Anaspides richardsoni, Ahyong, 2016

Ahyong, Shane T., 2016, The Tasmanian Mountain Shrimps, Anaspides Thomson, 1894 (Crustacea, Syncarida, Anaspididae), Records of the Australian Museum 68 (7), pp. 313-364 : 336-346

Anaspides richardsoni sp. nov.

Figs 18–23 , 35D–F , 36

Anaspides tasmaniae . — Calman, 1897: 802 [“Lake Field” (= Mt Field) specimens only (OM Iv.1394, below)]. — Smith, 1909a: 64, 70 (Mt Read, Mt Field). — Williams, 1965b: 333–334, fig. 1, 2; 1965a: 106, tab. 6. —Swain & Reid, 1983: 163–171. —Richardson & Swain, 1989: 277, tab. 1, app. 1. —Jarman & Elliot, 2000: fig. 4 (clade E, part), tab. 1 (part, Central Plateau). —Gooderham & Tsyrlin, 2002: 73 (unnumbered colour figure). — Richter et al., 2002: 341, 347, fig. 31–33.

Anaspides spinulae . — O’Brien, 1990: frontispiece, pl. 1.

Anaspides sp. (telson ‘normal’ type). — Eberhard et al., 1991: 48 (Mole Creek caves).

Type material. (Mt Field). HOLOTYPE: AM P72839, ♂ (33 mm), between Newdegate Pass and Mawson Plateau, coll. J. Kunze, 27 Feb 1974 . PARATYPES: AM P72840, 1♀ ( TL 32 mm), between Newdegate Pass and Mawson Plateau, 42°40.2' S 146°33.6'E, coll. J. Kunze, 27 Feb 1974 ; AM P72841, 4♂♂ (23–27 mm), 5 juv. ♂♂ (17–21 mm), 4♀♀ (24–31 mm), 5 juv. ♀♀ (14–20 mm),between Newdegate Pass and Mawson Plateau, 42°40.2'S 146°33.6'E,coll. J. Kunze, 27 Feb 1974 ; ZRC 2016.0491 , 1♂ (30 mm), 1♀ (31 mm), between Newdegate Pass and Mawson Plateau, 42°40.2'S 146°33.6' E, coll. J. Kunze, 27 Feb 1974 ; TMAG G6174 , 1♂ (27 mm), 2 juv. ♂♂ (14–16 mm), 1♀ (34 mm), Mt Field ,small tarn above University ski lodge, 42°40.134'S 146°34.152' E, tarn in herbfield with dolerite rocky outcrops, dipnet, 1240 m asl, coll . R. Mollison , 27Apr 2011 ; TMAG G6363 , 1♀ (45 mm), midway between Newdegate Pass and Newdegate Tarn, 42°39.6'S 146°33.6' E, in small tarn, 1200 m asl, coll. I. Wilson & J. Ong, 25 Jan 1970 ; TMAG G6419 , 2♂♂ (25–27 mm), 21 juv. ♂♂ (11–21 mm), 1♀ (22 mm), 37 juv. ♀♀ (11–21 mm), Newdegate Pass , 42°39.6' S 146°33.0' E, sample 2, coll. I. Wilson & J. Ong, 25 Jan 1970 ; TMAG G6445 , 5♂♂ (20–21 mm), 6 juv. ♂♂ (13–19 mm), 2♀♀ (24–29 mm), 5 juv. ♀♀ (13–15 mm), Newdegate Pass , 42°39.6'S 146°33.0'E,sample 1,coll. I. Wilson & J. Ong, 25 Jan 1970 ; TMAG G6429 , 11 ♂♂ (22–32 mm), 3 juv. ♂♂ (18–21 mm), 25♀♀ (22–34 mm), 5 juv. ♀♀ (14–21 mm), near Sitzmark Lodge , Mt Field, 42°40.8'S 146°34.8'E, tarn, coll. C.Reid, 15–18 Oct 1974 ; USNM 1277685 , 1♂ (26 mm), 1 juv. ♂ (17 mm), 3♀♀ (25–30 mm), near Sitzmark Lodge , Mt Field, 42°40.8'S 146°34.8'E, tarn, coll. C.Reid, 15–18 Oct 1974 .

Other material examined. Gunns Plain to Black Bluff: AM P99296, 1 juv. ♂ (12 mm), Great Western Cave ( GP27 ), Gunns Plains karst area, NW Tasmania, 41°17.8'S 146°00.3'E, from riffle pool 250 m upstream, 1296-24, 109 m asl, coll. A. Clarke, 29 Dec 1996 ; QVM 10 :13975, 1♀ (23 mm), Paddy’s Lake,below Black Bluff, near Loongana , Nietta South , 41°27.2'S 145°57.6'E, 1070 m asl,coll . T. Hume , 5 May 1972 ; TMAG G6398 , 14♂♂ (20–24 mm), 1 juv. ♂ (16 mm), 32♀♀ (19–30 mm), 3 juv. ♀♀ (16–17 mm), Paddy’s Lake, Black Bluff , 41°27.2' S 145°57.6'E, 1070 m asl, coll. C. Binks, 25Apr 1979 ; TMAG G6169 , 1 juv.♂ (13 mm), 6 juv. ♀♀ (10–17 mm), Vale of Belvoir , 41°32.95'S 145°53.54'E, coll . R. Mollison , 16 Mar 2010 .

