Chthonius (Chthonius) raridentatus Hadzi 1930
Creators: name: Gardini, Giulio
Publisher: name: Zenodo
Year: 2014
Subjects: Biodiversity
Taxonomy
Animalia
Arthropoda
Arachnida
Pseudoscorpiones
Chthoniidae
Chthonius
Chthonius raridentatus

description: Chthonius (C.) raridentatus Hadži, 1930
(Figs 59–89, 90)
Chthonius cavernarum Ellingsen 1909: 214 (in part).
Chthonius cavernarum (not Ellingsen 1909): Beier 1928: 313 (misidentification); Beier 1931: 91 (misidentification); Beier 1932: 53, fig. 61 (misidentification, in part: see C. cavernarum).
Chthonius (C.) raridentatus Hadži 1930: 134, figs 10–11, 12a–d, 13a–e; Beier 1939b: 15, fig. 11.
Chthonius (C.) ellingseni Beier 1939b: 16 (n. syn.).
Chthonius (C.) brandmayri Callaini 1986: 237, figs 2a–e. (n. syn.).
Chthonius (Chthonius) jalzici Ćurčić 1988: 17, figs 2–8 (n. syn.).
Chthonius (Chthonius) aff. ellingseni: Ćurčić 1988: 22, figs 18–22.
Type locality: Slovenia, Kočevsko, Kočevje, Borovec pri Kočevski Reki, Babja luknja pri Borovcu no. 93 (= Frauenloch; Jama pri Borovcu) (45°33′07″ N 14°47′ 41″E).
Distribution. NE Italy, Austria, Slovenia, Croatia.
Diagnosis (♂♀). An eyed, epigean Chthonius (Chthonius) from the eastern Alps and northern Dinaric region that differs from other species of the C. heterodactylus group in the following combination of characters: carapace without epistome, chaetotaxy 4:6:4:2:2 (18), rarely anterior row m 4m and posterior row m 2m; chaetotaxy tergites I–IV: 4:4:4:4; genital opening of males flanked by aligned setae on each side; fixed chelal finger with 15–26 (♂) or 12–21 (♀) teeth with dental canals, usually with intercalary microdenticles, and with a spatulate distal paraxial seta; movable chelal finger usually with 20–28 (♂) or 16–26 (♀) low, broad, reclined teeth, increasingly reduced towards finger base, reaching back almost to sb; distal half of movable chelal finger rarely with 6–8 triangular reclined teeth that are wider than high; coupled sensilla pc of movable chelal finger usually between trichobothria st and sb; chela length 0.86–1.51 (♂) or 0.88–1.74 (♀), movable finger length 0.51–0.90 (♂) or 0.54–1.0 (♀).
Type material of Chthonius ellingseni examined. AUSTRIA — Carinthia: 2 ♀ (paralectotypes of Chthonius (C.) ellingseni Beier, 1939, present designation), “Annenheim am Ossiachersee / Kärnten. 1893 Sturany leg.” “ Chthonius cavernarum Ell. / det. Dr. Beier” (NHMW).
SLOVENIA— Gorenjska regija: 1 ♀ (lectotype of Chthonius (C.) ellingseni, present designation) 2 ♂, 4 T (paralectotypes of Chthonius (C.) ellingseni, present designation), “Crna Prst [Mt. Črna prst] Wochein [Bohinj] / 700-800 m / Juli-August 1899 / Penther leg. XXIV” “ Chthonius cavernarum Ell. / det. Dr. Beier” (NHMW).
Type material of Chthonius brandmayri examined. SLOVENIA— Obalno — Kraška regija: 1 ♂ (holotype of Chthonius brandmayri Callaini, 1986), “F4 / Istria—M. Taiano (Slavnik) / 1.IX.1975 faggeta chiara / leg. Brandmayr ” “ Chthonius brandmayri n. sp. / ♂ holotypus Callaini det.” (MSNV); 1 ♀ (paratype of Chthonius brandmayri Callaini), “Istria—M. Taiano (Slavnik) / 22.X.1975 terra rossa (F2) / leg. Brandmayr” “ Chthonius brandmayri n. sp. / ♀ paratypus Callaini det.” (MSNV).
Other material examined. CROATIA— Istarska županija: 3 ♂, 3 ♀, Učka (= Monte Maggiore), 1390 m a.s.l., 6.X.1990, F. Gasparo leg., meadow on edge of Fagus sylvatica wood; 1 ♂, id., 950 m a.s.l., 6.X.1990, F. Gasparo leg., pitfall traps. Ličko–senjska županija: 2 ♂, 1 ♀, Velebit, 4 km southern of Oltari, 1050 m a.s.l., 13.VI.1996, G. Colombetta leg., pitfall traps in Fagus sylvatica wood. Zagrebačka županija: 2 ♂, 1 ♀, 2 T, 1 D, Zagreb, Medvednica, Gornji Stenjevec, Veternica špilia, 28.XII.2003, R. Ozimec leg. (ROZ); 2 ♂, 1 ♀, id., 9.IX.2006, R. Ozimec leg. (ROZ); 4 ♂, 2 ♀, 5 D, id., 13.V.2007, Bedek, Bilandžija & Pavlek leg., traps (ROZ).
