Lepanus villosus Matthews, 1974

Gunter, Nicole L. & Weir, Thomas A., 2019, Revision of Australian species of the dung beetle genus Lepanus (Coleoptera: Scarabaeidae: Scarabaeinae): description of 10 new species from the L. australis, L. monteithi, and L. villosus species groups, Zootaxa 4695 (3), pp. 201-252 : 239-247

Lepanus villosus Matthews, 1974

( Figs 16A, 16C , 17D , 18D , 19D )

Lepanus villosus Matthews, 1974: 118 .

Types examined: Holotype: ♂ “ Palmerston N.P., Q. 1.iv.68, E. Matthews ” / “ Lepanus villosus Det. 1972 M, E.G. Matthews, HOLOTYPE ” / “ ANIC Database No. 25 057986” in ANIC [17°34’0”S, 145°45’0”E] . Paratypes: “Palmerston N.P., Q. 1.iv.68, E. Matthews ” / “ Lepanus villosus Det. 1972 M, E.G. Matthews, PARATYPE” / “ ANIC Database No. 25 057987” 2 specimens ♀ in ANIC [17°34’0”S, 145°45’0”E], ; “Tully, N.Q., 12.x.1969, R.J. Huppatz ” / “ Lepanus villosus Det. 1972 M, E.G. Matthews, PARATYPE” / “ ANIC Database No. 25 057988” 1 specimen ♀ in ANIC [17°36’0”S, 145°55’0”E] ; “QLD. <50m Seymour Ra., 10/vii1971, Taylor Feehan” / Berlesate anic 376, rainforest 17.26x146.0” / “ Lepanus villosus Det. 1972 , E.G. Matthews, PARATYPE” / “ ANIC Database No. 25 057989” 1 specimen ♂ in ANIC [17°26’0”S, 146°0’0”E] ; “ Gillies H’way, N.Q. 2/10. GB.” / “ PARATYPE ” / “EM.117.” / “ Lepanus villosus Det. 1972 , E.G. Matthews, PARATYPE” / “ ANIC Database No. 25 057990” 1 specimen ♂ in ANIC [17°12’0”S, 145°42’0”E] ; “ QLD. 750m Koombooloomba , 4/vii/1971 Taylor Feehan ” / “Ber- lesate ANIC 259 rainforest” / “ Lepanus villosus Det. 1972 , E.G. Matthews, PARATYPE” / “ ANIC Database No. 25 057991” 1 specimen ♂ in ANIC [17°49’0”S, 145°33’0”E] ; “ Davies Creek , Atherton Tbld, N.Qld, 10 vii 1970, J.H. Connell ” / “ Lepanus villosus Det. 1972 M, E.G. Matthews, PARATYPE” (25-058496 1 specimen ♀, 25-058497 1 specimen ♂, 25-058498 1 specimen ♂, 1 specimen ♀ (on same card), 25-058499 1 specimen ♀, 25-058500 1 speci- men ♂ in QM) [16°59’0”S, 145°33’0”E] .