Deloraine: TMAG 14391 /G135, 2♂♂ (22–24 mm), Deloraine ,coll. C.King, Nov 1937 . Mole Creek (Form 1) : SAMA C8481 , 2 immature ♂♂ (16–21 mm), 2♀♀ (30–39 mm), 1 indet juv. (9 mm), about 200 m from cave entrance, Marakoopa Cave , 41°34.9'S 146°17.3'E, 490 m asl, small stream, BS0464,coll. E. Hamilton Smith, 19 Nov 1963 ; USNM 1277682 , 3♂♂ (29–34 mm), 3 juv. ♂♂ (12–16 mm), 3♀♀ (21–30 mm), 2 juv. ♀♀ (16–17 mm), Marakoopa Cave , 0.3 m, stn 87-253, coll . T. Iliffe , 27 Dec 1987 ; TMAG G646 , 1♂ (34 mm), 1♀ (45 mm), Long Creek , Marakoopa Cave ( MC120 ), 4 Dec 1991 ; QVM 10 :13973, 1♀ (33 mm), Marakoopa Cave II (MC-015), 41°34.9'S 146°17.3'E, coll. K. Crocker, 1 Jul 1981 ; TMAG G6458 , 2♂♂ (31–32 mm), 2♀♀ (32–33 mm), Lake Passage , Marakoopa Cave II ( MC15-1 ), 41°34.9' S 146°17.3'E, 490 m asl, coll. A. Goede, 21 Aug 1982 ; TMAG G6459 , 2♂♂ (26–28 mm), 3♀♀ (21–32 mm), Lake Passage , Marakoopa Cave II ( MC15-1 ), 41°34.9'S 146°17.3'E, 490 m asl, coll. A. Goede, 21 Aug 1982 ; TMAG G6462 , 2♀♀ (36–41 mm), Prohibition Cave , 41°35.6'S 146°19.5'E, coll . R. Eberhard , 19 Apr 1991 .

Mole Creek (Form 2): AM P73038, 1♂ (37 mm), Honeycomb Cave ( MC107 ), lower pool, dark zone, 41°36.0'S 146°24.4'E, 1096.03,coll. S. Bunton, 27 Oct 1996 ; ZSRO 863 a, 1♀ (44 mm), Honeycomb Cave,KS 34, Ana 16 A011, 27 Feb 2013 ; ZSRO 863 b, 1♀ (46 mm), Honeycomb Cave, KS 34, Ana 17 A011, 27 Feb 2013 ; USNM 1277681 , 1♂ (33 mm), 1 juv. ♂ (22 mm), 7♀♀ (32–55 mm), Honeycomb Cave , 0–0.5 m, stn 87- Pot ( MC207-12 ), 41°36.6'S 146°22.5' E, 520 m asl ; TMAG G6464 , 1♀ (41 mm), Kellys Pot ( MC207-14 ), 41°36.6'S 146°22.5'E, 520 m asl ; TMAG G6465 , 1♂ (29 mm), 2♀♀ (30–37 mm), Kellys Pot ( MC207-13 ), 41°36.6'S 146°22.5'E, 520 m asl ; TMAG G6460 , 1♀ (37 mm), Mole Creek caves, in water pool,coll . R.A. Rafferty , 28 Apr 1938 ; TMAG G6457 , 1♂ (36 mm), 2♀♀ (37–46 mm), Herbert’s Pot ( MC202 ), 41°36.9'S 146°23.3'E, 500 m asl , CV40 , coll. S. Eberhard, 6 Apr 1985 ; AM P56373, 1♂ ( TL 35 mm), 1♀ ( TL 35 mm), Wet Cave near Caveside, 41°36'S 146°25'E, C.92 T, moderate flowing stream, disappears before cave entrance, cave with few pools, coll. W. Ponder et al., 18 Jan 1982 .

Mole Creek (Form 3): SAMA C8482 , 1♀ (24 mm), 2 juv. (7–9 mm), unnamed cave, Sassafras Creek ,about 50 m from entrance, 41°33.7' S 146°21.9'E, 270 m asl,clear pool, mud bottom,BS0457, 18 Nov 1963 ; TMAG G6454 , 3♂♂ (23–32) mm), 1♀ (23 mm), 5 juv.♀♀ (13–20 mm), Kubla Khan Cave , 41°33.2'S 146°17.6' E,coll. S. Eberhard , R. Swain & A.Richardson, 23 Sep 1981 ; TMAG G6455 , 1♂ (26 mm), 1 juv.♂ (20 mm), 2♀♀ (26–28 mm), 3 juv. ♀♀ (10–21 mm), Kubla Khan Cave , 41°33.2'S 146°17.6'E, coll. S. Eberhard, A. Richardson & R. Swain , 25 May 1986 ; TMAG G6456 , 1♂ (29 mm), 2♀♀ (23–39 mm), Kubla Khan Cave , 41°33.2'S 146°17.6'E, in river, c. 500 m into cave, 320 m asl, coll. S. Eberhard, 12 Nov 1983 ; TMAG G6466 , 3♂♂ (25–30 mm), 3♀♀ (23–28 mm), Kubla Khan Cave ( MC1-30 ), 41°33.2'S 146°17.6'E ; USNM 1277679 , 1♂ (25 mm), Kubla Khan Cave , 0–0.5 m, stn 87-250, coll . T. Iliffe , 26 Dec 1987 ; USNM 1277683 , 3♂♂ (23–26 mm), 5♀♀ (28–40 mm), Mayberry area , Mole Creek, water cave, from sinkhole near farmhouse, baited traps, 0.2m, stn 87-251, coll . T. Iliffe , 27 Dec 1987 .

West Coast Range : TMAG G6411 , 1♂ (26 mm) , 3 juv. ♂♂ (16–20 mm) , 2 juv. ♀♀ (20–21 mm), tarn II, S of Lake Tyndall , 41°57.1'S 145°35.2'E, 980 m asl, coll. C.J. Binks & B. Knott, 16 Jan 1973 ; TMAG G6312 , 8♂♂ (23–25 mm) , 3 juv. ♂♂ (16–21 mm) , 8♀♀ (23–30 mm), 3 juv. ♀♀ (15–18 mm), tarn S of Lake Tyndall , 41°57.1'S 145°35.2, 980 m asl, coll. C.J. Binks & B. Knott, 16 Jan 1973 ; TMAG G6362 , 1 juv.♂ (18 mm) , 2 juv. ♀♀ (16–17 mm), Lake Sandra, Mt Murchison , 41°49.9'S 145°35.8'E, 940 m asl, 10 Dec 1973 ; TMAG G6385 , 1♀ (damaged, c. 26 mm), Lake Sandra, Mt Murchison , 41°49.9' S 145°35.8'E, 940 m asl,coll. W. Fulton, Nov 1983 ; TMAG G6392 , 1♀ (27 mm), N of Geikie, Tyndall Range , alpine plateau above climbing route and camp, 41°57.4'S 145°35.0'E, 1000 m asl, coll. C.J. Binks & B. Knott, 16 Jan 1973 .