ITALY — Veneto: Treviso Prov.— 4 ♂, 1 ♀, Susegana, Castello San Salvatore, Ricovero difensivo 74 V /TV, 150 m a.s.l., 1.IX.2002, V. Forte & S. Vanin leg. (SVP); 2 ♂, 1 T, id., Ricovero difensivo 60 V /TV, 107 m a.s.l., 1.IX.2002, V. Forte & S. Vanin leg. (SVP); 1 ♂, id., 15.III.2003 (SVP). Friuli: Pordenone Prov. — 1 ♂, 1 ♀, Barcis, Montelonga slope N, 800/ 900 m a.s.l., 11.IX.2006, G. Governatori leg.; 1 ♀, Claut, Rifugio Pussa, 24.VI.2000, G. Agazzi leg.; 1 ♂, 1 ♀, Clauzetto, doline of Caverna Mainarda, 540 m a.s.l., 29.VI.1989, F. Gasparo leg.; 1 ♀, id., Grotte Verdi 116 Fr/PN, 540 m a.s.l., 29.IX.1990, F. Gasparo leg.; 1 ♂, Meduno, 650 m a.s.l., 9.VI.1985, M. Seriani leg.; 1 ♂ (C. brandmayri, G. Gardini det. 2002), Sequals, peat bog of Sequals, 200 m a.s.l., UM31, 14.X.2001, G. Colombetta leg., trap in wood (MFNB); 1 ♂, Tramonti di Sotto, Buse dal Picissat 221 Fr/PN, 434 m a.s.l., 30.IX.1982, F. Gasparo leg.; 1 ♂, Zoppola, Parco Fluviale del Meduna, 10.VIII.1991, G. Governatori leg. Friuli: Udine Prov. —1 ♀, Artegna, 250 m a.s.l., 11.VIII.1992, F. Gasparo leg.; 2 ♂, Attimis, Porzus, 750 m a.s.l., 10.VII.1990, F. Gasparo leg.; 7 ♀, Chiusaforte, between Chiusaforte and Sella Nevea, 700 m a.s.l., 18.IX.1991, F. Gasparo leg.; 1 T, id., 6.XI.1991, F. Gasparo leg., pitfall trap; 1 ♀, Chiusaforte, Sella Nevea, 1170 m a.s.l., 5.IX.1990, G. Governatori leg.; 4 ♂, 6 ♀, id., 1190 m a.s.l., 21.VII.1992, F. Gasparo leg.; 1 ♂, id., 1180 m a.s.l., 31.V.1989, F. Gasparo leg.; 1 ♂, Cividale del Friuli, Castelmonte, 600 m a.s.l., 20.VII.1989, F. Gasparo leg., Fagus sylvatica wood; 3 ♂, 1 T (C. brandmayri, G. Gardini det. 2002), Fagagna, Quadris wet meadows, 171 m a.s.l., UM51, 14.X.2001, G. Colombetta leg., traps wet wood (MFNB); 2 ♀, 1 T (C. brandmayri, G. Gardini det. 2002), id., 21.VI.2001, G. Colombetta leg., traps wet wood (MFNB); 1 ♂, Grimacco, Grotta a E di Grimacco Superiore 2967 Fr/UD, 21.II.1997, G. Governatori leg.; 1 ♂, 1 ♀, Grimacco, Valle del T. Cosizza near Clodig, 1.IV.1990, F. Stoch leg.; 2 ♂, 3 ♀, Lusevera, near Casera Morandin, 340 m a.s.l., 6.X.1997, G. Governatori leg.; 3 ♂, Lusevera, Passo di Tanamea, 850 m a.s.l., 23.VIII.1989, F. Gasparo leg.; 1 T (C. brandmayri, G. Gardini det. 2002), id., UM72, 5.VIII.1992, G. Governatori leg., Fagus sylvatica wood (MFNB); 2 ♂, 1 ♀, 1 T, id., 24.IX.1997, F. Gasparo leg., Picea and Fagus wood; 3 ♂, 4 T (C. brandmayri, G. Gardini det. 2008), id., 850 m a.s.l., UM72, 19.VII/ 5.X.2006, G. Colombetta leg., pitfall traps in meadow (MFNB); 1 T (C. brandmayri, G. Gardini det. 2008), id., 850 m a.s.l., UM72, 12.VII/ 14.VIII.2007, L. Dorigo leg., pitfall trap in open meadow (MFNB); 1 ♂ (C. brandmayri, G. Gardini det. 2008), Lusevera, Pian di Tapou, 900 m a.s.l., UM63, 29.VI/ 6.VIII.2006, G. Colombetta & L. Dorigo leg., pitfall trap in Fagus sylvatica wood (MFNB); 1 ♀ (C. brandmayri, G. Gardini det. 2008), id., 4.IX/ 5.X.2006, G. Colombetta leg., pitfall trap in Fagus sylvatica wood (MFNB); 1 ♂, 3 T, 4 P (C. brandmayri, G. Gardini det. 2008), id., 17.IV/ 21.VI.2007, L. Dorigo leg., Fagus sylvatica wood (MFNB); 2 ♀, 1 T (C. brandmayri, G. Gardini det. 2002), id., 13.VII/ 29.VIII.2007, L. Dorigo leg., pitfall traps in Fagus sylvatica wood (MFNB); 1 ♂ (C. brandmayri, G. Gardini det. 2002), Majano, peat bog of Casarola, 160 m a.s.l., UM51, 14.X.2001, G. Colombetta leg., traps in wood (MFNB); 1 ♀, Masarolis, 10.V.1998, M. Grottolo leg.; 2 ♂ (C. brandmayri, G. Gardini det. 2008), Moggio Udinese, slope of Monte Zovet, Stavoli Cuel Lung Alto, 700 m a.s.l., UM53, 15.V/ 16.VII.2001, G. Governatori & A. Dall’Asta leg., pitfall traps in Fagus sylvatica wood (MFNB); 1 ♂ (C. brandmayri, G. Gardini det. 2008), id., 20.VIII/ 26.IX.2001, G. Governatori leg., pitfall trap in Fagus sylvatica wood (MFNB); 2 T (C. brandmayri, G. Gardini det. 2008), id., 22–27.VII.2002, A. Dall’Asta leg., litter leaf, Fagus sylvatica wood (MFNB); 3 ♂, 2 ♀, 1 T, 2 D (C. brandmayri, G. Gardini det. 2002), Moggio Udinese, Stavoli Tugliezzo, Rio Lavarie, 470 m a.s.l., UM53, 26.IX/ 30.X.2001, G. Governatori & A. Dall’Asta leg., cone traps in Fagus sylvatica wood (MFNB); 1 ♂, 2 ♀, 4 T (C. brandmayri, G. Gardini det. 2002), Moruzzo, peat bog of Borgo Pegoraro, 185 m a.s.l., UM50, 21.VI.2001, G. Colombetta