Other material examined. 532 specimens from 81 localities were examined. Cape Tribulation transect Site 5A, 16°4’41”S, 145°27’0”E, 1–31.i.1983, (T44339 1 specimen in QM) ; Cape Tribulation transect Site 5A, 16°4’41”S, 145°27’0”E, 5–9.i.1983, (T44480–T44482 3 specimens in QM) ; Cape Tribulation transect Site 5A, 16°4’41”S, 145°27’0”E, 20–23.iv.1983, (T44483–T44484 2 specimens in QM) ; 1.5 km Eby, N Mount Sorrow , [16°5’0”S, 145°27’0”E], 25.iii.1984, A. Calder & T.A. Weir (25-059588 2 specimens in ANIC) ; Thompson Creek, ( Grove ), 16°6’31”S, 145°26’25”E, 12.ii–15.iii.1998, (T85668 1 specimen in QM) ; Thompson Creek, ( Grove ), 16°6’31”S, 145°26’25”E, 15.iii–7.v.1998, (T88828 1 specimen in QM) ; Cooper Creek, ( Grove ), 16°9’10”S, 145°24’19”E, 11.i–10.ii.1998, (T73113–T73119 7 specimens in QM) ; Cooper Creek, ( Grove ), 16°9’10”S, 145°24’19”E, 10.ii– 16.iii.1998, (T85663–T85667, T85669, T85670, T85672–T85673, T88823–T88825 12 specimens in QM) ; Cooper Creek, ( Grove ), 16°9’10”S, 145°24’19”E, 16.iii –7.v.1998, (T88819–T88821 3 specimens in QM) ; Cooper Creek, ( Grove ), 16°9’10”S, 145°24’19”E, 11.ix–24.x.1998, (T88822, T88826 2 specimens in QM) ; Pimm’s Block, ( Grove ), 16°11’20”S, 145°24’18”E, 13.iii–8.v.1998, (T88829, T88831–T88833 4 specimens in QM) ; Pimm’s Block, ( Grove ), 16°11’20”S, 145°24’18”E, 10.ix–24.x.1998, (T88830, T88834 2 specimens in QM) ; Hutchinson Creek, ( Grove ), 16°13’4”S, 145°23’57”E, 13.i–11.ii.1998, (T73120–T73121 2 specimens in QM) ; Hutchinson Creek, ( Grove ), 16°13’4”S, 145°23’57”E, 11.ii–14.iii.1998, (T85671 1 specimen in QM) ; Hutchinson Creek, ( Grove ), 16°13’4”S, 145°23’57”E, 14.iii–8.v.1998, (T88827 1 specimen in QM) ; Alexandra Bay , [16°12’0”S, 145°28’0”E], 24.vi.1971, R.W. Taylor & J. Feehan, (25-057996 1 specimen in ANIC) ; Lync-Haven, Daintree area , 16°12’7”S, 145°24’32”E, 35 m, 2–10.xii.2012, T.A. Weir & G.B. Monteith, (25-057655 1 specimen in ANIC) ; Lync-Haven, Daintree area , 16°12’7”S, 145°24’32”E, 35 m, 6–9.xii.2012, G.B. Monteith (25-057653 1 specimen in ANIC) ; Thornton Range near Daintree , [16°14’0”S, 145°24’0”E], 200 m, 12–18.vii.1982, S. & J. Peck (25-059578 6 specimens in ANIC) ; Buchanan Creek, ( Grove ), 16°14’39”S, 145°25’54”E, 12.iii–8.v.1998, (T88818 1 specimen in QM) ; Humbug Landing, Daintree , 16°15’13”S, 145°20’54”E, 14–18.v.1998, (T66779–T66780 2 specimens in QM) ; Mount Gorton , 16°57’0”S, 145°53’0”E, 18.xi.1993, (T44315–T44321 7 specimens in QM) ; Mount Murray Prior , 16°56’0”S, 145°51’0”E, 7.xii.1995 (T44324 1 specimen in QM) ; Mount Murray Prior , 16°56’0”S, 145°51’0”E, 31.x.1995 – 8.xii.1995, (T44419–T44423 5 specimens in QM) ; Mount Murray Prior , 16°56’0”S, 145°51’0”E, 7–8.xii.1998, (T70933–T70954 21 specimens in QM) ; Chujeba Peak summit, 16°56’14”S, 145°39’19”E, 16.xii.1989, (T44340 1 specimen in QM) ; Davies Creek Road via Mareeba , [17°1’0”S, 145°35’0”E], 9.x–17.x.1976, R.I. Storey (25-059595 3 specimens in ANIC, 2 specimens in QDPC) ; Davies Creek Road, via Mareeba , 17°2’33”S, 