ , QM); Near Rosedale, -24.631128 151.913664 , 27.x.1974 , H. Frauca, ANIC29034948; ANIC29034965 ( 1♂ , 1♀ , ANIC); Six Mile Creek, W of Elimbah, -27.016667 152.933333 , 15.ix.1985 , G. and A. Daniels, GDCBReg_ 19569 ( 1♀ , QM); Stanthorpe, -28.657928 151.930344 , 22.ix.1979 , S.J. Johnson, GDCBReg_19532 ( 1♂ , GDCB); Summit of Mount Greville, -28.074028 152.502008 , 393m , 3.i.2015 , J.H., A.M. and A.W. Skevington, CNC451749 ( 1♂ , CNC); Tambourine Mountain, -27.923086 153.171717 , 28.x.1912 , H. Hacker, QM_REG._NO._T223184 ( 1♀ , QM); The Blunder, Brisbane, -27.526975 153.040322 , 9.x.1966 ; 18.ix.1966 ; 25–26.ix.1966 , C.F. Ashby, ANIC29034944; ANIC29034964; ANIC29034966; CNC_ Diptera 253707 ( 1♂ , 3♀♀ , ANIC); Woombye, near Nam- bour, -26.666911 152.966606 , 11–16.x.1965 , D.H. Colless, ANIC29034963 ( 1♀ , ANIC); along Rd to 18 Mile Swamp, N. Stradbroke Island, -27.516667 153.483333 , 17.ix.1989 , M.A. Schneider, UQIC_Reg94771 ( 1♀ , QM); South Australia : Adelaide Area, Mount Lofty, -34.984972 138.708333 , 4.x.1997 , J. Skevington, CNC_ Diptera 254973 ( 1♂ , QM); Adelaide, Belair National Park, -35.000000 138.633333 , Hilltopping, 4.x.1997 , J. & A. Skevington, C. Lambkin, S. Winterton, UQIC_Reg94522–94531 ( 8♂♂ , 2♀♀ , QM); Dutchmans Stern Conserva- tion Park, -32.296944 137.983058 , summit, 7.x.1997 , S. Winterton, J.&A. Skevington, C. Lambkin, UQIC_ Reg94537 ( 1♂ , QM); Para Wirra Recreational Park, -34.701739 138.824928 , 8.x.2014 , A. Marquart, SLAM trap, CNC587054 ( 1♀ , CNC); Sandy Creek Conservation Park, 8km NE Gawler, -34.603333 138.855833 , 29.ix–4.x.1997 , S. Winterton, J.&A. Skevington, C. Lambkin, Malaise trap, malaise in heath, UQIC_Reg94542 ( 1♂ , QM); South Para Reserve, -34.676314 138.876983 , 20.x.2015 , A. Marquart, SLAM trap, CNC587060 ( 1♀ , CNC); Tasmania : 1km SEE Gladstone, -40.580000 148.010000 , 29.i.1963 , I.D. Naumann & J.C. Cardale, CNC_ Diptera 253802 ( 1♀ , ANIC); 9 km SE Miena, -42.000000 146.766667 , 1060m , 11.i.1992 , G. and A. Daniels, GDCBReg_19616 ( 1♀ , QM); Eaglehawk Neck, -43.019722 147.925000 , 24.iv.1916 , G.H. Hardy, 12, CNC_ Diptera 253801 ( 1♀ , ANIC); Erriba, -41.425350 146.137136 , 30.i.1949 , E.F. Riek, ANIC29034960; ANIC29034967; CNC_ Diptera 253710 ( 3♀♀ , ANIC); Franklin-Gordon Wild River National Park, Donaghy’s Hill, -42.197778 145.931944 , 10–11.i.2015 , J.H., A.M. and A.W. Skevington, CNC385297; CNC385305; CNC385318; CNC451889–451892; CNC451894; CNC457453 ( 5♂♂ , 4♀♀ , CNC); Hobart, -42.882139 147.327194 , 16.xii.1917 , G.H. Hardy, 45, CNC_Dip- tera253800 ( 1♀ , ANIC); Lyell Highway, Frankin R nature trail, -42.215700 146.019800 , 11.i.2015 , S.J. Grove, hand collected, TMAG-F11616 ( 1♂ , TMAG); N Bruny Island; Waterview Hill, -43.093056 147.351111 , E Hilltop, 31.xii.1998 , J. & A. Skevington, M. Mathieson, UQIC_Reg94797 ( 1♂ , QM); North of Corinna, -41.588514 145.189764 , 12.i.2015 , J.H., A.M. and A.W. Skevington, CNC457433 ( 1♀ , CNC); Oonah, -41.224222 145.618608 , 30.i.1949 , E.F. Riek, ANIC29034971 ( 1♀ , ANIC); Pump house Bay Camping Ground Arthurs Lake, -41.983333 146.866667 , 12.ii.1992 , D.S. Horning, Jr., in flight, CNC_ Diptera 253698 ( 1♀ , CNC); Samford, 1959, C.F. Ashby, Date is ambiguous reads ”6.12.59”, ANIC29034972 ( 1♀ , ANIC); Warra Long Term Ecological Research Site 147, -43.1 146.683333, 17.xi–16.xii.1998 , malaise trap, UQIC_Reg94607–94608 ( 2♀♀ , QM); Victoria : Ararat, - 37.284372 142.932553 , H.W. Davey, CNC_ Diptera 253811 ( 1♀ , ANIC); Grampians National Park, Reed’s Look- out, -37.148111 142.448261 , 23.x.2014 , J.H., A.M. and A.W. Skevington, CNC384097; CNC457381 ( 2♂♂ , CNC); Heathmont, -37.829989 145.242019 , CNC_ Diptera 253778 ( 1♀ , CNC); Mt. Buffalo N.P., Dicksons Falls, -36.763417 146.788092 , 1440m , 29.i.2006 , Jeff_Skevington_Specimen43805–43806 ( 2♂♂ , QM); Yarra Ranges N.P., Mt. Don- na Buang track, -37.709467 145.676642 , 4.ii.2006 , M.P. van Zuijen, Jeff_Skevington_Specimen43808 ( 1♀ , QM); Western Australia : 2 km North of Pemberton, -34.426158 116.039058 , 29.ix.1987 , D.K. McAlpine, AustralianMuseumK407981 ( 1♀ , AM); Geraldton, -28.777372 114.614972 , ix.1926 , E.W. Ferguson, CNC_ Diptera 253812 ( 1♀ , ANIC); Barrington, Allyn Range, -41.356572 146.279756 , Leptospermum , ii.1925 , Label unclear: ”Tps Feb ‘25 SU ZooExp”, CNC_ Diptera 253810 ( 1♀ , ANIC); Mt. Wellington, -42.890000 147.230000 , 2.xii.2016 , G.H. Hardy, Date ambiguous: ”2.12.16 /48”, CNC_ Diptera 253798 ( 1♂ , ANIC). Discussion: Psilota cuprea doubtlessly encompasses multiple species as described here. Initial DNA barcode analysis indicates that at least nine species are represented ( Figure 43 ). Further DNA barcoding and male genitalia dissection are required to adequately separate the full range of specimens into species and will be tackled in a future revision. Furthermore, there are several other species of Psilota described from Australia that may or may not be synonyms of P. cuprea , including P. erythrogaster , P. rubra , and P. rubriventris (See ‘species not here recognized’ below). Because it is impossible to tell what name should properly be assigned to different barcode clusters without a full revision of this complex, these species have not been treated individually.As mentioned in the introduction, an individual from this complex was reared from the “rotten caudex of Xanthorrhoea resinosa ”. The holotype of this species was not available for study.