leg., traps in wood (MFNB); 1 ♀, 2 T, 1 D (C. brandmayri, G. Gardini det. 2002), id., 16.X.2001, G. Colombetta leg., traps in wood (MFNB); 2 ♂, 1 ♀, Musi, 700 m a.s.l., 1.VI.1986, A. Morisi leg.; 2 ♂, 11 ♀, Musi, northern slope of Gran Monte, 1150 m a.s.l., 27.VI.1995, G. Governatori leg.; 4 ♂, 3 ♀, id., 22.VI.1995, G. Governatori leg., Fagus sylvatica wood; 1 ♀, near Musi, 1.VI.1986, R. Sciaky leg.; 1 ♂, Nimis, Grotta di Monteprato 59 Fr/UD, 525 m a.s.l., 26.III.1983, F. Gasparo leg. (together with 1 ♂ of C. cavernarum); 1 ♂, id., 4.VII.1988; 1 ♀, id., 6.VII.1989; 1 ♂, id., 15.VI.1997; 1 ♀, Nimis, Nongruella, 450 m a.s.l., 29.IX.1997, F. Gasparo leg.; 1 ♂, 1 ♀, Osoppo, 200 m a.s.l., 15.XI.1990, F. Gasparo leg., trap 30 cm deep; 1 ♂ (C. exarmatus Hadži, Beier det.), Ovaro, Vals, 8.VIII.1950, A. Valle leg. (MSNV); 1 ♀, Paluzza, Grotta presso la centrale di Timau 89 Fr/UD, 970 m a.s.l., 13.XII.2003, G. Governatori leg.; 2 ♂, 3 ♀, 2 T (C. brandmayri, G. Gardini det. 2002), Pagnacco, peat bog of Lazzacco, 195 m a.s.l., UM51, 21.VI.2001, G. Colombetta leg., traps in wood (MFNB); 4 ♂, 2 ♀ 1D (C. brandmayri, G. Gardini det. 2002), id., 16.X.2001, G. Colombetta leg., traps in wood (MFNB); 1 ♂, Pontebba, Passo Pramollo, 1550 m a.s.l., 4.VII.1991, F. Gasparo leg., pasture with Abies; 2 ♀, Pontebba, 600 m a.s.l., 20.IX.1990, F. Gasparo leg.; 1 ♂, Pontebba, between Pontebba and Passo Pramollo, 1070 m a.s.l., 4.VII.1991, F. Gasparo leg.; 1 ♂, 1 T (C. brandmayri, G. Gardini det. 2002), Porpetto, Fraghis swamp, 8 m a.s.l., UL68, 26.VI.2001, G. Colombetta leg., traps in wood (MFNB); 2 ♂, 1 ♀, 1 T (C. brandmayri, G. Gardini det. 2002), id., 17.X.2001, G. Colombetta leg., traps in wood (MFNB); 4 ♂, Resia, Coritis, 620 m a.s.l., 27.V.1992, F. Gasparo leg.; 6 ♂, 4 ♀, Resia, Lischiazze, 580/ 600 m a.s.l., 8.VI.1997, F. Gasparo leg., Fagus sylvatica wood; 2 ♂, 4 ♀, 1 T, 1 D (C. brandmayri, G. Gardini det. 2008), Resia, Monte Zaiavor, near Bocchette di Zaiavor, 1450 m a.s.l., UM73, 19.VII/ 12.IX.2006, L. Dorigo leg., pitfall traps in meadow (MFNB); 6 ♂, 4 ♀ (C. brandmayri, G. Gardini det. 2008), id., 17.IV/ 6.VI.2007, L. Dorigo leg., pitfall traps in alpine meadow (MFNB); 1 ♂, 2 ♀, 3 T, 1 D (C. brandmayri, G. Gardini det. 2008), id., 12.VII/ 4.X.2007, L. Dorigo leg., pitfall traps in alpine meadow (MFNB); 1 ♀ (C. brandmayri, G. Gardini det. 2002), Resia, Sant’Anna di Carnizza, 1060 m a.s.l., UM73, 18.V/ 19.VI.2001, G. Governatori leg., pitfall trap in meadow (MFNB); 1 ♀ (C. brandmayri, G. Gardini det. 2002), id., 19.VI.2001, G. Governatori leg., leaf litter in Fagus sylvatica wood (MFNB); 2 ♂, 2 ♀, Resia, Sella Carnizza, 1050 m a.s.l., 2.V.1990, F. Gasparo leg.; 3 ♂ (C. brandmayri, G. Gardini det. 2002), Resia, Sella Carnizza, 1080–1090 m a.s.l., UM73, 18.V/ 19.VI.2001, G. Governatori & A. Dall’Asta leg., pitfall traps in Fagus sylvatica wood (MFNB); 1 T, 1 D (C. brandmayri, G. Gardini det. 2002), id., 1086 m a.s.l., 9.VIII/ 21.IX.2002, A Dall’Asta & L. Lapini leg., leaf litter in Fagus sylvatica wood (MFNB); 1 ♂ (C. brandmayri, G. Gardini det. 2002), Resia, above Clen, near Casera Coot, 950 m a.s.l., UM73, 26.IX/ 30.X.2001, A. Dall’Asta & G. Governatori leg., cone traps in Fagus sylvatica wood (MFNB); 5 ♂, 3 ♀, 3 T, 1 D (C. brandmayri, G. Gardini det. 2002), id., above Clen, between Casera Coot and Slatina Superiore, 940 m a.s.l., UM73, 27.VI/ 30.X.2001, G. Governatori leg., cone traps in pasture (MFNB); 1 ♂, 1 T (C. brandmayri, G. Gardini det. 2002), id., 1000 m a.s.l., UM73, 19.VI/ 20.VIII.2001, G. Governatori & A. Dall’Asta leg., pitfall traps in Fagus sylvatica wood (MFNB); 2 ♂ (C. brandmayri, G. Gardini det. 2002), id., 26.IX/ 30.X.2001, G. Governatori leg., pitfall traps in Fagus sylvatica wood (MFNB); 3 ♂, 2 ♀, 1 T (C. brandmayri, G. Gardini det. 2008), id., Val Uccea, near Stalli Rasuga, 950 m a.s.l., UM73, 6.VI/ 16.X.2007, G. Colombetta, L. Dorigo & P. Glerean leg., pitfall traps and under stones in Fagus sylvatica wood (MFNB); 1 ♂ (C. brandmayri, G. Gardini det. 2008), id., Val Uccea, near Stavoli Tanatemea, 980 m a.s.l., 29.VI/ 2.VIII.2006, G. Colombetta & L. Dorigo leg., pitfall traps in Fagus sylvatica wood (MFNB); 1 ♂, 3 ♀, 7 T, 1 D (C. brandmayri, G. Gardini det. 2008), id., Val