145°36’51”E, 26.xi.1985, (T44341 x 1 in QM) ; Davies Creek Road, via Mareeba , 17°2’33”S, 145°36’51”E, 4–13.xii.1988, (T44414–T44415 2 specimens in QM) ; Isley Creek , upper, 17°2’41”S, 145°39’57”E, 29–30.xi.1993, (T44330– T44338 9 specimens in QM) ; Mount Tiptree , [17°3’0”S, 145°37’0”E], 13.vii.1984, R.B. Halliday (25-059585 1 specimen in ANIC) ; Mount Tiptree , 17°3’0”S, 145°38’0”E], 730 m, 29.vi.1971, R.W. Taylor & J. Feehan SAMA25- 027824 2 specimens in SAMA ); North Bell Peak summit, 17°5’19”S, 145°52’44”E, 16.ix.1981, (T44328 1 specimen in QM) ; North Bell Peak summit, 17°5’19”S, 145°52’44”E, 22.xi.1990, (T44342–T44345 3 specimens in QM) ; North Bell Peak summit, 17°5’19”S, 145°52’44”E, 20–22.xi.1990, (T44365–T44370 6 specimens in QM) ; Lamb Range , 6 km NNE Tinaroo Falls, 17°6’40”S, 145°33’58”E, 1159 m, 16.xi.2009, G.B. Monteith & Turco, (25- 060851 3 specimens in ANIC aslso as COL960- COL962) ; Danbulla State Forest Robson Creek Road 1.5 km N of road to Tinaroo , [17°7’0”S, 145°37’0”E], 27.iv–2.v.1998, Miledge,G., (25- 060767 1 specimen in AM) ; Danbulla S.F., 13 km NE of Yungaburra [Danbulla Forestry Reserve], [17°10’0”S, 145°38’0”E], 13.i–13.ii.1987, R.I. Storey & De Faveri, (25-059594 1 specimen in QDPC) ; Danbulla State Forest , 13 km NE of Yungaburra [Danbulla Forestry Reserve], [17°10’0”S, 145°38’0”E], 23.iv–27.v.1987, R.I. Storey & De Faveri, (25-059604 1 specimen in ANIC) ; Danbulla State Forest , 13 km NE of Yungaburra [Danbulla Forestry Reserve], [17°10’0”S, 145°38’0”E], 23.vi–28.vii.1987, R.I. Storey & De Faveri, (25- 032825 1 specimen in QDPC) ; Danbulla State Forest , 13 km NE of Yungaburra [Danbulla Forestry Reserve], [17°10’0”S, 145°38’0”E], 28.vii–3.ix.1987, R.I. Storey & De Faveri, (25- 032826 1 specimen in QDPC) ; Cathedral Fig , 17°10’52”S, 145°39’26”E, 10.xii.1988, (T44326 1 specimen in QM) ; Graham Range , summit, 17°16’24”S, 145°57’58”E, 1.xi–8.xii.1995, (T44404–T44405 2 specimens in QM) ; Graham Range , 350 m, 17°16’37”S, 145°57’48”E, 1.xi–8.xii.1995, (T54656–T54657 2 specimens in QM) ; Boar Pock- et Road , approximately 8 km N of Gillies Highway [Gillies Road], [17°12’0”S, 145°42’0”E], 13.xii.1969, J.G. Brooks, (25-057998 1 specimen in ANIC) ; Boar Pocket Road , approximately 8 km N of Gillies Highway [Gillies Road], [17°12’0”S, 145°42’0”E], 9.xii.1969, J.G. Brooks, (25-057997 1 specimen in ANIC) ; Boar Pocket Road 3–9 mi, 17°11’57”S, 145°40’8”E, 2500 feet, 8.iv.1974, J.G. Brooks, (CMNEN-00030586-00030588 in CMNC) ; Bellenden Ker Range, 0.5 km S Cable Tower 7, [17°16’0”S, 145°55’0”E], 500 m, 25–31.x.1981, Earthwatch, Queensland Museum , (25- 032830 1 specimen in QDPC, 25-059582 x 5 in ANIC) ; Bellenden Ker Range , 1 km S Cable Tower 6, 17°16’0”S, 145°53’0”E], 500 m, 17–24.x.1981, Earthwatch, Queensland Museum, (25-057999 2 specimens in ANIC) ; Bellenden Ker Range , 1 km S Cable Tower 6, 17°16’0”S, 145°53’0”E], 500 m, 25–31.x.1981, Earthwatch, Queensland Museum, (25-059583 1 specimen in ANIC) ; Lake Barrine National Park , [17°16’0”S, 145°39’0”E], 7.vii.1984, R.B. Halliday (25-059587 1 specimen in ANIC) ; Wright Creek, Lake Eacham , [17°16’0”S, 145°37’0”E], 750 m, 31.vii.1982, S. & J. Peck (25-058000 2 specimens in ANIC) ; Wright Creek, Lake Eacham , [17°16’0”S, 145°37’0”E], 720 m, 26–30.vii.1982, S. & J. Peck (25-059579 2 specimens in ANIC) ; Baldy Mountain Road Atherton , [17°17’0”S, 145°24’0”E], 12.xii.1976, R.I. Storey (25- 032828 1 specimen in QDPC) ; Baldy Mountain near Atherton [Mount Baldy], [17°17’0”S, 145°27’0”E], xii.1977 – i.1978, R.I. Storey (25- 032827 2 specimens in QDPC) ; Eacham National Park [Lake Eacham National Park], [17°17’0”S, 145°37’0”E], 760 m, 21.iii.1975, R.W.Taylor (25-057994 1 specimen in ANIC) ; Bartle Frere, Gourka Road , 17°22’38”S, 145°44’34”E, 28–30.xi.1998, (T70929–T70932 4 specimens in QM) ; Bartle Frere, West Base , 17°22’47”S, 145°45’51”E, 10.i–31.iii.1995, (T44447–T44452 6 specimens in QM) ; Bartle Frere, West Base , 17°22’47”S, 145°45’51”E, 7.iii–15.v.1995, (T44453–T44464, T55904 11 specimens in QM) ; Bartle Frere, West Base , 17°22’47”S, 145°45’51”E, 25.xi.1994 – 10.i.1995, (T44465–T44479 15 specimens in QM) ; Lamins Hill , 17°22’56”S, 145°42’7”E, 1.xii.1993 – 25.ii.1994, (T44413 1 specimen in QM) ; Bartle Frere W Base , [17°23’0”S, 145°46’0”E], 750 m, 28.xi.1998, G.B. Monteith (25- 060849 1 specimen in ANIC) ; Mount Bartle Frere W side [Bartle Frere South Peak], 17°24’0”S, 145°49’0”E], 700 m, 30.vii.1982, S. & J. Peck (25-059580 2 specimens in ANIC) ; Mount Bartle Frere W side [Bartle Frere South Peak], [17°24’0”S, 145°49’0”E], 700 m, 30.vii–4.viii.1982, S. & J. Peck (25-059581 1 specimen in ANIC) ; Hugh Nelson Range , 21 km S of Atherton, [17°25’0”S, 145°28’0”E], 12.x–5.xi.1983, R.I. Storey & Brown (25-059601 1 specimen in ANIC) ; Hugh Nelson Range , 21 km S of Atherton, [17°25’0”S, 145°28’0”E], 13.iii–1.v.1984, R.I. Storey & Brown (25-059603 1 specimen in QDPC) ; Hugh Nelson Range , 21 km S of Atherton, [17°25’0”S, 145°28’0”E], 21.vi–26.vii.1984, R.I. Storey & Brown (25-059602 x 1 in ANIC) ; Hugh Nelson Range , 21 km S of Atherton, [17°25’0”S, 145°28’0”E], 5.xi–1.xii.1983, R.I. Storey & Brown (25- 032829 1 specimen in QDPC) ; Topaz, Hughes Road, ( Breeden house), 17°26’4”S, 145°42’2”E, 1.vii–30.ix.1993, (T44426 1 specimen in QM) ; 21 km S of Atherton , 17°26’41”S, 145°28’35”E, 5–14.xii.1988, (T44416 1 specimen in QM) ; Hugh Nelson Range , [17°27’0”S, 145°29’0”E], 1150 m, 3.vii–1.viii.1995, P. Zborowski (25-058038 1 specimen in ANIC) ; Josephine Falls, Mount Bartle Frere [Bartle Frere], [17°27’0”S, 145°52’0”E], 10.vii.1984, R.B. Halliday (25-059586 2 specimens in ANIC) ; Longlands Gap , 17°27’14”S, 145°28’26”E, 1120 m, 18–19.ix.2010, G.B. Monteith (25-059606 1 specimen, 25- 060866 3 specimens in ANIC also as COL1123-COL1125) ; tower near The Crater , 17°27’22”S, 145°29’13”E, 1235m, 21–23.i.2012, G.B. Monteith (25-055609 7 specimens in ANIC, 25-053847 2 specimens in ANIC) ; Tower S of Crater National Park, 17°27’23”S, 145°29’12”E, 