Cradle Mountain Lake St Clair National Park: TMAG G6352 , 1♂ (21 mm), 1♀ (23 mm), Twisted Lake , Cradle Mountain, 41°40.22'S 145°58.09'E, 1116 m asl, coll. D. O’Brien, 8 Mar 1990 ; SAMA C6303 , 1 juv. ♂ (18 mm), top of Cradle Mountain , 41°40.8'S 145°57.0'E,small pool in creek, 4850 ft asl [1455 m],coll. A.Kowanko, Dec 1967 ; SAM C6304 , 2♀♀ (21–32 mm),top of Cradle Mountain , 41°40.8'S 145°57.0' E, small pool in creek, 4850 ft asl [1455 m],coll. A.Kowanko, Dec 1967 ; QVM 10 :13835, 1♂ (20 mm), Sutton’s Tarn , Cradle Mountain, 41°41.01'S 145°55.98'E, 1089 m asl, coll. Kingston, 1991 ; TMAG G6350 , 1♂ (26 mm), 1♀ (24 mm), Sutton’s Tarn , near Kitchen Hut, Cradle Mountain, 41°41.0'S 145°56.0'E, 1089 m asl, coll. D. O’Brien, 8 Mar 1990 ; TMAG G6351 , 2♀♀ (30–31 mm), Sutton’s Tarn , near Kitchen Hut, Cradle Mountain, 41°41.0'S 145°56.0'E, 1089 m asl, coll. D. O’Brien, 8 Mar 1990 ; TMAG G6381 , 2♂♂ (27–31 mm), 12♀♀ (21–32 mm), Mt Doris , tarn on S side, 41°52.6'S 146°02.7'E, 1170 m asl, coll. B. Knott, 12 May 1970 ; AM P72843, 1 juv. ♂ (21 mm), 1♀ (23 mm), Mt Ossa–Mt Doris Ridge , Mt Ossa, 41°52.2'S 146°04.8'E, 1255 m asl, coll. C. Sands ; AM P82858, 2♂♂ (23–24 mm), 4 juv. ♀♀ (14–21 mm), 8♀♀ (23–30 mm), 5 juv. ♀♀ (13–18 mm), S of Mt Doris,runnel crossing Mt Ossa track, 41°52.2'S 146°01.8' E,deep runnel in grassy lawn in alpine shrubbery,FW18,coll. A.Richardson & P.A. Serov, 28 Jan 1990 ; TMAG G6413 , 6♂♂ (24–29 mm), 6♀♀ (26–34 mm), 1 juv. ♀ (16 mm), near summit of Cathedral ( NE of summit), 41°53.4'S 146°07.2'E, c. 1350 m asl, tarn, small, clear & with rocky bottom, sample 6, coll. C.J. Binks & B. Knott, 31 Jan 1972 ; WAM C11772, 1♂ (21 mm), 3♀♀ (21–23 mm), Cradle Mountain district , coll. A. Connell, 1939 ; WAM C58160, 1 juv. ♂ (16 mm), 4♀♀ (25–39 mm), Cradle Mountain-Lake St Clair, National Park, 16 Oct 1947 .