Uccea, near Torrente Uccea, 650 m a.s.l., UM72, 13.VII/ 5.X.2006, G. Colombetta & L. Dorigo leg., pitfall traps (MFNB); 5 ♂, 2 ♀, 10 T (C. brandmayri, G. Gardini det. 2008), id., 17.IV.2006 / 6.VI.2007, G. Colombetta & L. Dorigo leg., pitfall traps (MFNB); 5 ♂, 7 ♀, 4 T, 1 D (C. brandmayri, G. Gardini det. 2008), id., 6.VI/ 16.X.2007, G. Colombetta, L. Dorigo & P. Glerean leg., pitfall traps (MFNB); 1 ♂, Rigolato, 730 m a.s.l., 20.IX.1996, F. Gasparo leg., Picea abies wood; 1 ♀, Savogna, Cepletischis, Grotta La Mitica 2907 Fr/UD, 525 m a.s.l., 15.V.1998, G. Governatori leg.; 1 ♀, near Savogna, 250 m a.s.l., 26.VI.1994, S. Zoia leg., Fagus sylvatica wood; 6 ♂, 2 ♀, Savogna, Monte Matajur, 1200 m a.s.l., 25.V.1986, S. Zoia leg.; 1 ♀, Sella Nevea, slope W Monte Poviz, 1885 m a.s.l., 16.IX/ 15.X.1990, G. Governatori leg., alpine meadow (MFNB); 1 ♂, id., slope NE Monte Canin, 1220 m a.s.l., 13.V/ 14.VI.1991, G. Governatori leg. (MFNB); 2 ♂, 1 ♀, Stregna, 9.V.1998, M. Grottolo leg.; 1 ♂, Taipana, Monteaperta, slope S Gran Monte, 700 m a.s.l., 10.IX/ 10.X.1994, G. Governatori leg., meadow (MFNB); 1 ♀, Taipana, Grotta Barloch 61 Fr/UD, 20.IV.2001, M. Grottolo leg.; 4 ♂, 4 ♀, Taipana, Platischis, 700 m a.s.l., 9.V.1990, F. Gasparo leg.; 2 ♀, Tarvisio, Coccau, 750 m a.s.l., 18.VII.1991, F. Gasparo leg.; 1 ♀, Torreano, Prestento, 190 m a.s.l., 10.VII.1993, F. Gasparo leg.; 1 ♀, near Uccea, 780 m a.s.l., 1.VI.1986, S. Zoia leg.; 5 ♂, Venzone, Piani, 270 m a.s.l., 9.VIII.1995, F. Gasparo leg., Ostrya carpinifolia wood; 2 ♂ (C. brandmayri, G. Gardini det. 2008), id., Monte Plauris, near Malga Confin, 1315 m a.s.l., UM63, 19.VI/ 25.VII.2007, G. Colombetta & L. Dorigo leg., leaf litter and pitfall trap in Fagus sylvatica wood (MFNB); 2 ♂ (C. brandmayri, G. Gardini det. 2008), id., 4.IX/ 9.X.2007, G. Colombetta leg., pitfall traps in Fagus sylvatica wood (MFNB); 1 ♂, 3 ♀, 1 T (C. brandmayri, G. Gardini det. 2008), id., Monte Plauris, near Casera Ungarina, 1320 m a.s.l., UM63, 1.IX.2006 / 30.IV.2007, G. Colombetta & L. Dorigo leg., pitfall traps in Fagus sylvatica wood (MFNB); 1 ♂, 3 ♀, id., 30.IV/ 19.VI.2007, G. Colombetta & L. Dorigo leg., pitfall trap in Fagus sylvatica wood; 1 ♂, 1 ♀ (C. brandmayri, G. Gardini det. 2008), id., 19.VI/ 25.VII.2007, G. Colombetta & L. Dorigo leg., pitfall traps in Fagus sylvatica wood (MFNB); 1 ♂ (C. brandmayri, G. Gardini det. 2008), id., Val Venzonassa, Borgo Prabunello, 765 m a.s.l., UM63, 20.VI/ 2.VIII.2006, G. Colombetta & L. Dorigo, pitfall trap in wood (MFNB); 1 ♀ (C. brandmayri, G. Gardini det. 2008), id., 30.IV/ 19.VI.2007, G. Colombetta & L. Dorigo leg., pitfall trap in Orno–Fagetum (MFNB); 1 ♀ (C. brandmayri, G. Gardini det. 2008), id., Val Venzonassa, near Casera Frassin, 630 m a.s.l., UM63, 6.IX/ 9.X.2007, G. Colombetta leg., pitfall in Fagus sylvatica wood (MFNB); 1 ♂, Villa Santina, 400 m a.s.l., 30.IV.1991, F. Gasparo leg., Carpinus wood. Venezia Giulia: Trieste Prov.— 1 ♀ (C. ellingseni, V. Mahnert det. 1984), Duino–Aurisina, Grotta di Ternovizza 242 VG/TS, 275 m a.s.l., 15.VIII.1981, F. Gasparo leg. (MCST); 4 ♂, 2 ♀, 3 T (C. ellingseni, V. Mahnert det. 1984), Monrupino, Caverna dei Ciclami 2433 VG/TS, 324 m a.s.l., 12.VII.1981, F. Gasparo leg. (MCST); 1 ♂, 2 ♀ (♂♀ C. brandmayri, ♀ C. raridentatus, G. Gardini det. 1986, 2002), San Dorligo della Valle, Grotta di Crogle 2716 VG/TS, 370 m a.s.l., 10.X.1982, F. Gasparo leg., entrance and inside the cave; 1 ♀ (C. raridentatus, G. Gardini det. 2002), id., 7.V.1988, F. Gasparo leg.; 2 ♀, id., 23.II/ 13.VI.2009, F. Stoch & G. Tomasin leg. (MCST); 2 ♀ (C. ellingseni, V. Mahnert det. 1984), Sgonico, Grotta dell’Orso 7 VG/TS, 224 m a.s.l., 4.X. and 9.XI.1980, F. Gasparo leg. (MCST); 6 ♂, 7 ♀ (C. brandmayri, G. Gardini det. 2002), id., Grotta dell’Orso 7 VG/TS, 30.VIII.1987, G. Gardini & R. Rizzerio leg., entrance; 1 ♀ (C. brandmayri, G. Gardini det. 2002), id., 29.III.1991, M. Bodon leg., entrance; 2 ♂, 1 ♀ (C. brandmayri, G. Gardini det. 2002), Trieste, Sercetova Jama 290 VG/TS, 310 m a.s.l., 24.X.1982, F. Gasparo leg.; 2 ♂, 2 T (C. brandmayri, G. Gardini det. 2002), id., 14.VII.1985, F. Gasparo leg.; 2 ♂ (C. brandmayri, G. Gardini det. 2002), Trieste, Vicolo delle Rose 72, 120 m a.s.l., 18.V.1986, F. Gasparo leg.