25.xi.1994 – 10.i.1995, (T44346–T44351 6 specimens in QM) ; Tower S of Crater National Park, 17°27’23”S, 145°29’12”E, 10.i–31.iii.1995, (T44352–T44359 8 specimens in QM) ; Tower S of Crater National Park, 17°27’23”S, 145°29’12”E, 7.iii–15.v.1995, (T44360–T44364 5 specimens in QM) ; Longlands Gap , 17°27’46”S, 145°29’32”E, 30.x.1991 – 23.vii.1992, (T44389 1 specimen in QM) ; Millaa Millaa Falls (below), [17°29’0”S, 145°36’0”E], 760 m, 18–20.iv.1999, C. & I. Reid (25-059605 1 specimen in ANIC) ; Millaa Millaa Falls , 17°29’53”S, 145°36’32”E, 6–9.ii.1999, (T76229,T76231–T76232 3 specimens in QM) ; Kjellberg Road turnoff, 17°31’57”S, 145°36’27”E, 7.iii–15.v.1995, (T44380 1 specimen in QM) ; Mount Fisher, Kjellberg Road , 17°32’34”S, 145°33’31”E, 27.iv.1982, (T44322–T44323 2 specimens in QM) ; Mount Fisher, Kjellberg Road , 17°32’34”S, 145°33’31”E, 3.v.1983, (T44329 1 specimen in QM) ; Mount Fisher, Kjellberg Road , 17°32’34”S, 145°33’31”E, 27–29.iv.1982, (T44417–T44418, T44425 3 specimens in QM) ; Kjellberg Road, State Forest boundary, 17°32’9”S, 145°34’39”E, 6–9.ii.1999, (T76233 1 specimen in QM) ; Maalan State Forest, Ravenshoe Road , 17°35’23”S, 145°35’40”E, 10.i–31.iii.1995, (T44427–T44431 5 specimens in QM) ; Maalan State Forest, Ravenshoe Road , 17°35’23”S, 145°35’40”E, 7.iii–15.v.1995, (T44435–T44438 4 specimens in QM) ; Maalan State Forest, Ravenshoe Road , 17°35’23”S, 145°35’40”E, 25.xi.1994 – 10.i.1995, (T44439–T44446 8 specimens in QM) ; Maalan State Forest, Ravenshoe Road , 17°35’23”S, 145°35’40”E, 6–9.ii.1999, (T76219– T76227 9 specimens in QM) ; Henrietta Creek Palmerston National Park , [17°35’54”S, 145°45’30”E], 22–24.iv.1973, R.I. Storey (CMNEN-00030582-00030585 in CMNC) ; Millaa Millaa Falls , [17°30’0”S, 145°37’0”E], 760 m, 11.v.1974, R.I. Storey (25- 032831 1 specimen in QDPC) ; Millaa Millaa Falls , [17°30’0”S, 145°37’0”E], 760 m, 13. vii–15.viii.1990, R.I. Storey & H.K. Halfpapp, (25- 032818 3 specimens in QDPC) ; Millaa Millaa Falls , [17°30’0”S, 145°37’0”E], 760 m, 13.iii–10.iv.1990, R.I. Storey & H.K. Halfpapp, (25- 032816 3 specimens in QDPC) ; Millaa Millaa Falls , [17°30’0”S, 145°37’0”E], 760 m, 15.viii–12.ix.1990, R.I. Storey & H.K. Halfpapp, (25- 032819 2 specimens in QDPC) ; Millaa Millaa Falls , [17°30’0”S, 145°37’0”E], 760 m, 4.i–7.ii.1990, R.I. Storey & H.K. Halfpapp, (25- 032815 1 specimen in QDPC) ; Millaa Millaa Falls , [17°30’0”S, 145°37’0”E], 760 m, 6.vi–13.vii.1990, R.I. Storey & H.K. Halfpapp, (25- 032817 2 specimens in QDPC) ; BS2 Mount Fisher , [17°33’0”S, 145°32’0”E], 1150 m, 31.i–27.ii.1996, L. Umback (25-059610 1 specimen in ANIC) ; Mount Fisher , [17°33’0”S, 145°32’0”E], 1150 m, 1.xii.1994 – 3.i.1995, P. Zborowski (25-059613 1 specimen in ANIC) ; Mount Fisher , [17°33’0”S, 145°32’0”E], 1150 m, 2.viii–4.ix.1995, L. Umback (25-058007 3 specimens in ANIC) ; Mount Fisher , [17°33’0”S, 145°32’0”E], 1150 m, 2.vi–4.vii.1995, P. Zborowski (25-058036 1 specimen in ANIC) ; Mount Fisher , [17°33’0”S, 145°32’0”E], 1150 m, 