Physiphora alceae Preyssler 1791
Creators: name: Elena P. Kameneva
name: Valery A. Kroneyev
Publisher: name: Zenodo
Year: 2016
Subjects: Biodiversity
Taxonomy
Animalia
Arthropoda
Insecta
Diptera
Ulidiidae
Physiphora
Physiphora alceae

description: Physiphora alceae (Preyssler 1791)
Figures 28–40
Musca alceae Preyssler, 1791: 129.
Physiphora alceae: Rozkošný, Chvála & Pont, 1982: 353; Zuska, 1997: 70; Kameneva, 2000: 156; 2001: 156; Chen & Kameneva, 2007: 17; Kameneva & Korneyev, 2010: 622; Kameneva & Greve, 2013.
Musca demandata Fabricius, 1798: 564.
Tephritis demandata Fabricius, 1805: 324.
Ulidia demandata: Meigen, 1826: 386; Macquart, 1835: 504.
Chloria demandata: Schiner, 1864: 386.
Chrysomyza demandata: Becker, 1905: 106; Hendel, 1910: 4, 21; 1913b: 217; Lamb, 1914: 315; Séguy, 1934: 84.
Physiphora demandata: Hennig, 1940: 11; Steyskal, 1977: 167; 1980: 576; Zaitzev, 1984: 60; Evenhuis, 1989: 481.
Chrysomyza splendida Fallén, 1817: 4.
Ulidia bicolor Schiner, 1864: 86; Séguy, 1934: 84; Zaitzev, 1984: 60; attributed to Macquart, 1835: 504 (Macquart’s “U. bicolore” was used clearly as a vernacular French name of Ulidia demandata; consequently mentioned “ U. bicolor Macquart ”, always as a synonym of demandata, is unavailable).
Ulidia smaragdi Walker, 1849: 1059.
Material. Non-type. Afghanistan: “ O. Afghanistan, Sarobi 1100 m ”, 13. viii.1981, 1 ♂ (Ebert) (ZSSM); Polichomri 700 m, 28. v.1956, 2 ♀, 8. vi.1956, 1 ♀ (Amsel) (SMNK); Armenia: “Kotaih prope Eilar”, 4. vii.1926, 1 ♀; slope of Alagöz mt., Piragan, 4800', 16. vi.1924, 1 ♂ (A. Shelkovnikov); prope Erivan, 13. vii.1924, 2 ♂ (S. Paramonov), idem, 16. vi.1926, 1 ♀ (A. Shelkovnikov), idem, 22. v.1927, 4 ♂, 2♀; Arazdajan [39.7448802N 44.8246479E], 26. vi.1933, 1 ♀ (S. Paramonov) (SIZK); Austria: “Wien / 7.[18]40”, 3♂ ♀; (with black-and-gray paper label) 2♂, 1♀ (Coll. Loew) (MNKB); Admont, 1♀ (MTD); Azerbaijan: Lenkoran, 31.vii–1. viii.1900, 2 ♀: (F.Karsch) (MNKB); Ordubad, 29. v.1924, 1 ♀ (S. Paramonov) (SIZK); Botswana: “bei Maun, Krokodilfarm”, 21. ii.1998, 1 ♂ (Göllner) (MNKB); Bulgaria: Kiten, oak wood + pig farm, 41.14N 27.48E, 17. vii.1987, 1 ♂, 1♀; Sliven, 5km N, 42.45N 26.17E, 21. vi.1987, 1 ♂, 1♀ (M. Barták) (MBC); China: Xinjiang, Kashgar, 15.10.1991, 1♀ (W.Mey) (MNKB); “Schansi”, 1♂ (MTD); “Tchangk.tch” [Great Wall of China], 16. vi.1916, 1 ♂ (Licent); Shanxi: Hongtong, 14. vii.1916, 1 ♂, 1♀ (Licent) (MHNP); Croatia: Javornik, 1423 m, 10. vi.1910, 1 ♀ (Meusel); Cmi Padez, 423 m, 22. ix.1910, 1 ♀ (Meusel); Dalmatien, 14.v. “221891”, 1♂; Draga 18.vi.1887 1 