Walls of Jerusalem National Park (non-“ spinulae ” form): TMAG G6448 , 28 juv. ♂♂ (13–18 mm), 49 juv. ♀♀ (12–19 mm), 1 indet juv.(5 mm), Jaffa Vale, at Dixon’s Hut, Walls of Jerusalem , 41°40.77' S 146°06.37'E, 1250 m asl,coll .? R. Swain ,1969 ; TMAG G6449 , 2♂♂ (25–28 mm), 2 juv. ♂♂ (18–19 mm), 4 juv. ♀♀ (15–17 mm), Jaffa Vale, at Dixon’s Hut, Walls of Jerusalem , 41°40.77'S 146°06.37' E, 1250 m asl, coll .? R. Swain , 1969 ; TMAG G6319 , 2♂♂ (23–29 mm), 1♀ (20 mm), 1 juv. ♀ (13 mm), pool 50 m W of Herod’s Gate Pool , 41°48.7'S 146°16.8'E, 1220 m asl,coll. S.Smith, 16 Apr 1990 ; TMAG G6380 , 6 juv. ♂♂ (13–16 mm), 8 juv. ♀♀ (12–19 mm), Lake Adelaide track between Fish Rock and Herod’s Gate , Walls of Jerusalem , 41°48.7'S 146°16.8'E, runnel, 1200 m asl, coll. B. Knott, 18 Nov 1971 ; TMAG G6368 , 5♂♂ (24–26 mm), 10 juv. ♂♂ (14–21 mm), 8♀♀ (25–35 mm), 9 juv. ♀♀ (11–20 mm), near Herod’s Gate, Walls of Jerusalem , 41°48.7'S 146°17.4'E, small creek, 1200 m asl, coll. B. Knott, 17 Nov 1971 ; TMAG G6446 , 2♂♂ (25–27 mm), 1 juv. ♂ (17 mm), 2♀♀ (26–30 mm), 10 juv. ♀♀ (14–18 mm), Zion Gate west, Walls of Jerusalem , 41°48.9'S 146°19.6'E, 1280 m asl, coll .? R. Swain , 1969 ; TMAG G6393 , 1♂ (24 mm), 10 juv. ♂♂ (11–18 mm), 2♀♀ (24–33 mm), 13 juv. ♀♀ (15–18 mm), Saddle through to east wall, Walls of Jerusalem , 41°48.9'S 146°19.2'E, 1230 m asl, coll. J. Bludhorn, 18 Nov 1971 ; TMAG G6418 , 4♂♂ (29–32 mm), 4 juv.♂♂ (16–23 mm), 2♀♀ (26–29 mm), 2 juv.♀♀ (13–18 mm), E side of Zion Gate, Walls of Jerusalem , 41°49.0'S 146°19.6'E, 1240 m asl, coll .? R. Swain , 1969 ; TMAG G6345 , 3♂♂ (21–22 mm), 1♀ (25 mm), 20 m below Gate of Chain, Walls of Jerusalem , 41°49.12' S 146°18.39' E, pool in pencil pines, 1300 m asl, coll. S. Smith, 16 Apr 1990 ; TMAG G6346 , 1♂ (21 mm), 3♀♀ (20–22 mm), 1 juv. ♀ (14 mm), Gate of Chain, 50 m below ridge, Walls of Jerusalem , 41°49.13' S 146°18.53'E, 1280 m asl, coll. S. Smith, 16 Apr 1990 ; TMAG G6318 , 2♂♂ (24–27 mm), 1 juv. ♂ (13 mm), 2♀♀ (26–30 mm), 100 m E of ridge, Gate of Chains, Walls of Jerusalem , 41°49.13' S 146°18.60'E, 1260 m asl, coll. S. Smith, 16 Apr 1990 ; TMAG G6440 , 1♂ (23 mm), 1 juv. ♂ (16 mm), 1♀ (23 mm), 6 juv. ♀♀ (12–16 mm),near Lake Ball, Walls of Jerusalem , 41°49.23'S 146°17.95'E, 1240 m asl, coll . V. Thorp , 3 Apr 1972 ; TMAG G6320 , ♂ (26 mm), 1 juv. ♂ (15 mm), 2♀♀ (25–29 mm), 1 juv. ♀ (16 mm), 200 m below Pool of Bethesda, Walls of Jerusalem , 41°49.34'S 146°17.81'E,runnel,#1, 1260 m asl, coll. S. Smith, 14 Apr 1990 ; TMAG G6348 , 6♂♂ (25–32 mm), 1♀ (32 mm), 50 m SE of Damascus Gate , 41°49.61'S 146°17.59'E, pool in grassland, coll . R. Smith , 1350 m asl, 15 Apr 1990 ; TMAG G6376 , 3 juv. ♂♂ (15–16 mm), 11 juv. ♀♀ (12–17 mm), Damascus Gate , 41°49.61'S 146°17.95'E, coll . R. Swain , 1969 ; TMAG G6422 , 1♂ (29 mm), 1 juv. ♂ (18 mm), 2♀♀ (26–28 mm), 6 juv. ♀♀ (12–15 mm), Damascus Gate, Walls of Jerusalem, runnel draining into Lake Calvine , 41°49.99' S 146°18.16'E, 1340 m asl, coll .? R. Swain , 1969 ; TMAG G6394 , 9 juv. ♂♂ (15–20 mm), 1♀ (24 mm), 8 juv. ♀♀ (13–18 mm),near Junction Lake immediately under Moraine retaining Lake Meston , Walls of Jerusalem , 41°55.35' S 146°11.52' E, bog drainage pool (chiefly Gleichenia ), “sample 2”, 950 m asl, coll. C. Binks & B. Knott, 31 Jan 1972 ; TMAG G6397 , 7 juv. ♂♂ (10–22 mm), 1♀ (29 mm), 5 juv. ♀♀ (17–21 mm), creek draining into Junction Lake, second of four or five drainage systems on broad plain from Lake Meston, Walls of Jerusalem , 41°55.35' S 146°11.52'E,clear pools under Nothofagus lined creek, “sample 3”, 950 m asl, coll. C. Binks & B. Knott, 31 Jan 1972 .

Walls of Jerusalem National Park (mixed “ spinulae ” and non-“ spinulae ” forms): TMAG G6396 , 4♂♂ (21–22 mm), 6 juv. ♂♂ (15–18 mm), 1♀ (24 mm), 2 juv. ♀♀ (17–20 mm), Zion Vale, Walls of Jerusalem , 41°48.64' S 146°18.83' E, pool, 1200 m asl , coll. R. Swain , 1969 ; TMAG G6379 , 1♂ (25 mm), 2 juv. ♂♂ (18–22 mm), 2♀♀ (24–25 mm), Maximal Creek, W of Herod’s Gate, Walls of Jerusalem , 41°48.62'S 146°15.94'E, 1150 m asl, coll. B. Knott, 16 Nov 1971 .

Walls of Jerusalem National Park (“ spinulae ” form): WAM C11771, 4 juv. ♂♂ (shrivelled, c. 13–16 mm), 1♀ (shrivelled, c. 19 mm), Jones Tarn , at foot of Western Wall, Walls of Jerusalem, 41°49.02'S 146°18.46'E, approx. 4200 feet [1260 m], 26 Apr 1935 ; TMAG G6347 , 4 juv. ♂♂ (9–12 mm), 7 juv. ♀♀ (11–15 mm), 100 m E of ridge, Gate of Chain, Walls of Jerusalem, 41°49.13'S 146°18.53'E, 16 Apr 1990 ; TMAG G6391 , 9 juv. ♂♂ (14–18 mm), 1♀ (27 mm), 11 juv. ♀♀ (13–17 mm),SE end Cloister Lagoon , 41°54.26'S 146°10.66'E,from small drainage pools in predominantly Gleichenia covered bog, sample 4, 1100 m asl, coll. C.J. Binks & B. Knott, 31 Jan 1972 ; TMAG G6434 , 7♂♂ (23–30 mm, 5♀♀ (24–31 mm),Pool of Bethesda,Walls of Jerusalem, 41°49.28' S 146°17.95'E, 1270 m asl, coll . R. Swain ,1969 ; TMAG G6349 , 2♂♂ (21–23 mm), 1 juv. ♂ (15 mm), 3♀♀ (26–27 mm), E edge Pool of Bethesda , 41°49.28'S 146°17.88'E, 1270 m asl, coll. S. Smith, 16 Apr 1990 ; TMAG G6427 , 1♂ (24 mm), 19 juv. ♂♂ (12–18 mm), 11 juv. ♀♀ (12–20 mm), 6 indet juv. (6–7 mm), 41°53.01'S 146°09.23' E,from dirty,deep pool system draining through pineapple grass into Lake Chalice , 1020 m asl, “sample 5”, coll. C. Binks & B. Knott, 31 Jan 1972 ; TMAG G6330 , 3♂♂ (22–23 mm), 1♀ (24 mm), [no label] Lake Chalice , 41°52.84'S 146°08.87'E, coll. W. Fulton, 2 Feb 1988 .