2–30.xi.1995, L. Umback (25-058013 1 specimen in ANIC) ; Mount Fisher , [17°33’0”S, 145°32’0”E], 1150 m, 21.iii–5.iv.1995, P. Zborowski (25-058021 1 specimen, 25-058022 2 specimens, 25-058023 2 specimens, 25-058024 3 specimens in ANIC) ; Mount Fisher , [17°33’0”S, 145°32’0”E], 1150 m, 3.i–4.ii.1995, P. Zborowski (25-059611 2 specimens in ANIC) ; Mount Fisher , [17°33’0”S, 145°32’0”E], 1150 m, 3.v–2.vi.1995, P. Zborowski (25-058034 3 specimens, 25-058035 4 specimens in ANIC) ; Mount Fisher , [17°33’0”S, 145°32’0”E], 1150 m, 3.x–2.xi.1995, L. Umback (25-058011 2 specimens in ANIC) ; Mount Fisher , [17°33’0”S, 145°32’0”E], 1150 m, 4.ii–21.iii.1995, P. Zborowski (25-059612 2 specimens in ANIC) ; Mount Fisher , [17°33’0”S, 145°32’0”E], 1150 m, 4.vii–2.viii.1995, P. Zborowski (25-058001 2 specimens, 25-058002 1 specimen in ANIC) ; Mount Fisher , [17°33’0”S, 145°32’0”E], 1150 m, 4.ix–3.x.1995, L. Umback (25-058008 1 specimen, 25-058009 9 specimens in ANIC) ; Mount Fisher , [17°33’0”S, 145°32’0”E], 1150 m, 5.iv–3.v.1995, P. Zborowski (25-058025 3 specimens, 25-058026 5 specimens, 25-058027 3 specimens, 25-058028 1 specimen, 25-058029 3 specimens, 25-058030 2 specimens in ANIC) ; Henrietta Creek Palmerston National Park [Plamerston National Park], [17°34’0”S, 145°45’0”E], 22–24.iv.1973, R.I. Storey (25- 032824 6 specimens in QDPC, 25-058501 8 specimens in QM) ; BS3 Massey Creek , [17°37’0”S, 145°34’0”E], 1000 m, 1.xii.1994 – 3.i.1995, P. Zborowski (25-059608 5 specimens in ANIC) ; BS3 Massey Creek , [17°37’0”S, 145°34’0”E], 1000 m, 3.i–4.ii.1995, P. Zborowski (25-059609 1 specimen in ANIC) ; BS3 Massey Creek , [17°37’0”S, 145°34’0”E], 1000 m, 4.ii–6.iii.1995, P. Zborowski (25-059614 3 spec- imens in ANIC) ; Massey Creek , [17°37’0”S, 145°34’0”E], 1000 m, 2.viii–4.ix.1995, L. Umback (25-058006 6 specimens in ANIC) ; Massey Creek , [17°37’0”S, 145°34’0”E], 1000 m, 2.vi–4.vii.1995, P. Zborowski (25-058037 1 specimen, 25-058039 5 specimens, 25-058040 5 specimens in ANIC) ; Massey Creek , [17°37’0”S, 145°34’0”E], 1000 m, 2.xi–30.xi.1995, L. Umback (25-058014 1 specimen, 25-058015 3 specimens in ANIC) ; Massey Creek , [17°37’0”S, 145°34’0”E], 1000 m, 3.v–2.vi.1995, P. Zborowski (25-058031 1 specimen, 25-058032 2 specimens, 25-058033 2 specimens in ANIC) ; Massey Creek , [17°37’0”S, 145°34’0”E], 1000 m, 3.x–2.xi.1995, L. Umback (25-058012 14 specimens in ANIC) ; Massey Creek , [17°37’0”S, 145°34’0”E], 1000 m, 4.vii–2.viii.1995, P. Zborowski (25-058003 6 specimens, 25-058004 3 specimens, 25-058005 2 specimens in ANIC) ; Massey Creek , [17°37’0”S, 145°34’0”E], 1000 m, 4.ix–3.x.1995, L. Umback (25-058010 9 specimens in ANIC) ; Massey Creek , [17°37’0”S, 145°34’0”E], 1000 m, 6.iii–5.iv.1995, P. Zborowski (25-058016 3 specimens, 25-058017 1 specimen, 25-058018 2 specimens, 25-058019 12 specimens, 25-058020 17 specimen in ANIC) ; South Johnstone Forestry Camp , 5 km W, 17°38’12”S, 145°40’15”E, 1.xii.1993 – 25.ii.1994, (T44403 1 specimen in QM) ; Vine Creek, Majors Mountain , 