SLOVENIA— Gorenjska regija: 2 ♀, “140 Chthonius / cavernarum Ell. n. sp. / Austria: Krain Sever” [= “Bischoflacker Grotte” (see Ellingsen 1909), not identified, but presumably either Migutovo brezno, no. 5 (451 m a.s.l., 46°09′36.7″N 14°17′51.7″E) or Marijino brezno no. 6 (429 m a.s.l., 46°09′45″N 14°17′50″E), both located near Škofja Loka] (paralectotypes of C. cavernarum, see Remarks under C. cavernarum) (ZMUN); 1 ♀, Cerklje na Gorenjskem, Pšata, Urhova jama (= Urhovo brezno), n. 505, 10.I.1979, T. Novak & M. Štangelj leg., trap (PMSL); 2 ♂, Bled, castle’s hill, 580 m a.s.l., 25.IV.1992, F. Gasparo leg.; 1 ♀, 1 T, 17 D, Crngrob, Škofja Loka, Štinetova jama, n. 240, 25.VII.1980, T. Novak & I. Sivec leg. (PMSL). Goriška regija: 3 ♂, 1 T, between Predmeja and Lokve, 1130 m a.s.l., 1.VIII.1992, F. Gasparo leg., Fagus sylvatica wood; 6 ♂, 8 ♀, between Godovič and Črni Vrh, 700 m a.s.l., 8.IX.1991, F. Gasparo leg., Fagus sylvatica wood; 2 ♂, 4 ♀, Godovič, southeastern of Idrija, 600 m a.s.l., 24.VI.1990, F. Gasparo leg.; 1 ♂, 3 ♀, Idrija, 350 m a.s.l., 28.X.1990, F. Gasparo leg., mixed wood with Fagus; 4 ♂, 2 ♀, 2 T, Most na Soči, near Jama pod Smoganico, n. 823, 15.X.1968, G. & A. Vigna leg., under stones in Fagus sylvatica wood; 1 ♂, 2 ♀, id., Jama pod Smoganico, n. 823, 26.VI.1996, M. Grottolo leg., entrance; 2 ♂, id., 500 m a.s.l., 15.IX.2003, F. Gasparo leg., entrance. Jugovzhodna Slovenija regija: 1 ♂, 2 ♀, Mozelj, Kočevje, 500 m a.s.l., 11.V.1993, F. Gasparo leg.; 1 ♀, 300 m NNE of Ograja, 3.V.2008, L. Slana Novak & T. Novak leg.; 2 ♀, near Trebnje, 26.VIII.1987, G. Gardini & R. Rizzerio leg. Koroška regija: 1 ♂, 1 T, Suhi dol, Slovenj Gradec, Rdeški vrh, Rdečka jama, n. 3488, 10.VII.1977, V. Kuštor & T. Novak leg., Berlese (PMSL); 1 ♂, Suhi dol, Slovenj Gradec, Skobirjeva votlica, n. 3956, 7.VII.1978, T. Novak & T. Prošek leg. (PMSL). Notranjsko–Kraška regija: 4 ♂, 1 ♀, Podgrad (= Castelnuovo d’Istria), 570 m a.s.l., 20.IV.1992, F. Gasparo leg.; 1 ♂, Hrušica (= Selva di Piro), 22.VI.1982, M. Seriani leg.; 1 ♀, Postojna, Polhovica (5 km northwestern from Postojna), 600 m a.s.l., 15.VII.1990, F. Gasparo leg., pitfall traps; 3 ♂, 1 ♀, id., 19.IV.1992, F. Gasparo leg.; 1 ♂, 1 ♀, id., 28.IX.1997, F. Gasparo leg.; 1 ♂, 2 ♀, Postojna, Rakov Škocjan, 520 m a.s.l., 5.X.1991, F. Gasparo leg. (together with 2 ♂ of C. cavernarum); 2 ♂, 1 ♀, id., 9.V.1993, F. Gasparo leg., mixed wood; 1 ♂, 1 ♀, Kalič near Postojna, 780 m a.s.l., 10.V.1992, F. Gasparo leg.; 10 ♂, 1 T, Postojna, northeastern slope of Mt. Nanos, 1150 m a.s.l., 4.VII.1993, F. Gasparo leg., Fagus sylvatica wood; 4 ♂, Postojna, 3 km northwestern of Postojna, 600 m a.s.l., 6.V.1990, F. Gasparo leg.; 4 ♂, 2 ♀, between Postojna and Planina, 600 m a.s.l., 2.VI.1991, F. Gasparo leg.; 1 ♂, 1 ♀, Postojna, Lož, 600 m a.s.l., 14.X.1990, F. Gasparo leg. (together with 1 ♂, 4 ♀ of C. cavernarum); 1 ♂, 2 ♀, 1 T, Postojna, Šmihel pod Nanosom, Šmihel, 25.V.1980, T. Novak leg. (PMSL); 10 ♂, 4 ♀, Snežnik, Sviščaki, 1250 m a.s.l., 25.VIII.1991, F. Gasparo leg. (together with 3 ♂, 6 ♀ of C. cavernarum); 1 ♀, Postojna, Zguba jama, 9.XII.2012, T. Novak leg., pitfall trap (PMSL). Obalno–Kraška regija: 1 ♀, Divača, Škocjanske jame (= Grotte di San Canziano), 29.VIII.1987, G. Gardini & R. Rizzerio leg., entrance. Osrednjeslovenska regija: 1 ♀, 250 m northeastern of V. Stena, 439 m a.s.l., 3.V.2008, L. Slana Novak & T. Novak leg., Fagus sylvatica wood; 1 ♂, 300 m northwestern of Čemšenik, 21.VII.2002, L. Slana Novak & T. Novak leg., Fraxinus, Carpinus, Tilia wood; 2 D, Krtina, Domžale, Boštonova jama, n. 757, 16/ 18.VII.1980, I. Sivec & T. Novak leg., Berlese (PMSL); 1 ♀, Grosuplje, Velika Račna, Jama Virsnica, n. 571, 349 m a.s.l., 24.VIII.1995, F. Gasparo leg.; 1 T, Ihan, Domžale, Ihanščica, n. 46, 18.VII.1980, T. Novak & I. Sivec leg., Berlese (PMSL); 5 ♀, Šmartno pod Šmarno Goro, Matjaževa jama, 29.VIII.1987, G. Gardini & R. Rizzerio leg., entrance. Podravska regija: 2 ♀, Laška Vas, Štore, Bojanski vrh, Fantovska jama II, n. 3967, 13.X.1978, T. Novak & I. Sivec leg. (PMSL). Savinjska regija: 2 ♀, Gruska Lesično, Kozje, Gruška jama, n. 1374, 16.X.1978, T. Novak & I. Sivec leg. (PMSL); 1 ♀, Završe, Dolič, Velenje, Mt. Tisnik, Jama Školjka, n. 3311, 14.X.1977, V. Kuštor & T. Novak leg., pitfall trap (PMSL); 1 ♀, 1 T, Suho, Šentjur near Celje, Jama v kamnolomu pri Suhem, n. 4632, 21.VII.1978, T. Novak & I. Sivec leg., pitfall trap (PMSL); 1 ♀, Rimske toplice, Mt. Grmada, Kapelarjevo