Western Lakes—Great Western Tiers: AM P57906, 1♀ (21 mm), Western Bluff , 41°37'S 146°17'E, 26 Jan 1964 ; QVM 10 :13977, 1♀ (36 mm), 1 juv. ♀ (18 mm), Jacks Lagoon , 5 km SW of Lake Mackenzie, 41°42.1'S 146°20.6' E,“abundant”,“no fish”, 1260 m asl, coll. E . V. Terry , 7 Apr 1989 ; WAM C58161, 3♂♂ (26–29 mm), 2♀♀ (25–26 mm), Ironstone Mountain , Northwestern Tiers, 41°42.8'S 146°28.5'E, from temporary creek, 100 m asl, coll. I. Gooch, 1946 ; QVM 10 :13978, 1♂ (23 mm), 2♀♀ (23–31 mm), Meander Falls , 41°44.1'S 146°30.3' E,coll. S. Merry, 1957 ; QVM 10 :49162 (“spinulae” form), 1♂ (23 mm), 5♀♀ (23–28 mm), Lake Johnny , 41°43.6'S 146°25.2'E, 1210 m asl,coll. W. Fulton, 9 Dec 1987 ; QVM 10 :49163 (“spinulae” form), 2♂♂ (18–20 mm), 5♀♀ (23–26 mm), Lake Halkyard , 41°44.2'S 146°18.9'E, 1210 m asl, coll. W. Fulton, 9 Dec 1987 ; QVM 10 :49164 (“spinulae” form), 2♂♂ (22–23 mm), 5♀♀ (20–30 mm), Lake Fox , 41°43.9'S 146°24.8'E, 1230 m asl, coll. W. Fulton, 9 Dec 1987 ; QVM 10 :49165, 3♂♂ (28–31 mm), 8♀♀ (24–29 mm), 3 juv. ♂♂ (20–21 mm), 15 juv. ♀♀ (11–19 mm), 300 m W of Lake Fox , 41°43.9'S 146°24.00'E, pool, 1210 m asl, coll. W. Fulton, 9 Nov 1987 ; QVM 10 :13980, 2 juv. ♂♂ (17–18 mm), 1 juv. ♀ (19 mm), stream on top of Western Tier near Lake Lucy Long, 41°42.2' S 146°26.0' E, 1190 m asl, coll. J. Simmons, 3 Mar 1963 ; TMAG G6438 , 3♂♂ (25–29 mm), 4 juv. ♂♂ (17–22 mm), 1♀ (28 mm), 7 juv. ♀♀ (14–21 mm), edge of Great Western Tiers near Pine Lake, 41°44.7' S 146°43.0'E, runnel draining opposite way, 1220 m asl, coll. B. Knott, 1970 ; WAM C58166, 1♀ (31 mm),creek north of Pine Lake ,coll. G.E.Nicholls, 27 Jan 1947 ; WAM C58165, 7♂♂ (19–23 mm), 4 juv. ♂♂ (17– 17 mm), 7♀♀ (19–27 mm), 4 juv. ♀♀ (16–18 mm), N end of Pine Lake , 41°44.2' S 146°42.2'E, from linked puddle, coll. G.E.Nicholls, 27 Jan 1947 ; TMAG G6443 , 1♂ (21 mm), 16 juv.♂♂ (16–20 mm), 8♀♀ (19–26 mm), 10 juv. ♀♀ (14–19 mm), Pine Lake , 41°44.6' S 146°42.0'E, 1190 m asl, coll . R. Swain , 22 Nov 1969 ; TMAG G6420 , 19♂♂ (20–24 mm), 9 juv.♂♂ (9–18 mm), 19♀♀ (18–22 mm), 6 juv. ♀♀ (14–19 mm), creeks entering Pine Lake , 41°44.6'S 146°42.0' E, coll . R. Swain , 3 Feb 1969 ; AM P97845, 3 juv. ♂♂ (7–12 mm), 15 juv.♀♀ (6–11 mm), Halfmoon Creek , roadcrossing below Pine Lake, 41°45.01'S 146°42.75'E, from rocky pools,hand sieves, 1159 m asl,TAS-516,coll. G.D.F.Wilson & S.J.Keable, 9 Mar 2001 ; TMAG 14371 /G115, 1♀ (29 mm), stream, 1 mile N of Rainbow Chalet, Great Lake , 41°46.0'S 146°33.7'E, 3000 ft asl [914m],coll. J. Pearson, Apr 1939 ; TMAG G1309 , numerous juv, Breona , 41°47' S 146°42'E, 1060 m, from well,coll. G.E.Nicholls, 6 Feb 1945 ; TMAG G1310 , 8 mature ♂♂ (18–24 mm), 6♀♀ (21–35 mm), Breona , from well, coll. G.E. Nicholls, 6 Feb 1945 ; TMAG G1314 , 1♂ (28 mm), 3 juv. ♂♂ (12–17 mm), 1♀ (26 mm), 2 juv. ♀♀ (11–15 mm), Breona , Great Lake, north-east, in creek supply hotel, coll. G.E.Nicholls, 23 Dec 1943 ; TMAG G1311 , 1♀ (19 mm) and numerous juv, Breona , from well behind Stewart’s house, coll. G.E. Nicholls, 6 Feb 1945 ; TMAG G1312 , 1♂ (20 mm) and numerous juv, Breona , from waterhole draining creek,coll. G.E.Nicholls, 6 Feb 1945 ; WAM C11773, 1♂ (c. 18 mm), 1 juv. ♂ (c.mm), 1♀ (22 mm), 3 juv.♀♀ (c. 10–16mm), Stewarts Water Hole , Breona, coll. G.E.Nicholls, 8 Jan 1946 ; WAM C58164, 1♂ (18 mm), 3 juv. ♂♂ (12–14 mm), 1♀ (21 mm), 7 juv. ♀♀ (6–15 mm), Breona , log behind (north) Stewart’s Cottage, 30 Jan 1947 ; WAM C11769, 2♂♂ (18–22 mm), 2 juv.♂♂ (13–16 mm), 1♀ (21 mm), 5 juv.♀♀ (7–20 mm), creeks at Breona , Great Lake, 41°47'S 146°42'E, coll. Mr Stewart et al., 25 Jan 1947 ; WAM C11774, 2♂♂ (21–23 mm), 1 juv. ♂ (17 mm), 1♀ (18 mm), 2 juv. ♀♀ (9–10 mm), Breona , wells,coll. G.E.Nicholls, 6 Feb1945 ; WAM C58158, 27 juv.♂♂ (11–18 mm), 43 juv. ♀♀ (8–17 mm), indet juv. (6–7 mm), Breona , Great Lake, puddles near haulage, coll. G.E. Nicholls, 28 Jan 1947 ; WAM C11770, 8♂♂ (24–31 mm), 8♀♀ (27–42 mm), Brandons [=Brandum], Great Lake, 41°49.6'S 146°40.5' E, 1034 m asl, Easter 1946 ; WAM C11775, 1 juv.♂ (16 mm, 1 juv.