ex.; St.Pasova, 11. viii.1887, 1 ♀; Spitzberg, 17.viii.1926, “061”, 1♂; Zagreb, 3.ix.1886, 18. ix.1887, 2 ♂ (MNKB); Rovinj, 28. v.1961, 1 ♀ (Ulrich) (ZFIB); Czech Republic: Praha: Šárka, corn field, 50.06N 11.19E, 300m, 12. ix.1985, 1 ♂; Prokopské údolí, fly trap, 50.02N 11.23E, 27.iv–7. v.1987, 1 ♂; Kunice—garden, Malaise trap, 49.56N 14.40E, 430m, 17–19.viii.1985, idem, cheese trap, 25.vii–13. viii.1985, 2 ♂, 2♀ (M. Barták); Nová Rabyně, 49.49N 14.26E, 350m, 17. x.1988, 1 ♀; Budyné—Ochří, 3km W, damp meadow, 50.24N 14.06E, 190m, 27. vi.1987, 1 ♂; Jíloviště— 2km SW near brook, 49.56N 14.19E, 300 m, 22. ix.1983, 1 ♂; Karlštěin env., deciduous forest, 49.57N 14.11E, 350 m, 28. vii.1985, 1 ♂; U. Janovice— 5km W, near pond, 49.53N 15.03E, 460 m, 19. ix.1991, 2 ♂, 1♀; idem, damp meadow, 5–12. x.1995, 1 ♂ (M. Barták) (MBC); Egypt: “ Aegypten / Ambukohl”, 1♀ (Ehrenlarg); [green square label], Coll. H. Loew, 338, 1♀; “Alexandrien / 44154.XI”, 2♂, 2♀; “Kairo XI / 44296”, 1♀ (MNKB); Fayed, 17. v.1943, 1 ♀ (Priesner) (ZSSM); “ Aegypt ”, xii.1903, 1 ♀ (MTD); Kafr-el-Sheikh, garden vegetation, 30.48N 31.14E, 28. iii.1995, 1 ♂; Cairo, El-Marg, margin of field, 30.16N 31.23E, 29. iii.1995, 1 ♂; Cairo, 20 km S, semidesert, 29.52N 31.17E, 31. iii.1995, 2 ♂, 2♀ (M. Barták) (MBC); France: Corsica, 24. vi.1899, 1 ♂ (Ajaccio); Bautzen, 2♂, 2♀; Digne, vi.1908, 1 ♂; “Franzn s bd”, 2♂, 1♀ (Kowarz); (MTD); Mont Dophin, pine wood, 44.41N 6.37E, 11. vii.1990, 1 ♂ (M. Barták) (MBC); “Villefranche”, 2♂ (DEI); Gambia: 3km NW Central Banjul garden at Wadner Beach Hotel, Loc. No 1A, 19:00–21:00, 21. ii.1977, 15 ♂, 10♀; at light in vegetation along mangrove swamp, loc. No. 2, 21–22. ii.1977, 5 ♂, 3♀; idem, loc. No 2, 23. ii.1977, 2 ♂, 1♀, idem, loc. No 2, 25. ii.1997, 1 ♀ (Cederholm, Danielsson, Larsson, Mireström, Norling & Samuelsson) (ZMLU); Georgia: “Tiflis” [Tbilisi], 1♂ (DEI); Germany: Berlin, 21 ex. with different data; [yellowish paper square], 2♀ (Coll. H.Loew) (MNKB); Erfurt, 1928, 3 ♂, 3♀ (v. Roeder S.); Berlin, Umgbg, 2♂; Dresden, 2♀; Gr. Garten, 1♀; Leipzig, 1♀; Bromberg, 9. viii.1915, 1 ♂, 1♀ (Meyer); Lössnitz, ix.1985, 1 ♂ (MNKB); Greece: Korfu, Zitadell, Gebüsch, 6. v.1958, 2 ♀ (Boness) (ZFIB); Gr. Insel Lesbos, NW, Ayia Paraskevi, Kulturland, Quercus, 19. v.1996, 1 ♂ (Grimm) (SMNS); Hungary: Kenzel, 1♂ (Thalhammer) (MTD); Zákániszék, 20 km W Szeged, 26. vii.1993, 1 ♂, 1♀ Bekes, Komadi, 29. vii.1993, 1 ♂, 3♀ (D. & T. Osten) (SMNS); India: South India, Bangalore, 15. iv.1936, 31 ♂♀ (Nathan) (BINH); Bombay, 26. x.1901, 1 ♂ (Uzel) (NHMW); Iran: Shemiran, viii.1959, 4 ♂, 3♀ (Barnett) (USNM); Iraq: Basra, 20, 21. x.1954, 2 ♂, 1♀ (Lindemann) (ZSSM); “ Mesopotamia ”, 1917, 5 ♂, (D’Beaufort) (BMNH: Wellcome coll., 1990–10); Baghdad, Hotel Babylon, garden, light trap, 1. v.1988, 1 ♀ (Olejnícek) (MBC); Israel: Jericho, 30. v.2000, 1 ♀ (Merz) (NHMG); Rehoboth bei Jaffa, 27.vii.1931 – 24.v.1935, 40♂ ♀ (Aharoni); Nahatal, 26. iii.1935, 1 ♂ (Aharoni); Jerusalem, Scopusberg, 1♂ (Aharoni); Karmel Mts., 10 km S Haifa, 14. v.1996, 1 ♂ (Schmid-Egger) (SMNS); Italy: Sicily: “Syrakus”, 21.v, 7♂ ♀; “Syrakus 15.6” 1♂; “Bouchi”, 1♂: (MNKB); Italy: Venezia—Lido, 18. viii.1932, 1 ♀; Bozen 13. vi.1890, 5 ♂; “Coll. / Gerst.” 2♂, 3♀; “N Italien Garda, See Gargnana”, 21. ix.1923, 1 ♂, 2♀ (G. Enderlein) (MNKB); Ferrara, along river, 44.54N 11.38.E, 7. viii.1988, 1 ♂; Castiglione D.Or, ca. along river, 43.00N 11.37E, 6. viii.1988, 1 ♀ (M. Barták) (MBC); Jordan: Irbid Adm., Unit, Qaren, ca. 18 km, N Dier Alla, 17. vi.1987, 7 ♂♀ (Zack) (USNM); Kyrgyzstan: Kazarman, 26. vi.1964, 1 ♀ (L. Peck); Chon-Aryk prope Bishkek [42.7892282N 74.5747232E], 24.07.1994, 1♂ (D. Milko) (SIZK); Latvia: “Kurland, Libau” [=Liepaja], 1♀ (DEI); Lebanon: Ksara am Libanon, 2. vi.1920, 1 ♀ (Aharoni) (SMNS);♂: Beyrut (MNKB); Madagascar: Majunga, i–ii.[18]93, 1 ♂: (Voeltrkow) (MNKB); Behara, iv.1937, 1 ♀ (A. Seyrig) (MRAC); Tanatan, iv.1930, 3 ♀ (A. Seyrig); de Fénérive a Soanierara, 1927, 2 ♂, 2♀ (G. Petit) (MHNP); Malta: Mellieha Bay, Gadira, 19– 31. viii.1998, 1 ♂ (B. Petersen) (ZMUC); Moldova: Chisinău, Institute of Biocontrol, on window, 12.ix.1987; idem [46.9663137N 28.8820267E], 13. vii.1987, 2 ♂ (V. Korneyev) (SIZK); Balabanesti nr. Vadul-lui-Vodă [47.0401821N 29.1376305E], 22–26. vii.1988, 1 ♂ (V. Korneyev) (SIZK). Morocco: Anti-Atlas, Tizi n'Bachkoun, 1600 m, 1. vii.1987, 1 ♂ (Schacht) (ZSSM); Casablanca / Quedenfieldt (MNKB); Tanger, 1♂ (DEI); Nepal: Katmandu, 1400 m, 14. iv.1962, 1 ♂ (Ebert, Falkner) (ZSSM); Oman: Al Misfa desert, 6. iv.1985, 1 ♀ (FAÖ) (ZMLU); Pakistan: Peshawar, N.W. Frontier Prov., vi.1959, 4 ♂, 4♀ (Barnett) (USNM); Poland: “Gleiwits” [Gliwice], “6.8. 317”, 1♂, 7♀; 3♂, 2♀ (Coll. Loew) (MNKB); Portugal: “ Portugal ”, 1♀ (Hoffmannsegg); “Lusitan / Hoffsg.”, 1♂ (MNKB); Romania: “Bucarest”, 2♂ (A. C. Montandon) (ZSSM); Russia: Stavropol: Spakovskoye, 4. viii.1988, 1 ♂ (V. Korneyev) (SIZK); Volgograd: “Sarepta” [Volgograd], 3♂, 1♀ (MNKB); Samara, 1♂; Orenburg, vi.1909, 1 ♂ (MTD); Saudi Arabia: Er Riad, 22.v.1957, 14.x.1957, 13.iii.1959, 25. v.1959, 3 ♂, 2♀ (Diehl) (ZSSM); Riad, 700 m, 28. ii.1958, 1 ♀ (Diehl) (SMNK); Riyadh, R. Abo-zohayrah, sp. 3, in stems of Phoenix dactylifera [date palm], 9.xi.1986 (CIE 18484) (BMNH); Seychelles: Farcoire, sea shore, on fish, 16. viii.1984, 1 ♂, 3♀ (no collector) (SIZK); Slovakia: Velké Leváre — 3km N, near brook, 48.32N 16.59E, 27. vii.1986, 2 ♂ (M. Barták) (MBC); South Africa: W Cape, Cape Town, 33º56′S 18º28′E, larvae infesting growth tip of date palm, 19. v.2008, 1 ♂ (dissected), 4♀ (G. Tribe) (SANC; SIZK); Free State Bloemfontain; Franklin Game Reserve, Naval Hill, radio tower, 26º06.277′S 26º13.852′E, sweeping, 20. xii.2008, 1 ♂ (A. Kirk-Spriggs) (BMSA); Spain: Elche, v.1933, 1 ♀; Sierra Espuna, 1200 m, 1♂, 1♀: v.1933 (Hering) (MNKB); Rio Big. Del Sangua, 6. v.1955, 1 ♀ (Mannheims) (ZFIB); E. Prov. Salamanca, Castillejo, 17. ix.1986, 1 ♀ (Viejo) (SMNS); Canary Is.: Teneriffe, Las Nercedes, a. Euphorbien, 3.vii.1926, 41 specimens (Liebe) (MNKB); Tordera, along river, 41.45N 2.45E, 5. vii.1990, 1 ♂ (M. Barták) (MBC); Sudan: Wad Medani, compost of sweet potatoes etc., 22. v.1941, 2 ♀ (D.J.Lewis); caught in a trap, 16. i.1945, 4 ♂; [London School of Hygiene and Medicine] (BMNH: 1996–140); Syria: Malula, 27. vi.1994, 1 ♀ (M. Hradský) (MBC); Tadjikistan: 2♂: “Pamir” (MNKB); Prov. Dushanbe, Res.
Tigerbalken, a fluss vachtsch b. Nishni Piandsh, vi.1986, 2 ♂ (Muche) (ZSSM); Fayzabad Distr., 22. vii.1982, 1 ♂ (Isametdinov) (SIZK); Tunisia: “ v–vi.1913 ”, 1♂ (NHMW); Taberka area, 7–18 v.1988, 1 ♀ (Zool. Mus. Copenhagen Exp.) (ZMUC); Turkey: East, 70 km E of Erzincan, Yollerüstü, 19.vi. 187, 1♀ (T. Osten) (SNMS); Turkmenistan: Kerki, 2♀ (MTD); Berzengi S of Ashgabat, 24.04.1988, 1♀ (A. Antropov) (SIZK); Ashgabat, 17. vi.1928, 1 ♀, 10. vii.1928, 6 ♀, (Vlasov); Kushka, 5. vi.1926, 3 ♂, 1♀ (S. Paramonov) (SIZK); Ukraine: Kyiv Reg.: Kyiv, 16. vi.1920, 1 ♀ (S. Paramonov), 13. ix.1944, 1 ♀ (Bilanovskiy), [idem, in room,] 1. ix.1931, 1 ♀ [Bilanovskiy]; Maliutianka nr. Fastiv, 5. ix.1920, 1 ♀ (S. Paramonov); Irpin, 26. vii.1995, 1 ♀ (S. Korneyev) (SIZK); Cherkasy Reg,: Moshnohir’ya [49.4558512N 31.7360687E], 30. viii.1988, 1 ♀ (S. Zrazhevsky); Kaniv, Nature Reserve [49.7250063N 31.5322959E]: dung heap, 30. vi.1957, 1 ♂, greenhouse, 5. vi.1958, 1 ♂, 3♀, 16. vi.1958, 2 ♀, 22. vii.1959, 2 ♂, 7♀ (O. Viktorov-Nabokov), idem, vi.2007, 1 ♀ (S. Korneyev) (SIZK); Odesa Reg.: Ananyiv, 11. vii.1921, 1 ♀ (S. Paramonov) (SIZK); Kherson Reg.: Novooleksandrivka nr. Kalanchak [46.2684565N 33.4056044E], 23. vii.1987, 2 ♂, 3♀ (V. Korneyev) (SIZK); Zaporizhzha Reg.: “Osypenko” [=Berdiansk, 46.7624431N 36.7959595E], vi.1946 (Gulinov) (SIZK); Crymea: Yalta, 16. ix.1923, 2 ♂ (S. Paramonov); Mischor, viii.1926, 1 ♂ (Musytschenko) (SIZK); United Arab Emirates: Al-Ajban, 8♂, 5♀, 6–25.vii.2006, MT; 2♂, 13♀, 26.ii–2.iv.2006, MT; 4♂, 26.ii–27.iii.2006, LT; 1♂, 1♀; 6–22.v.2006, LT. Bithnah, 15♂, 7♀; 31.xii.2005 – 2.ii.2006, LT, Fujairah, 2♂, 2♀, 19.iv–2.v.2003, LT; 1♂, 1♀; 2–30.i.2006, LT; 32♂, 33♀, 28.ii–21.iii.2006, LT; 3♂, 6♀, 28.ii–1.iv.2006, LT; 13♂, 13♀, 20–27.v.2006, LT. Hatta, 43♂, 12♀, 19–28.iii.2006, LT; 30♂, 11♀, 8–26.iv.2006, LT; 28♂, 18♀, 24– 30.v.2006, LT. Near Mahafiz, 1♂, 2♀, 10–29.xii.2005, LT; 2♂, 3♀, 29.xii.2005 – 7.i.2006, LT. Sharjah-Khor Kalba, near tunnel, 6♂, 3♀, 16–31.i.2006, LT; 6♂, 10♀, 7–22.iii.2006, LT. NARC, near Sweihan, 24♂, 5♀, 16.xi–21.xii.2005, LT. Wadi Maidaq, 11♂, 2♀, 27.iv–4.v.2006, LT. Wadi Safad, 13♂, 4♀, 27.xi– 22.xii.2005, LT (leg. A. van Harten) (SIZK); USA: Minnesota: Waconia, 8. ii.1932, 1 ♂, 1♀ (Parten) (MNKB); Utah: Fielding, 14. ix.1926, 1 ♀; Logan, 2♂; Spanish Fork, 2♂, 2♀; Springville, 1♂; Ogden, 10. viii.1958, 7 ♂♀; Roy, 6♂ ♀; Benjamin, 2. viii.1958, 1 ♂ (DEI); Uzbekistan: Chimgan, damp meadow, 800m, 41.37N 70.00E, 18. v.1989, 1 ♀; Chirchiq, along river, 41.32N 69.38E, 19. v.1989, 1 ♂, Chinaz, along Syrdaria, 40.53N 68.43E, 20. v.1998, 2 ♂, 1♀; Samarqand, pasture, 39.39N 67.01E, 22. v.1989, 1 ♂; Zeravshan Reserve, deciduous wood, 39.38N 67.08E, 24. v.1989, 1 ♀ (M. Barták) (MBC); Kokand, Fergana, 1♂ (MTD); Yem en: Sana's, 4. vi.1987, 2 ♂ (Mühle) (ZSSM).
Diagnosis. P. alceae can be recognized from the combination of mostly brownish yellow frons and face, facial carina low, with entire white microtrichose band, scutum and scutellum subshining, finely rugulose, with dull greenish sheen, fore metatarsus mostly white; preglans of phallus with chain of partly fused black spines (Fig. 31 a). It can be differentiated with certainty from superficially similar P. azurea, P. hendeli sp. n. and P. orinigra sp. n. from the characters as given in the Key (couplets 27–30) and especially by the structure of the phallic preglans (Fig. 31 a).
Description. Head (Figs. 29–30). Frons 1.2 times as long as wide, brownish yellow, rarely to dark brown, satin shining, very sparsely and finely, almost inconspicuously setulose, with round parafrontal microtichose spots and two pairs of calluses (oval swellings) posterior to its middle and slightly concave anterior half. Vertical plates usually black, with greenish or dark blue sheen, bearing 2 pairs of black, short, slightly reclinate orbital setae. Ocellar triangle black.