17°40’58”S, 145°32’2”E, 4–6.ii.1999, (T76234–T76236 3 specimens in QM) ; Vine Creek, Majors Mountain , 17°40’58”S, 145°32’2”E, 15–18.iv.1999, (T79131 1 specimen in QM) ; Charmillin Creek crossing, 17°42’3”S, 145°31’21”E, 8.xii.1989 – 5.i.1990, (T44373–T44374 23 specimens in QM) ; Charmillin Creek crossing, 17°42’3”S, 145°31’21”E, 5–11.ii.1998, (T62197 1 specimen in QM) ; Ravenshoe State Forest, Tully Falls Road, [17°43’0”S, 145°33’0”E], 10.x–15.xi.1987, A. Walford-Huggins (25-059584 2 specimens in ANIC) ; Ravenshoe State Forest, Tully Falls Road, [17°43’0”S, 145°33’0”E], 15.xi–12.xii.1987, A. Walford-Huggins (25-059589 3 specimens in ANIC) ; Tully Falls State Forest 18 km SSW Ravenshoe, [17°46’0”S, 145°31’0”E], 730 m, 1.x– 5.xi.1987, R.I. Storey & Dickinson, (25- 032820 1 specimen in QDPC, 25-059590 6 specimens in ANIC) ; Tully Falls State Forest 18 km SSW Ravenshoe, [17°46’0”S, 145°31’0”E], 730 m, 20.iv–25.v.1988, R.I. Storey & Dickinson, (25-059591 1 specimen in ANIC) ; Tully Falls State Forest 18 km SSW Ravenshoe, [17°46’0”S, 145°31’0”E], 730 m, 5.i–7.ii.1988, R.I. Storey & Dickinson, (25- 032822 1 specimen in QDPC, 25-059598 1 specimen in ANIC) ; Tully Falls State Forest 18 km SSW Ravenshoe, [17°46’0”S, 145°31’0”E], 730 m, 7.xii.1987 – 7.i.1988, R.I. Storey & Dickinson, (25-059597 2 specimens in ANIC) ; Tully Falls State Forest 18 km SSW Ravenshoe, [17°46’0”S, 145°31’0”E], 730 m, 7.xi–7.xii.1987, R.I. Storey & Dickinson, (25- 032821 2 specimens in QDPC, 25-059592 4 specimens in ANIC) ; Tully Falls State Forest 18 km SSW Ravenshoe, [17°46’0”S, 145°31’0”E], 730 m, 9.ii– 3.iii.1988, R.I. Storey & Dickinson, (25-059600 2 specimens in ANIC) ; Tully Falls State Forest 11 km SSW Ravenshoe, [17°47’0”S, 145°34’0”E], 900 m, 1.x–5.xi.1987, R.I. Storey & Dickinson, (25- 032823 1 specimen in QDPC, 25-059593 1 specimen in ANIC) ; Tully Falls State Forest 11 km SSW Ravenshoe, [17°47’0”S, 145°34’0”E], 900 m, 20.iv–25.v.1988, R.I. Storey & Dickinson, (25-059599 1 specimen in ANIC) ; Tully Falls State Forest 11 km SSW Ravenshoe, [17°47’0”S, 145°34’0”E], 900 m, 7.xii.1987 – 7.i.1988, R.I. Storey & Dickinson, (25-059596 2 specimens in ANIC) ; Tully Falls State Forest via Ravenshoe , [17°47’0”S, 145°34’0”E], 20–21.iv.1973, R.I. Storey (25-058502 1 specimen in QM) ; Tully Falls , 17°47’0”S, 145°33’27”E, 8.xii.1989 - 5.i.1990, (T44379 1 specimen in QM) ; Red Road turnoff on Tully Falls Road , 17°49’19”S, 145°32’42”E, 8.xii. 198 9–5.i.1990, (T44371 1 specimen in QM) ; Boulder Creek track, 200–600 m, 17°51’2”S, 145°54’29”E, 24–27.x.1983, (T44390-T44397 8 specimens in QM) ; Laceys Creek , [17°52’0”S, 146°4’0”E], 40 m, 11.vii.1971, R.W. Taylor J.E. Feehan, (25-057995 1 specimen in ANIC) ; Tully , [17°53’0”S, 145°55’0”E], 200 m, 17.x–22.xi.1994, (25-059607 1 specimen in ANIC) ; Tully River Crossing , 10 km S Koombooloomba Dam, 17°55’32”S, 145°37’38”E, 8.xii.1989 – 4.i.1990, (T44375–T44378 4 specimens in QM) ; Mount Tyson , 17°55’35”S, 145°54’34”E, 7.v.1983, (T44327 1 specimen in QM)