♀ (16 mm), Reynolds Neck , Great Lake, 41°51.1'S 146°41.4'E, 1034 m asl,coll. A.Pike,1935 ; TMAG G1630 , 4♂♂ (22–23 mm), 1♀ (23 mm), near Shannon Lagoon , 41°59.6'S 146°44.1'E, pool in swamp, 1040 m asl, coll. J.W. Evans, 13 Dec 1936 ; TMAG G131 , 2 juv. ♀♀ (15–21 mm), mouth of creek, Great Lake , coll. A.W.G.Powell, 29 Mar 1937 ; NMV J42435 , 2♀♀ (25.0–36.0 mm), Great Lake , coll. F.E. Burbury, 1942 ; NMV J42443 , 1♀ (24.0 mm), Great Lake , coll. F.F.Wilson, Jan 1933 ; AM P99298, 2 juv.♂♂ (10–11 mm), 12 juv.♀♀ (8–14 mm), Sandbanks Tier , 41°50'28.69"S 146°51'10.95"E, 1150 m asl, stream from under boulderfield, coll. S. Jarman ; QVM 10 :22606, 1♀ (34 mm), Lake River Valley , coll. C. Spencer, 1997 ; WAM C58156, 33♂♂ (18–31 mm), 48 juv. ♂♂ (12–18 mm), 23♀♀ (19–34 mm), 174 juv. ♀♀ (9–18 mm), 5 miles N of Breona , 41°42.2'S 146°43.4'E, creek,coll. G.E.Nicholls, 7 Feb 1945 ; NMV J42444 , 4 juv. ♀♀ (12.0–16.0 mm), Ouse River , 5 miles W of Miena, coll. A. Neboiss, 28 Feb 1967 ; USNM 1277684 , 1 juv. ♂ (15 mm), Ouse River , 5 miles W of Miena,coll. A.Neboiss, 28 Feb 1967 ; TMAG G6369 (“spinulae” and non-“spinulae” forms), 8♂♂ (19–22 mm), 4 juv. ♂♂ (15–17 mm), 10 ♀♀ (18–27 mm), 1 juv.♀ (15 mm), Ouse River ,semi creek, 22 Nov 1969 ; AM P11873, 1♀ (24 mm), 4–4½ miles S of Miena, in spring off road between Miena and Bothwell, about 20 yards to left of road near old notice on tree, marked “water”, 41°59.8'S 146°47.2'E, coll. J. Waterhouse, c. 1930 ; TMAG G6428 , 16♂♂ (23–33 mm), 7♀♀ (32–35 mm), small stream running into Ouse River on Lake Auga Road , 41°52.61'S 146°36.28'E, drift sample, 1150 m asl, coll. P. Davies, Nov, 1985 ; AM P14158, 6 juv. (slide preparations),near Little Pine Lagoon , 42°00.0'S 146°35.7'E, 1000 m asl, stream, coll. W.D. Williams, 5 Feb 1963 ; QVM 10 :13981, 17♂♂ (19–25 mm), 1 juv. ♂ (18 mm), 7♀♀ (22–26 mm), 5 juv. ♀♀ (10–18 mm), about 10 miles from Great Lake on Missing Link Road , between Great Lake and Bronte Park, from little stream on S side of McKenzies Tier leading into Little Pine River, 42°02.2'S 146°33.0' E, coll. Sgt. McIntyre, 1958 viaArthur Fleming ; SAMA C8446 , 4 juv.♂♂ (13–18 mm), 2♀♀ (36–41 mm), 9 juv. ♀♀ (9–21 mm), about 10 km N of Bronte Park , 42°04'S 146°29'E, from small stream above Pine Tier Dam, coll. W. Zeidler, 16 Jul 2001 ; AM P99297, 1♂ (21 mm), Pine Tier Lagoon , near shore, 42°04.38'S 146°29.30' E, coll. S. Richter & C. Wirkner, 28 Feb 2006 ; ZSRO 372 , 1 juv. ♂ (11 mm), 7 juv. ♀♀ (6–12 mm), Pine Tier Lagoon , near shore, 42°04.38'S 146°29.30'E, coll. S. Richter & C. Wirkner, 28 Feb 2006 ; ZSRO, 1♀ (38 mm), tributary of Travellers Rest River , coll. A. Richardson, 4 Aug 2016 ; TMAG G6321 , 2♀♀ (21–28 mm), Clarence Lagoon , 40–50 m from outflow, 42°05.12'S 146°18.87' E, 980 m asl, coll. D. O’Brien & R. Kirkwood , 30 Mar 1990 ; TMAG G6439 , 2♂♂ (24–29 mm), 5♀♀ (21–36 mm), 6 juv.♀♀ (11–12 mm), Clarence Lagoon , 42°05.2' S 146°19.2'E, on scuba, 980 m asl, coll . R. Mawbey 11 Jan 1984 ; TMAG G6442 , 3♂♂ (21–23 mm), 2♀♀ (26–29 mm), Clarence Lagoon , 42°05.12' S 146°18.87' E, stomach of brook trout, 980 m asl, coll. P.A. Tyler, 5 Aug 1970 ; TMAG G6355 , 1♀ (32 mm), Clarence Lagoon , 42°05.2'S 146°19.2'E, outflow creek, 980 m asl, in FBA net, coll . R. Mawbey , 13 May 1985 ; TMAG G6359 , 3♂♂ (22–24 mm), 1 juv. ♂ (12 mm), 1♀ (30 mm; non-“spinulae” form), 1 juv.♀ (13 mm), Clarence Lagoon , near start of Clarence River, 42°05.1'S 146°18.9' E, rocky shoreline, 980 m asl, coll. W. Fulton, 22 May 1976 ; TMAG G6367 (non-“spinulae” form), 4♂♂ (22–26 mm), 6 juv. ♂♂ (18–21 mm), 2♀♀ (23–24 mm), 4 juv. ♀♀ (10–21 mm), 2 indet juv. (7 mm), moorland pools draining into Clarence Lagoon , 42°04.42'S 146°18.80'E, 980 m asl, coll . R. Swain , 21 Dec 1982 ; TMAG G6353 , 15 juv.♂♂ (12–15 mm), 13 juv.♀♀ (11–14 mm), Silver Plains Creek , Alma Tier, 42°07.19'S 147°04.57'E, 920 m asl, coll . R. Mawbey , 11 Mar 1979 ; TMAG G6360 , 1♂ (30 mm), 1♀ (36 mm), “The Groves”, Oatlands, 2 Mar 1969 ; AM P14160, 1♂ (slide preparation), 1♀ (slide preparation),near Tarraleah , 42°17.2'S 146°26.3'E, 470 m, stream, coll. W.D. Williams, 7 Feb 1963 .

