Revision of Australian species of the dung beetle genus Lepanus (Coleoptera: Scarabaeidae: Scarabaeinae): review of the L. ustulatus, L. storeyi, and L. nitidus species groups and description of eight new species Author Gunter, Nicole L. 0000-0002-8355-0862 ngunter@cmnh.org Author Weir, Thomas A. 0000-0002-8355-0862 ngunter@cmnh.org text Zootaxa 2021 2021-02-03 4923 1 1 66 journal article 8243 10.11646/zootaxa.4923.1.1 0a027439-c601-4533-bd03-f0e5d4442801 1175-5326 4496757 41D057A7-F855-47C2-A598-8000CFD83A5B Lepanus dichrous ( Gillet, 1925 ) ( Figs. 13D, H, J, L ; 14B ; 15B ; 16B ; 17A, C ) Panelus dichrous Gillet, 1925: 4 ; Paulian 1935: 119 , 1939: 70 . Lepanus dichrous : Matthews 1974: 111 . Type material digitally examined. Holotype : “Queensl. Mjöberg” (printed) / “Ma– landa” (printed) / “Typ.” (printed) / dichrous Gillet type .” (handwritten in black, with “ type ” in red ink) / “ Typus ” (printed) / “261 68” (“261” printed, “68” handwritten) / “3555 E92 +” (printed) / “ NHRS _ JLKB 000027315 ” (printed) [ 17°21’00”S , 145°35’00”E ] . Other material examined. 831 specimens from over 50 localities were also examined. Queensland : North Bell Peak summit, 17°05’19”S , 145°52’44”E , 15–16.ix.1981 (T55406–T55421 16 specimens in QM ) ; North Bell Peak summit, 17°05’19”S , 145°52’44”E , 16.ix.1981 (T55422 1 specimen in QM ) ; North Bell Peak summit, 17°05’19”S , 145°52’44”E , 19–22.xi.1990 (T55423–T55435, T55835 14 specimens in QM ) ; Robson Creek , Atherton [ 17°07’00”S , 145°37’00”E ], 17.i.1996 , R.L. Kitching (25-058288 1 specimen in ANIC ) ; Robson Creek , Atherton [ 17°07’00”S , 145°37’00”E ], 18.i.1996 , R.J. Kitching & H. Mitchell (25-058287 1 specimen in ANIC ) ; Robson Creek [ 17°08’00”S , 145°38’00”E ] (25-065483 1 specimen also as COL3003, 25-065484 1 specimen also as COL3004 specimens in ANIC ) ; Danbulla Forestry Reserve , 13 km NE of Yungaburra [ 17°10’00”S , 145°38’00”E ], 13.ii–6.iii.1987 , R.I. Storey & De Faveri (25-032545 1 specimen in QDPC ) ; Danbulla Forestry Reserve , 13 km NE of Yungaburra [ 17°10’00”S , 145°38’00”E ], 13.i–13.ii.1987 , R.I. Storey & De Faveri , (25-058283 2 specimens in ANIC ) ; Danbulla Forestry Reserve , 13 km NE of Yungaburra [ 17°10’00”S , 145°38’00”E ], 26.iii– 23.iv.1987 , R.I. Storey & De Faveri , (25-058285 1 specimen in ANIC ) ; Danbulla Forestry Reserve , 13 km NE of Yungaburra [ 17°10’00”S , 145°38’00”E ], 28.vii–3.ix.1987 , R.I. Storey & De Faveri , (25-058284 1 specimen in ANIC ) ; Danbulla Forestry Reserve , 13 km NE of Yungaburra [ 17°10’00”S , 145°38’00”E ], 6–26.iii.1987 , R.I. Storey & De Faveri (25-058286 2 specimens in ANIC ) ; Danbulla Scientific Reserve , 17°11’09”S , 145°39’36”E , 10.xii.1995 7.ii.1996 (T55306 specimen in QM ) ; Malanda Falls , 17°21’22”S , 145°35’13”E , 750 m , 23–24.i.2012 , G.B. Monteith (25-053935 17 specimens , 25-053936 24 specimens in ANIC ) ; Bartle Frere , Gourka road, 17°22’38”S , 145°44’34”E , 28–30.xi.1998 (T70869–T70877 9 specimens in QM ) ; Bartle Frere , west base, 17°22’47”S , 145°45’51”E , 19–20.xi.2001 (T111969–T111984 