Description. Uniformly brown in colour. Antennal clubs yellow to white.

Total length: 2.2–2.8 mm. Holotype measurements: total length 2.4 mm, elytral width 1.5 mm.

Male. Head. Surface smooth and nitid, with dense ocellate punctures each bearing a seta. Very broad U-shaped between clypeal teeth that are short but not upturned. Genal angle weakly defined. Dorsal part of the eyes narrow, separated by an interocular space approximately 11 times the eye width (34:3). Head 1.5 x wider than long (45:30).

Prothorax: Pronotal surface smooth and nitid, with dense ocellate punctures each bearing an obvious posteiorly directed seta. Punctures along the basal margin horseshoe shaped differing from those on disc. Anterior angles obtuse. Basal margin slightly angulate. Hypomeral surface reticulate and punctate. Hypomeral stria about two thirds the lenth of hypomeron with a row of fine setae present between hypomeral stria and lateral edge. Pronotum 1.8

x wider than long (68:38).

Elytra: Intervals reticulate with fine meshes, and with a double row of recurved setae. Striae geminate, appearing double. A well-developed humeral callus between striae 6 and 7, and a basal depression between striae 4 and 5. Stria 8 ending basally in a deep pit and lateral carina indented at this point. Elytra 1.05 x wider than long (87:82).

Legs: Apical digit of foretibiae broad with a small comb of setae on inner apex and very small apical spur on outer edge. Metatibiae with a notch on the inner edge.

Abdomen: Pygidium flat, reticulate, punctate and with setae and with an overhanging basal edge. Abdominal ventrites reticulate right across.

Pterothorax: Medial lobe of metaventrite punctate and narrowly margined between the mesocoxae. Lateral lobe of metaventrite with ocellate punctures. Meso-metaventral suture very slightly arched. Mesoventrite smooth with punctation. Mesepimeron reticulate and punctate. Metanepisternum reticulate and punctate.

Hind wings: Fully developed.

Female. Metatibae without a notch on the inner edge, otherwise as per the L. villosus species group.

Distribution. Known from rainforests of the Australian Wet Tropics bioregion from Cape Tribulation to Tully ( Fig. 20 ). The majority of specimens have been collected from mid to high elevation forest (500–1200 m) but some are known from forest at sea level.

Comments. Almost 550 specimens were examined for this revision of which almost all have been collected in flight intercept traps, pitfall traps, or in berlesed leaf and log litter. The few specimens that have data for baited pitfalls indicate a preference of kangaroo dung, but other attractants include human faeces, carrion, mushroom, and fish baits. Ebert et al. (2019) only collected a single specimen of L. villosus in their 10-way food preference study and it came to a kangaroo dung baited pitfall trap. Specimens collected from Lambs Range and Longlands Gap (25- 060866 and 25- 060851) have been sequenced and included in the phylogenetic analyses ( Gunter & Weir 2017; Gunter et al. 2019). In the latter more comprehensively sampled study, L. villosus is recovered as sister to L. pseudovillosus new species, in a monophyletic clade that also contained L. vestitus .

Figures

FIGURE 16. Some diagnostic characters of the Lepanus villosus species group. A, lateral view of elytron of Lepanus villosus, white arrow points to deep pit at the base of stria 8; B, lateral view of elytron of Lepanus vestitus, note the absence of any deep pit; C, elytron of Lepanus villosus dotted line showing humeral callus between striae 6 and 7 and white arrow showing depression at base of striae 4 and 5; D, male metatibiae of Lepanus vestitus, notch on inner edge; E, male protibia of Lepanus pseudovillosus, showing typical configuration for the species group including the small comb of setae (grey arrow); F, close up of elytral interval surface of Lepanus reticulatus, showing reticulation of short longitudinal striations; G, close up of elytral interval surface of Lepanus vestitus showing reticulations of fine meshes.

FIGURE 17. Habitus of species within the Lepanus villosus species group. A, Lepanus pseudovillosus; B, Lepanus reticulatus; C, Lepanus vestitus; D, Lepanus villosus. All specimens in comparative scale. Scale bar 1 mm.

FIGURE 18. Left, dorsal, and right view of aedeagi of species within the Lepanus villosus species group. A, Lepanus pseudovillosus; B, Lepanus reticulatus; C, Lepanus vestitus; D, Lepanus villosus. All specimens in comparative scale. Scale bar 0.2 mm.

FIGURE 19. Pygidial configuration of species within the Lepanus villosus species group. A, Lepanus pseudovillosus; B, Lepanus reticulatus; C, Lepanus vestitus; D, Lepanus villosus. All specimens in comparative scale. Scale bar 1 mm.

FIGURE 20. Map of known records including predicted distribution estimated with BIOCLIM models for A, Lepanus villosus; B, Lepanus pseudovillosus. QLD, Queensland.