Face reddish yellow to reddish brown, dorsal half of facial carina and antennal grooves entirely gray microtrichose without bare areas. Lunule shining orange to brown. Facial ridge, parafacial and gena shining orange or brown, gena 1/3 times as high as eye; only parafacial with narrow white microtrichose stripe along anteroventral eye margin. Epistome brownish yellow, rarely (a specimen from Czech Republic) dark brown with greenish sheen. Occiput black, with yellowish brown area behind ocellar triangle and postgena; orbit between posterodorsal eye margin and row of black postocular setae with very narrow white microtrichose stripe or without it. Medial vertical seta half as long as frons width, 1.5 times as long as lateral vertical and 3–5 times as long as ocellar, orbital and postocellar setae. Antenna reddish brown, greyish microtrichose; flagellomere 1 rounded apically, 1.5 times as long as wide; arista bare, yellow in basal 1/4, remainder black. Clypeus orange to brown. Palp brown to black, microtrichose, with moderately long black setulae. Mouthparts black.
Thorax (Fig. 30). Scutum and scutellum brown to black, with dull green metallic sheen, finely rugulose, except antepronotum, posterior surface of postpronotal lobe and notopleuron, as well as pleura strongly shining; posterodorsal part of anepisternum shagreened; supra-alar area and tympanal fossa matt black, postscutellum black, gray microtrichose; postero-ventral margin of scutellum without microtrichose area. Mesonotal scutum with short medial row of setulae in anterior portion, pair of regular dorsocentral and intra-alar rows, all setae very fine and short, black; pair of hair-like acrostichal and dorsocentral seta. One postprononal, 2 postsutural supra-alar, one intra-alar and one postalar setae strong, black.
Scutellum with golden sheen, with very fine and short black setulae and 2 pairs of black scutellar setae.
Wing. Entirely hyaline, with pale yellow veins; cell r4+5 almost closed, not forming petiole; postero-apical extension of cell cup 1.5 times as long as vein A1+CuA2, and 4 times as long as transverse section of vein CuA2. Costal vein from middle of costal cell to middle of r1 cell with alternate thickened and thin setae in antero-dorsal and antero-ventral rows. Length: 2.5–4.8 mm.
Legs. Black except fore tarsus with basitarsomere creamy yellow in basal 3/4, mid- and hind tarsi yellow; all setae black; fore femur postero-ventrally with 5–7 thickened, but rather short setae in apical half.
Abdomen. Both tergites and sternites brown or black-brown, with very weak bluish or greenish reflection, with black setulae; femaleabdominal tergite 2 with pair of matt gray spots (= dimple-like structures) laterally; tergite 5 in both male and female conspicuously shagreened, subshining black, sometimes with deep blue reflection.
Male postabdomen brown to black; epandrium as on Figs. 33–34, phallus with stipe twice as long as preglans and glans; preglans with chain of partly fused black spines (Fig. 31 a); glans as on Figs. 31 and 32.
Female terminalia: eversible membrane (Fig. 35) with two pairs of long taeniae and membrane between them cowered by fine monodentate scales; posterior part of membrane with fine multidentate scales; aculeus (Figs. 36– 37) 6.5–8 times as long as wide at base; vagina (Figs. 38–39) with simple, rod-like ventral receptacle; 3 spherical spermathecae (Fig. 40).
Distribution: Widespread in the Palaearctic Region, except North; introduced to the Nearctic, Neotropical and Australasian Region as synanthropic species; quite rarely in the Afrotropical Region.
Biology. Larvae feed in rotting date palm stems, dung and dung/vegetal compost. It is believed to be widespread with cattle due to shift of its original larval feeding in the date palm stems to the dung. Remarks. One of the most common ulidiid species in the temperate and arid zones of the Palaearctic Region. Type specimens of most nominal species have not been examined, but since it is the only species occurring in Middle and Northern Europe, its identity is clear.
description: Published as part of Elena P. Kameneva & Valery A. Kroneyev, 2016, Revision of the Genus Physiphora Fallén 1810 (Diptera: Ulidiidae: Ulidiinae), pp. 1-88 in Zootaxa 4087 (1) on pages 13-17, DOI: 10.11646/zootaxa.4087.1.1, http://zenodo.org/record/399511