16 specimens in QM ) ; Bartle Frere , west base, 17°22’47”S , 145°45’51”E , 25.xi.1994 10.i.1995 (T55845–T55857 13 specimens in QM ) ; Bartle Frere , west base, 17°22’47”S , 145°45’51”E , 10.i–31.iii.1995 (T55858–T55864 7 specimens in QM ) ; Bartle Frere , west base, 17°22’47”S , 145°45’51”E , 7.iii–15.v.1995 (T55865–T55889 25 specimens in QM ) ; Bartle Frere , west base, 17°22’47”S , 145°45’51”E , 28–30.xi.1998 (T70860–T70868 9 specimens in QM ) ; Bartle Frere west base [ 17°23’00”S , 145°46’00”E ], 750 m , 28.xi.1998 , G.B. Monteith (25-060847 7 specimens also as COL587– 589 specimens in ANIC ) ; Bartle Frere west base [ 17°23’00”S , 145°46’00”E ], 750 m , 28–29.xi.1998 , G.B. Monteith (25-060850 2 specimens also as COL616– 17 in ANIC ) ; Malanda , 3.5 km S, 17°23’27”S , 145°35’46”E , 6–9.ii.1999 (T75465– T75470, T75485 7 specimens in QM ) ; Topaz , Pei Road ( Frith house in QM ) , 17°24’23”S , 145°41’05”E , 6.xii.1993 25.ii.1994 (T55220–T55226 7 specimens in QM ) ; The Crater National Park , 17°25’48”S , 145°29’24”E , 980 m , 18–19.ix.2010 , G.B. Monteith (25-053860 1 specimen also as COL1613, 25-059235 2 specimens , 25-060796 1 specimen , 25-060886 15 specimens also as COL1160 and COL1161, 25-053932 11 specimens in ANIC ) ; The Crater National Park , 17°25’48”S , 145°29’24”E , 980 m , 22–23.i.2012 , G.B. Monteith (25-053933 20 specimens in ANIC ) ; Thiaki , main rainforest, 17°25’55”S , 145°30’58”E , 990 m , 21–23.i.2012 , G.B. Monteith (25-053849 1 specimen in ANIC ) ; Thiaki cottage, 17°26’02”S , 145°30’40”E , 1000 m , 19–23.i.2012 , G.B. Monteith (25-053843 2 specimens in ANIC ) ; Topaz National Park , SE of Malanda [ 17°26’00”S , 145°43’00”E ], 720 m , 28.vii.1982 , S. Peck & J. Peck (25-058290 1 specimen in ANIC ) ; Hugh Nelson Range , 17°27’00”S , 145°29’00”E ], 1150 m , 6.iii–4.iv.1995 , P. Zborowski (25-058292 1 specimen in ANIC ) ; Longlands Gap , 3 ways road junction, 17°27’56”S , 145°28’30”E , 3–6.ii.1999 (T75487 specimens in QM ) ; BS1 Longlands Gap [ 17°28’00”S , 145°29’00”E ], 1150 m , 1.xii.1994 3.i.1995 , P. Zborowski (25-058291 1 specimen in ANIC ) ; BS1 Longlands Gap [ 17°28’00”S , 145°29’00”E ], 1150 m , 27.ii–26.iii.1996 , P. Zborowski (25-058293 1 specimen in ANIC ) ; Kenny Road , Atherton Tablelands , 17°28’13”S , 145°32’04”E , 25.xi.1994 10.i.1995 (T55296 specimens in QM ) ; Kenny Road , Atherton Tablelands , 17°28’13”S , 145°32’04”E , 10.i–31.iii.1995 (T55297–T55301 5 specimens in QM ) ; Kenny Road , Atherton Tablelands , 17°28’13”S , 145°32’04”E , 7.iii–15.v.1995 (T55302–T55303 2 specimens in QM ) ; Tumoulin Road , Evelyn , 17°29’52”S , 145°28’32”E , 5–11.ii.1998 (T57499 1 specimen , T60022–T60026 5 specimens in QM ) ; Millaa Millaa Falls , 17°29’53”S , 145°36’32”E , 5.v.1983 (T55230 1 specimen in QM ) ; Millaa Millaa Falls , 17°29’53”S , 145°36’32”E , 8–10.xii.2000 (T105998 1 specimen in QM ) ; Millaa Millaa Falls , 17°29’53”S , 145°36’32”E , 6– 9.ii.1999 (T75457–T75464 8 specimens in QM ) ; above Millaa Millaa Falls , 17°29’53”S , 145°36’47”E , 830 m , 23–25.i.2012 , G.B. Monteith (25-053937 32 specimens , 25-053938 41 specimens in ANIC ) ; above Millaa Millaa Falls , 17°29’56”S , 145°36’50”E , 830 m , 23–25.i.2012 , G.B. Monteith (25-053939 52 specimens , 25-053940 23 specimens in ANIC ) ; Millaa Millaa Falls (above) [ 17°30’00”S , 145°36’00”E ], 800 m , 18–20.iv.1999 , C. Reid & I. Reid (25-058295 4 specimens in ANIC ) ; Millaa Millaa Falls [ 17°30’00”S , 145°36’00”E ], 780 m , 11.v.1974 , R.I. Storey (25-058513 1 specimen in QM ) ; Millaa Millaa Falls [ 17°30’00”S , 145°36’00”E ], 780 m , 23–25.i.2012 , G.B. Monteith (25-053934 25 specimens in ANIC , 25-055612 6 specimens in ANIC ) ; Millaa Millaa Falls via Millaa Millaa [ 17°30’00”S , 145°37’00”E ], 21–22.iv.1973 , R.I. Storey (25-032538 2 specimens in QDPC , 25-058514 5 specimens in QM ) ; Millaa Millaa Falls via Millaa Millaa [ 17°30’00”S , 145°37’00”E ], 830 m , 15–16.ix.2010 , G.B. Monteith (25-059233 2 specimens , 25-060794 35 specimens , 25-060857 5 specimens in ANIC also as COL1104– 1108) ; Millaa Millaa Falls via Millaa Millaa [ 17°30’00”S , 145°37’00”E ], 760 m , 10.iv–4.v.1990 , R.I. Storey & K.H. Halfpapp (25-032548 2 specimens in QDPC ) ; Millaa Millaa Falls via Millaa Millaa [ 17°30’00”S , 145°37’00”E ], 760 m , 13.iii–10.iv.1990 , R.I. Storey & K.H. Halfpapp (25-032541 1 specimen in QDPC ) ; Millaa Millaa Falls via Millaa Millaa [ 17°30’00”S , 145°37’00”E ], 760 m , 15.viii–12.ix.1990 , R.I. Storey & K.H. Halfpapp (25-032544 3 specimens in QDPC ) ; Millaa Millaa Falls via Millaa Millaa [ 17°30’00”S , 145°37’00”E ], 760 m , 4.i–7.ii.1990 , R.I. Storey & K.H. Halfpapp (25-032540 2 specimens in QDPC ) ; Millaa Millaa Falls via Millaa Millaa [ 17°30’00”S , 145°37’00”E ], 760 m , 7.ii–13.iii.1990 , R.I. Storey & K.H. Halfpapp (25-032539 3 specimens in QDPC ) ; Kjellberg Road turnoff, 17°31’57”S , 145°36’27”E , 25.xi.1994 10.i.1995 (T55317 1 specimen in QM ) ; Kjellberg Road turnoff, 17°31’57”S , 145°36’27”E , 7.iii–15.v.1995 (T55318–T55328 11 specimens in QM ) ; Kjellberg Road turnoff, 17°31’57”S , 145°36’27”E , 10.i–31.iii.1995 (T55329–T55338 10 specimens in QM ) ; Kjellberg Road turnoff, 17°31’57”S , 145°36’27”E , 6–9.ii.1999 (T75443–T75446, T75451–T75456 10 specimens in QM ) ; Kjellberg Road , State Forest , boundary [ 17°32’00”S , 145°35’00”E ], 920 m , 6–9.ii.1999 , G.B. Monteith & D. Cook (25-060841 11 specimens in ANIC ) ; Mount Fisher , Kjellberg Road , 17°32’34”S , 145°33’31”E , 1.xii.1993 25.ii.1994 (T55339– T55340 2 specimens in QM ) ; Kjellberg Road , State Forest boundary, 17°32’09”S , 145°34’39”E , 6–9.ii.1999 (T75436–T75442, T75447–T75450 11 specimens in QM ) ; Wooroonooran National Park , 17°34’06”S , 145°42’21”E , 500 m , 9–14.ix.2004 , L. Masner (CMNEN-00030220–000030221 2 specimens in CMNC ) ; Henrietta Creek , Palmerston National Park [ 17°34’00”S , 145°45’00”E ], 22–24.iv.1973 , R.I. Storey (25-058512 4 specimens in QM , CMNEN-00030222 1 specimen in CMNC ) ; Beatrice River , 2 km S, 17°34’35”S , 145°41’25”E , 1.xii.1993 25.ii.1994 (T55304–T55305 2 specimens in QM ) ; Palmerston , Scenic Road , 17°34’44”S , 145°41’53”E , 650 m , 15–16.ix.2010 , G.B. Monteith (25-060795 15 specimens in ANIC ) ; Palmerston Forestry turnoff, 17°35’20”S , 145°42’14”E , 700 m , 15–16.ix.2010 , G.B. Monteith (25-059234 1 specimen , 25-060885 15 specimens in ANIC also as COL1157, COL1159) ; Maalan State Forest , Ravenshoe Road, 17°35’23”S , 145°35’40”E , 7.iii–15.v.1995 (T55104–T55133 30 specimens in QM ) ; Maalan State Forest , Ravenshoe Road, 17°35’23”S , 145°35’40”E , 10.i–31.iii.1995 (T55134– T55152 19 specimens in QM ) ; Maalan State Forest , Ravenshoe Road, 17°35’23”S , 145°35’40”E , 25.xi.1994 10.i.1995 (T55153–T55186 34 specimens in QM ) ; Maalan State Forest , Ravenshoe Road, 17°35’23”S , 145°35’40”E , 6–9.ii.1999 (T75471–T75484 14 specimens in QM ) ; Maalan Road , 2 km S Palmerston Highway , 17°35’56”S , 145°41’58”E , 7.iii–15.v.1995 (T55079–T55089 11, T55103 specimens in QM ) ; Maalan Road , 2 km S Palmerston Highway , 17°35’56”S , 145°41’58”E , 10.i. 1–7.iii.1995 (T55090–T55092 3 specimens in QM ) ; Maalan Road , 2 km S Palmerston Highway , 17°35’56”S , 145°41’58”E , 26.xi.1994 10.i.1995 (T55093–T55102 10 specimens in QM ) ; Henrietta Creek , Palmerston National Park , 17°35’58”S , 145°45’28”E , 30.xi.1992 15.iv.1993 (T111989 1 specimen in QM ) ; Palmerston National Park , west boundary, 17°33’36”S , 145°41’15”E , 1.xii.1993 25.ii.1994 (T55836– T55839 4 specimens in QM ) ; Mount Father Clancy , 800 m , 17°35’58”S , 145°38’04”E , 6–14.xii.1988 (T55840– T55844 5 specimens in QM ) ; road to Maalan Palmerston National Park [ 17°36’00”S , 145°40’00”E ], 23–24.iv.1978 , R.I. Storey (25-058511 17 specimens in QM , CMNEN–00030223 1 specimen in CMNC ) ; Rosina Creek , 14 km SE Millaa Millaa [ 17°36’00”S , 145°43’00”E ], 720 m , 24.vi–2.viii.1982 , S. Peck & J. Peck (25-058289 4 specimens in ANIC ) ; Palmerston National Park [ 17°37’00”S , 145°47’00”E ], 1.iv.1968 , E. Matthews (25-057885 4 specimens in ANIC ) ; South Johnstone Forestry Camp , 5 km W, 17°38’12”S , 145°40’15”E , 1.xii.1993 25.ii.1994 (T55214– T55217 4 specimens in QM ) ; Vine Creek , Majors Mountain , 17°40’58”S , 145°32’02”E , 4–6.ii.1999 (T75494, T76145, T76152 3 specimens in QM ) ; Vine Creek , Majors Mountain , 17°40’58”S , 145°32’02”E , 15–18.iv.1999 (T79112 1 specimen in QM ) ; Charmillin Creek crossing, 17°42’03”S , 145°31’21”E , 2–5.xii.1998 (T70886–T70888 3 specimens in QM ) ; Charmillin Creek crossing, 17°42’03”S , 145°31’21”E , 5–11.ii.1998 (T60017–T60021, T60425–T60433 14 specimens in QM ) ; Charmillin Creek , 17°42’54”S , 145°31’34”E , 880 m , 18–19.ix.2010 , G.B. Monteith (25-060856 2 specimens in ANIC ) ; Ravenshoe State Forest , Tully Falls Road [ 17°43’00”S , 145°33’00”E ], 15.xi–12.xii.1987 , A. Walford-Huggins (25-058294 2 specimens in ANIC ) ; Tully Falls State Forest , 18 km SSW Ravenshoe [ 17°46’00”S , 145°31’00”E ], 730 m , 1.x–5.xi.1987 , R.I. Storey & Dickinson (25-058272 5 specimens in ANIC , 25-032546 1specimen in QDPC ) ; Tully Falls State Forest , 18 km SSW Ravenshoe [ 17°46’00”S , 145°31’00”E ], 730 m , 20.iv–15.v.1988 , R.I. Storey & Dickinson (25-058274 