Mt Field National Park: OM Iv. 1394, 4♂♂ (21–30 mm), 3♀♀ (21–32 mm), Mt Field, 4000 ft asl [1200 m] ; AM P97846, 1♀ (38 mm), 1 juv.♀ (16 mm), Tarn Shelf , Rodway Range, coll. M.S.Moulds, 3 Feb 1992 ; AM P14159, 4 specimens (slide preparations c. 16–18 mm),near Lake Dobson , 42°41'S 146°36'E, stream, coll. W.D. Williams, 28 Jan 1963 ; AM P73058, 1♂ (28 mm), Wombat Moor , Mt Field, spring head of small spring fed sphagnum swamp on hillside, 42°40.98' S 146°36.48'E,TAS-484,coll. G.Wilson & S. Keable, 4 Mar 2001 ; AM P73059, 1♂ (25 mm), 3♀♀ (19–25 mm), Wombat Moor , Mt Field, spring head of small spring fed sphagnum swamp on hillside, 42°40.98'S 146°36.48'E,TAS-484,coll. G.Wilson & S. Keable, 4 Mar 2001 ; AM P97844, 1 juv.♂ (15 mm), 9 juv. ♀♀ (6–15 mm), spring flowing out of hillside near ski lodge at bottom of walking track skirting Lake Dobson and continuing to Mt Field West, 42°41.11'S 146°35.44'E,pool among water mosses, with phreatoicidean isopods,hand sieves, 1037 m asl,TAS-483,coll. G.D.F.Wilson & S.J.Keable, 4 Mar 2001 ; TMAG G6405 , 10 ♂♂ (25–33 mm), 5♀♀ (28–38 mm), Mawson Plateau , Mt Field, 42°41.4' S 146°35.1'E, 1270 m asl, coll. D. Hamilton, Apr 1970 ; TMAG G6371 , 1 juv. ♂ (18 mm), Mt Field West , 42°39.5' S 146°30.7'E, 1400 m asl, coll. M. Fenton, 20 Jan 1971 ; TMAG G6401 , 2♂♂ (27–29 mm), 5♀♀ (26–38 mm), Newdegate Pass , Mt Field, 42°39.5'S 146°33.3'E,coll. C. Reid, 23 May 1974 ; TMAG G6441 , 6♂♂ (24–29 mm), 2 juv. ♂♂ (9–10 mm), 6♀♀ (24–33 mm), between Mackenzie tarn and James Tarn, 42°40.2'S 146°33.7' E, small tarn, 1180 m asl, coll . T. Walker , Mar 1972 ; TMAG G133 , 1♀ (29 mm), Lake Fenton , 42°40.4'S 146°36.9'E, 1006 m asl, coll. Pott, Apr 1936 ; TMAG G6375 , 3♂♂ (29–31 mm), 2 juv. ♂♂ (15–25 mm), 1♀ (33 mm), Mt Mawson , 42°41.7'S 146°35.8' E, 1280 m asl, coll. D. Hamilton, Apr 1970 ; TMAG G6444 , 5♂♂ (25–29 mm), 1 juv. ♂ (19 mm), 5♀♀ (26–42 mm), 5 juv.♀♀ (8–22 mm), Mawson Plateau , 42°41.4'S 146°35.1'E, pool with mud bottom, 1267 m asl, coll . T. Walker , Mar 1972 ; TMAG G6415 , 2 juv. ♀♀ (13 mm), Mawson Plateau , 42°41.4'S 146°35.0'E, pool with rocky bottom, coll . T. Walker , Mar 1972 ; TMAG G6384 , 4♂♂ (25–30 mm), 7♀♀ (26–36 mm), 8 juv. ♀♀ (12–20 mm), Mawson Plateau , 42°41.4'S 146°35.0' E, pool with rocky bottom, coll . T. Walker , Mar 1972 ; TMAG G6432 , 1♂ (25 mm), 1 juv. ♂ (20 mm), 1♀ (31 mm), 6 juv. ♀♀ (8–23 mm), 1 indet juv. (7 mm),W side of Rodway Range , 42°40.2'S 146°32.4' E, pool with muddy bottom, coll . T. Walker , Mar 1972 ; TMAG G6373 , 3 indet juv. (7 mm), W side of Rodway Range , 42°40.2' S 146°32.4' E, pool with muddy bottom, coll . T. Walker , Mar 1972 ; QVM 10 :43547, 28♂♂ (20–32 mm), 18♀♀ (20–41 mm), 1 juv. ♀ (13mm), Mawson Plateau , Mt Field, 42°41.4'S 146°35.1'E, 1270 m asl,coll. C.Reid, 1975 ; TMAG G6314 , 7♀♀ (25–36 mm), Robert Tarn , Tarn Shelf, Mt Field, 42°40.7'S 146°34.2' E, 1200 m asl,coll . T. Walker , Mar 1972 ; TMAG G6372 , 10 ♂♂ (25–32 mm), 8 juv. ♂♂ (16–21 mm), 9♀♀ (20–32 mm), 11 juv. ♀♀ (10–22 mm), Robert Tarn , Tarn Shelf, Mt Field, 42°40.7' S 146°34.2'E, 1200 m asl, sample 1, tube 1(2), coll. I. Wilson & J. Ong, 25 Jan 1970 ; TMAG G6408 , 1♂ (31 mm), 1♀ (38 mm), Mt Field , midway between Newdegate Pass and Newdegate Tarn, 42°39.6'S 146°33.6'E, in small tarn, 1200 m asl, coll. I. Wilson & J. Ong, 25 Jan 1970 ; TMAG G6423 , 1♂ (27 mm), Tarn Shelf , Mt Field, pool on lodge side, 42°40.1'S 146°33.6' E, 1260 m asl, coll . T. Walker , Mar 1972 ; TMAG G6386 , 3 juv.♂♂ (15–18 mm), 2♀♀ (20–26 mm), 6 juv. ♀♀ (9–17 mm), Lake Dobson Road , 3 miles before Lake, 42°40.8' S 146°37.7'E, 1000 m asl,coll . R. Swain , 18 Dec 1969 ; TMAG G6410 , 4♂♂ (20–29 mm), 6 juv. ♂♂ (10–17 mm), 12♀♀ (19–28 mm), 10 juv. ♀♀ (11–14 mm), Tarn Shelf , near James Tarn, 42°40.3'S 146°33.8' E, small tarn with stream, sample 4, coll. I. Wilson & J. Ong, 25 Jan 1970 ; TMAG G6421 , 5♂♂ (22–26 mm), 4 juv. ♂♂ (13–17 mm), 3♀♀ (24–30 mm), 5 juv. ♀♀ (12–18 mm), Tarn Shelf , near Backhouse Tarn, 42°40.1'S 146°33.6'E, small tarn, sample 5, coll. I. Wilson & J. Ong, 25 Jan 1970 ; TMAG G6445 , 5♂♂ (20–21 mm), 6 juv. ♂♂ (13–19 mm), 2♀♀ (24–29 mm), 5 juv. ♀♀ (13–15 mm), Newdegate Pass , 42°39.6'S 146°33.0'E, sample 1, coll. I. Wilson & J. Ong, 25 Jan 1970 .