brezno, n. 4706, 10.VII.1978, T. Novak leg. (PMSL); 1 D, Liboje, Celje, Štadler’s forest, Ocvirkova jama v Štadlerjevem gozdu, n. 5348, 21.VII.1979, V. Kuštor & T. Novak leg., pitfall trap (PMSL); 1 ♂, 1 ♀, Ponikva, 25.VIII.1984, T. Novak & L. Slana Novak leg.; 1 T, Galicija, Velika Pirešica, Špegličeva jama, n. 3512, 7.I.1979, T. Novak leg., Berlese (PMSL); 1 ♀, id., 11.X.1978, T. Novak & I. Sivec leg. (PMSL); 1 ♂, id., 13.X.1978, T. Novak & I. Sivec leg. PMSL); 1 ♂, id., 9.I.1979, T. Novak & M. Štancelj leg., pitfall trap (PMSL); 2 ♂, 1 ♀, id., 2.V.1997, M. Grottolo leg. Spodnjeposavska regija: 1 ♀, Krško, 1 km southwestern of Anže, 290 m a.s.l., 14.X.2005, L. Slana Novak & T. Novak leg., Fagus sylvatica wood; 2 ♀, Brežice, 28.VIII.1987, G. Gardini & R. Rizzerio leg. (together with 1 T of C. cavernarum). Zasavska regija: 2 ♂, 2 ♀, Lipovica, Izlake, Jama v Lipovici (= Lovrinova jama), n. 1182, 21.VII.1978, I. Sivec & T. Novak leg. (PMSL).
Description of adults (♂♀). Integument pigmented, carapace, tergites, chelicerae and pedipalps brown or pale brown (lectotype and paralectotypes of C. ellingseni with integument decoloured); hispid granulation on lateral surfaces of carapace, on cheliceral palm, on base of cheliceral movable finger and on distal surface of pedipalpal hand. Carapace 1.0–1.1 (rarely 0.9 or 1.2) times longer than broad, posteriorly weakly constricted; anterior margin (Figs 59–60, 67–68, 75–76, 82) without epistome (rarely with weakly prominent epistome), with sharp denticles between median macrosetae; ocular area as in Figs 61–62, 69–70, 77, 83; anterior eyes with convex lens (diameter 0.040–0.055 mm), posterior eyes with flat lens or reduced to spots, all eyes with tapeta (usually evident in recently preserved specimens); distance from anterior eyes to anterior margin of carapace 0.030–0.050 mm; chaetotaxy 4:6:4:2:2 (18), rarely anterior row m 4m (see Hadži 1930: 135) and posterior row m 2m (♀ from Slovenia, Urhova jama, no. 505); length of anteromedian and anterolateral macrosetae respectively 0.13–0.19 and 0.06–0.11 mm, length of median macrosetae of the ocular row 0.16–0.22 mm, length of posterior macrosetae 0.13–0.19 mm. Standard chaetotaxy of tergites I–X 4:4:4:4:4:6:6:6:6:4; tergite I rarely with 2 setae (♂ holotype and ♀ paratype of C. brandmayri from Mt. Slavnik; 1 ♂ from Most na Soči, near Jama pod Smoganico, n. 823, 15.X.1968) or with 3 setae (1 ♀ from Italy, Masarolis; 1 ♀ from Slovenia, Most na Soči; 2 ♂, id., 15.IX.2003; 2 ♀ from Slovenia, between Godovič and Črni Vrh). Standard chaetotaxy of sternites II–X 10:(3)8–10(3):(2)7–8(2):8:6:6:6:6:7; sternite II rarely with 9 or 11 setae, IV rarely with 9 setae and 1 stigmatal microseta; V rarely with 9 setae; genital opening of males flanked by 7–8 (rarely 6 or 9) setae on each side. Chelicera (Figs 63–64, 71–72, 78–79, 84) 2.1–2.4 (usually 2.3) (♂) or 2.1–2.3 (usually 2.2) (♀) times as long as broad, palm with 6 setae and 1 microseta laterally (1–2 microsetae in ♀ from Slovenia, Jama Virsnica); fixed finger with 10–15 teeth (distal tooth and third from distal larger and sharp) and 2–4 proximal microtubercles; movable finger with an isolated subapical tooth (di), 7–12 teeth proximally reduced in size and 2–5 proximal tubercles; gl ratio 0.50–0.54, rarely 0.55–0.57; spinneret usually prominent, rarely weakly prominent (Fig. 63) or absent (Fig. 64), in males, prominent and apically rounded in females (Figs 71–72, 79, 84); rallum with 11 blades; serrulae interior and exterior respectively with 13–15 and 15–17 blades. Coxal setae: pedipalp 5 (including 2 on manducatory process), I 3 + 3 marginal microsetae (rarely 4 + 2 or 4 + 4 marginal microsetae), II 4 (rarely 5), III 5 (rarely 6), IV 6 (rarely 7); coxa II with 7–21 (usually 11–15) coxal spines, coxa III with 4–9 (usually 5–7) coxal spines; intercoxal tubercle bisetose (with one seta in ♂ from Slovenia, Mozelj). Pedipalp: femur 5.6–7.5 (♂) or 5.5–7.4 (♀) times as long as broad; chela (Figs 65–66, 73–74, 80–81, 85–89) 5.3–6.1 (♂) or 4.6–5.4 (♀) times as long as deep; hand of chela 1.8–2.2 (♂) or 1.6–2.0 (♀) times as long as deep; fixed chelal finger with marked sigmoid curvature in lateral view, with 15–26 (♂) or 12–21 (♀) teeth with dental canals, usually with 1– 3 intercalary microdenticles in most interdental spaces, rarely without intercalary microdenticles in females; first tooth reduced, distad of it or halfway between et and it (rarely at level of it in females); 