specimens in ANIC , 25-032572 1 specimen in QDPC ) ; Tully Falls State Forest , 18 km SSW Ravenshoe [ 17°46’00”S , 145°31’00”E ], 730 m , 20.iv–25.v.1988 , R.I. Storey & Dickinson (25-058269 2 specimens in ANIC ) ; Tully Falls State Forest , 18 km SSW Ravenshoe [ 17°46’00”S , 145°31’00”E ], 730 m , 3.iii–20.iv.1988 , R.I. Storey & Dickinson (25-058280 2 specimens in ANIC ) ; Tully Falls State Forest , 18 km SSW Ravenshoe [ 17°46’00”S , 145°31’00”E ], 730 m , 5.i–7.ii.1988 , R.I. Storey & Dickinson (25-058276 2 specimens in ANIC ) ; Tully Falls State Forest , 18 km SSW Ravenshoe [ 17°46’00”S , 145°31’00”E ], 730 m , 7.xii.1987 7.i.1988 , R.I. Storey & Dickinson (25-058279 1 specimen in ANIC ) ; Tully Falls State Forest , 18 km SSW Ravenshoe [ 17°46’00”S , 145°31’00”E ], 730 m , 7.xi–7.xii.1987 , R.I. Storey & Dickinson (25-058270 1 specimen in ANIC ) ; Tully Falls State Forest , 11 km SSW Ravenshoe [ 17°47’00”S , 145°34’00”E ], 900 m , 5.i.1988 7.ii.1988 , R.I. Storey & Dickinson (25-058275 1 specimen in ANIC ) ; Tully Falls State Forest , 11 km SSW Ravenshoe [ 17°47’00”S , 145°34’00”E ], 900 m , 5.xi–7.xii.1988 , R.I. Storey & Dickinson (25-058282 2 specimens in ANIC ) ; Tully Falls State Forest , 9.5 km SSW Ravenshoe [ 17°47’00”S , 145°34’00”E ], 1000 m , 3.iii–20.iv.1988 , R.I. Storey & Dickinson (25-058281 1 specimen in ANIC ) ; Tully Falls State Forest , 9.5 km SSW Ravenshoe [ 17°47’00”S , 145°34’00”E ], 1000 m , 7.xii.1987 7.i.1988 , R.I. Storey & Dickinson (25-058278 2 specimens in ANIC ) ; Tully Falls State Forest , 9.5 km SSW Ravenshoe [ 17°47’00”S , 145°34’00”E ], 1000 m , 7.i–9.ii.1988 , R.I. Storey & Dickinson (25-058273 3 specimens in ANIC ) ; Tully Falls State Forest , 9.5 km SSW Ravenshoe [ 17°47’00”S , 145°34’00”E ], 1000 m , 9.ii–3.iii.1988 , R.I. Storey & Dickinson (25-058277 1 specimen in ANIC ) ; Tully Falls State Forest , via Ravenshoe [ 17°47’00”S , 145°34’00”E ], 13.v.1974 , R.I. Storey (25-032536 1 specimen in QDPC , 25- 058510 1 specimen in QM ) ; Tully Falls State Forest , via Ravenshoe [ 17°47’00”S , 145°34’00”E ], 20–21.iv.1973 , R.I. Storey (25-032537 1 specimen in QDPC , 25-058509 11 specimens in QM ) ; Tully Falls State Forest [ 17°47’00”S , 145°34’00”E ], 11–12.xi.1976 , R.I. Storey (25-058271 4 specimens in ANIC ) ; Tully Falls , 30 mi NE Tully [ 17°47’00”S , 145°34’00”E ], 2500 feet , 5.vi.1972 , G.B. & S.R. Monteith (25-058505 1 specimen , 25-058506 1 specimen , 25-058507 1 specimen , 25-058508 1 specimen in QM [ paratypes of Lepanus nitidus ]) ; Tully Falls , Ravenshoe [ 17°47’00”S , 145°34’00”E ], 20.xi.1977 16.iv.1978 , R.I. Storey (25-032547 1 specimen in QDPC ) . Description. Colour ranging from uniformly light brown to shiny dark brown or with pronotum having orange tinges. Holotype is shiny dark brown with mainly orange pronotum. Antennal club with apical antennomere (3 rd ) yellowish white and basal 2 antennomeres (1st, 2nd) grey. Total length: 3.3–3.8 mm . Holotype measurements (taken from image): total length 3.8 mm ; elytral width 2.6 mm . Male. Head: Width to length ratio 29: 21. Broad V-shape between clypeal teeth. Surface without setae and finely punctate. Dorsal part of eyes moderate, separated by an interocular space approximately 10–11 times eye width (21: 2). Prothorax: Pronotum smooth, nitid, with very fine punctation towards the anterior and finely punctate posteriorly. Hypomeral surface finely reticulate. Hypomeral stria absent. Pronotum width to length ratio 47: 28. Elytra: Smooth and nitid. Intervals very finely punctate and with extremely fine setae. Apical area nitid with striae 1 and usually 2 superficial, appearing as a single line. Ratio of length of the elytra along suture to maximum elytral width 58: 64. Legs: as per L. nitidus species group. Ridge on ventral side of protibia not joined to second tooth by a fine carina. Abdomen: Rest of pygidium surrounding central disc with fine punctures, each bearing a seta; finely reticulate only towards the basal angles, and nitid otherwise. Pterothorax: Lateral lobe of metaventrite finely punctate, setose and more or less finely reticulate. Medial lobe of metaventrite very finely punctate, smooth and nitid between metacoxae and with a very small tubercle, located closer to metacoxae than mesocoxae. Male genitalia: Aedeagus with left paramere excised apically. Female. As per L. nitidus species group. Metaventral tubercle absent. Distribution. Known from rainforest of the Australian Wet Tropics (WET) IBRA bioregion [Zones 7, 8, 9, 10, 11, and 12] from around Danbulla to Ravenshoe ( Fig. 17A ), with all specimens from high elevations ( 650–1150 m ). Within the BIOCLIM model ( Fig. 17C ), there are some grids with suitable climate from which L. dichrous has not yet been recorded, however the absence of records may reflect the limited sampling due to accessibility of highelevation forests in these grids. Comments. Most specimens have been collected in dung or mushroom-baited pitfall traps or in flight intercept traps. Lepanus dichrous was one of the most abundant species collected in the food preference study of Ebert et al. (2019) with 203 specimens collected from all lures except both fermenting fruit baits. Both macropod and fowl dung baits collected approximately 25% of the total specimens each ( n = 49 and 52 respectively), while vertebrate carrion ( n = 26) and invertebrate carrion ( n = 22, 9, 21) were more attractive than cucumber ( n = 12) and mushroom ( n = 12) ( Ebert et al. 2019 ). These results support the generalist feeding habits of the species. Specimens from Charmillin Creek, Millaa Millaa Falls, Palmerston National Park, and Mount Hypipamee National Park were sequenced for this study (see Fig. 1 ). They form a close relationship with each other, but the relationship within the L. nitidus species group could not be resolved. In Bayesian analysis, it was recovered at the base of the clade containing L. vangerweni and L. carbinensis while in maximum likelihood analysis it was recovered at the base of the clade containing L. nitidus and L. kulki . Neither relationship was strongly supported (BA PP = 81, ML BS = 28).