11–24 (♂) or 9–19 (♀) large, sharp, weakly reclined teeth increasing in size as far as halfway along the finger, then reduced in size towards the base; 1–5 (♂) or 1–3 (♀) small proximal teeth with rounded tips and dental canals; base of fixed finger with 0–5 microtubercles; fixed finger at level of it/est with 3 (rarely 2 or 4) teeth occupying 0.1 mm (distance between successive apices 0.034–0.055 mm) in males, or with 2 (rarely 3) teeth occupying 0.1 mm (distance between successive apices 0.046–0.080 mm) in females; tip of fixed finger with apical sensilla af1 ‾ 2, distal paraxial seta spatulate, enlarged in the middle and becoming sharper apically; movable finger strongly curved in lateral view and always shorter than fixed finger; dentition of movable finger highly variable, even among individuals of the same population (e.g. Figs 80–81), usually with 20–28 (♂) or 16–26 (♀) low, broad, reclined teeth increasingly reduced towards the finger base, reaching back nearly to sb, proximally followed, between sb and b, by 2–8 vestigial teeth revealed only by the presence of dental canals (♂ from Italy, Ovaro, shows the highest reduction in number of teeth on movable finger: 9 normal + 4 vestigial); distal half of movable finger usually with 11–15 low, broad, reclined teeth (Figs 65–66, 80–81), rarely with 6–10 triangular, reclined teeth, wider than high (Figs 80, 88–89); coupled sensilla pc usually halfway between st and sb, rarely closer to st than to sb; tip of movable finger with apical sensilla am1 ‾ 2; trichobothria as illustrated (Figs 65–66, 73–74, 80–81, 85–89); eb/esb/ist placed in a straight line or with ist distad of the line eb/esb; ratio of movable finger/hand 1.6–1.8 (♂) or 1.5–1.8 (♀); ratio of femur/movable finger 1.0–1.2 (♂♀); ratio of femur/carapace 1.2–1.7 (♂♀).
Measurements (in mm). Body length 1.5–2.1 (♂) or 1.5–2.4 (♀). Carapace 0.40–0.61 × 0.40–0.55 (0.36–0.51 anteriorly) (♂) or 0.45–0.68 × 0.45–0.63 (0.43–0.61 anteriorly) (♀). Chelicera 0.35–0.53 × 0.16–0.23 (♂) or 0.42–0.65 × 0.19–0.28 (♀); movable finger length 0.20–0.28 (♂) or 0.21–0.36 (♀). Pedipalp: femur 0.57–1.05 × 0.09–0.14 (♂) or 0.55–1.18 × 0.10–0.16 (♀); chela 0.86–1.51 × 0.14–0.25 (♂) or 0.88–1.74 × 0.18–0.35 (♀); hand length 0.31–0.55 (♂) or 0.30–0.64 (♀); fixed finger length (from tip to eb) 0.56–1.0 (♂) or 0.57–1.14 (♀); movable finger length 0.51–0.90 (♂) or 0.54–1.0 (♀).
Description of tritonymph. Integument weakly pigmented, hispid granulation less marked than in adults. Carapace 1.0–1.05 times longer than broad, anterior margin without epistome, denticles as in adults; anterior eyes with convex lens, posterior ones reduced to eye-spots; chaetotaxy 4:6:4:2:2 (18), length of anteromedian and anterolateral macrosetae respectively 0.095 and 0.050 mm. Chaetotaxy of tergites as in adults. Chaetotaxy of sternites II–X 5:(2)8(2):(1)7(1):7:6:6:6:6:7. Chelicera 2.15 times as long as broad, palm with 5 setae and 1 microseta laterally; fixed finger with 10 teeth, the distal two larger, the following proximally reduced in size; movable finger with an isolated subapical tooth (di) and 10 teeth; gl ratio 0.56; spinneret prominent and apically rounded, as in females; rallum with 9 blades; serrula exterior with 13 blades. Coxal setae: pedipalp 5 (including 2 on manducatory process), I 3 + 2 marginal microsetae, II 4, III 5, IV 5; coxa II with 6–7 coxal spines, coxa III with 4 coxal spines; intercoxal tubercle bisetose. Pedipalp: femur 4.9–5.5 times as long as broad; chela 4.85–5.5 times as long as deep; hand 1.7–2.0 times as long as deep; fixed finger with 11–13 reclined teeth with canals, with 1 intercalary microdenticle in most interdental spaces; distal tooth reduced, at level of it, 1–2 proximal teeth low, with rounded tips; fixed finger at level of it/est with 4 teeth occupying 0.1 mm (distance between successive apices 0.027 mm); distal paraxial seta as in adults; movable finger with 16 low, reclined teeth increasingly reduced towards the finger base, reaching back halfway between st and b; ratio of movable finger/hand 1.65–1.7; ratio of pedipalpal femur/movable finger 1.1; ratio of pedipalpal femur/carapace 1.2–1.3.
Measurements (in mm). Body length 1.0–1.2. Carapace 0.32–0.38 × 0.32–0.36. Chelicera 0.28–0.30 × 0.13–0.14, movable finger length 0.15–0.16. Pedipalp: femur 0.42–0.47 × 0.085–0.095; chela 0.64–0.68 × 0.12–0.14; hand length 0.22–0.24; fixed finger length (from tip to eb) 0.42–0.43; movable finger length 0.37–0.38.
Remarks. Chthonius raridentatus was described by Hadži (1930) from three specimens from as many Slovenian localities: one paratype of unstated sex from a cave on Mt Krim, south of Ljubljana [cave Velika Pasica, n. 75, 700 m a.s.l., Zgornji Ig, Rakitna (T. Novak in litt., 12.V.2013)]; one paratype of unstated sex from the cave Boštonova jama near Domžale [= cave Boštonova jama, no. 757, Krtina, Domžale (T. Novak in litt., 12.V.2013)]; and the female holotype—“mounted on a slide and used as the type ” (Hadži 1930, p. 134, translated)—from Frauenloch cave near Borovec, Kočevje [= cave Babja luknja pri Borovcu, no. 93, 650 m a.s.l., Borovec, Kočevje (T. Novak in litt., 12.V.2013)]. These specimens cannot be found in the Institute of Zoology at the Univ. Ljubljana, the Slovenian Academy of Sciences and Arts, or the Prirodoslovni muzej Slovenije in Ljubljana (T. Novak & T. Trilar in litt., 4.XII.2012). It is possible that they are in the collections of the Institute of Zoology of University of Belgrade, but a request for information from that institution went unanswered. Hadži’s (1930) description of C. raridentatus contains some contradictions concerning the sex of the type specimens: although males are not explicitly mentioned among the material examined, it is reported that the “male is a little smaller and thinner, while showing no obvious differences (except those in the genital apparatus) with respect to the female” (Hadži 1930, p. 140, translated); also, captions of figures 10–13 (p. 142) refer to a male, even if there are evident errors with reference to the respective drawings, notably fig. 11, which clearly shows a female. Hadži (1930) considered C. raridentatus to be related to C. troglodytes Redikorzev, 1928, C. heterodactylus and, in particular, C. cavernarum, and emphasized that the only difference between C. raridentatus and C. cavernarum lies in dentition of the movable chelal finger.
C. raridentatus was redescribed by Beier (1939b; repeated in Beier 1963) based on a female from the cave Boštonova jama (Biospeologica balcanica no. 757) and compared to C. subterraneus meuseli Beier, 1939. Subsequent authors (see Harvey 2013) added solely faunistic data for C. raridentatus.
The redescription of C. cavernarum, based on a male from “Breznohöhle ober Bischoflak” (= cave Migutovo brezno no. 5 near Škofja Loka), Slovenia, allowed Beier (1939b) to describe Chthonius ellingseni on specimens of both sexes from Austria (Annenheim am Ossiachersee and Villach), Slovenia (Mt Črna prst, near Bohinjska Bistrica), Montenegro (Herceg-Hovi) and, doubtfully, from Crete. These specimens were previously misidentified, apart from the one from Villach, as C. cavernarum (Beier 1928, 1931, 1932; see Remarks under C. cavernarum). The record from Crete (Beier 1931), based on material from the Roewer collection, is probably erroneous due to the mislabelling of material from the eastern Alps or Slovenia (Helversen & Martens 1972). Because of the uncertainty concerning the correct identification of syntypes from Montenegro and Crete, a lectotype female has been designated from Slovenia (Mt Črna prst, near Bohinjska Bistrica). Ćurčić (1974) mistakenly reported the cave “Završka jama” as type locality of C. ellingseni.
Chthonius ellingseni was subsequently redescribed by Beier (1963), incorporating some morphological characters from Beier (1931) and others, including part of the measurements and the figure of the chela, from Beier (1932). According to Beier (1963), the main differences between C. raridentatus and C. ellingseni are found in the carapacal chaetotaxy (setae of basal row: 2 versus m 2m, respectively) and the dentition of the chelal fingers (fixed finger without intercalary microdenticles in C. raridentatus, versus fixed finger with intercalary microdenticles in C. ellingseni; fixed finger with 13 sharp teeth proximad of est, versus fixed finger with 19 sharp teeth proximad et; movable finger with reduced dentition, versus movable finger with evident reclined teeth distad of sb).
Chthonius brandmayri was described by Callaini (1986), from one male and two females collected from Slavnik, Slovenian Istria, and compared with many species of Chthonius (Chthonius) with a sigmoid fixed chelal finger from southeastern Europe and Anatolia, particularly C. subterraneus meuseli, but no comparison was made with C. raridentatus or C. ellingseni. The main characters proposed by Callaini (1991) to differentiate C. brandmayri from C. s. meuseli are the presence of 2 setae on tergite I (4 in C. s. meuseli) and the dentition of the distal half of the movable chelal finger (with reclined teeth, wider than high, in C. brandmayri; with erect teeth, higher than wide, in C. s. meuseli).
Chthonius jalzici was described by Ćurčić (1988) from a single female from the cave Veternica Špilja, near Zagreb, Croatia, and compared to C. raridentatus, from which it was stated to differ mainly in the number of setae in the posterior row of the carapace: 2 macrosetae and one microseta on each side in C. jalzici, 2 macrosetae in C. raridentatus. The holotype of C. jalzici is not deposited in the Hrvatski Narodni Zoološki Muzej in Zagreb, contrary to Ćurčić (1988) (R. Ozimec in litt., 10.II.2014).
Examination of material from many Italian, Slovenian and Croatian localities, previously referred to C. raridentatus, C. ellingseni, C. brandmayri or C. jalzici (including types of C. ellingseni and C. brandmayri, and topotypes of C. jalzici), has revealed remarkable intra- and interpopulational variability in most of the characters used in previous comparisons of these species. This concerns in particular the carapacal and tergal chaetotaxy, the presence/absence of interdental microdenticles on the fixed chelal finger (Figs 73–74), the position of first distal tooth on the fixed chelal finger relative to trichobothria it–et (Figs 73–74, 80–81, 86–89) and the dentition of the distal half of the movable chelal finger (Figs 80–81). The widest dimensional range was found in a sample from Most na Soči (Slovenia): length of pedipalpal chela 1.01–1.38 (♂) 1.02–1.64 (♀) mm, length of movable chelal finger 0.61–0.825 (♂) 0.62–0.94 (♀) mm (Figs 65–66, 73–74). The following synonymies are therefore proposed: Chthonius (C.) ellingseni Beier, 1939, C. (C.) brandmayri Callaini, 1986 and C. (C.) jalzici Ćurčić, 1988 are all junior subjective synonyms of C. raridentatus Hadži, 1930 (n. syn.).
Chthonius raridentatus is a troglophilic species known from the eastern Venetian pre-Alps to southern Austria (Carinthia), Slovenia and northern Croatia (Fig. 90). Among the species of Chthonius of the C. heterodactylus group, C. raridentatus seems to be most closely related to the sympatric (rarely syntopic) C. cavernarum. Differences between C. raridentatus and C. cavernarum are given in the above key.
description: Published as part of Gardini, Giulio, 2014, The species of the Chthonius heterodactylus group (Arachnida, Pseudoscorpiones, Chthoniidae) from the eastern Alps and the Carpathians, pp. 101-137 in Zootaxa 3887 (2) on pages 124-135, DOI: 10.11646/zootaxa.3887.2.1, http://zenodo.org/record/228058