Junee-Florentine Karst: QVM 10 :12158, 1 juv. ♀ (20 mm), Rift Cave ( JF34-5 ), Junee, Florentine Valley, 42°44.3' S 146°35.6' E, 680 m asl, coll. S.M. Eberhard, 4 Jan 1985 ; TMAG G6481 , 1 juv.♂ (12 mm), Growling Swallet Cave , Florentine Valley, 42°41.4'S 146°30.0'E, JF 36-39, 570 m asl , CV10 , coll. S. Eberhard, 2 Jun 1985 ; TMAG G6473 , 1♀ (41 mm), Growling Swallet Cave ( JF36 ), Florentine Valley, 42°41.4'S 146°30.0'E, 550 m asl , CV41 , coll. S. Eberhard, 14 Apr 1985 .

No data: AM P 11897, 2♀♀ (31–39 mm), Tasmania,coll. B. Plomley via J.Waterhouse, pre 1949; QVM 10:13484, 1♂ (24 mm), 2♀♀ (21–24 mm),unidentified cave, coll.S. Eberhard,?1993; QVM 10:13976, 1♀ (31 mm), coll.E.O.G. Scott;USNM 1277686, 1♀ (31 mm), from G.M. Thomson, no data .

Description. Eyes with well-developed cornea, pigmented, wider than stalk (epigean forms) to narrower than stalk in some subterranean forms, longer than half length of stalk; stalk with subparallel margins (except in some “spinose” specimens). Rostrum narrow in adults, apex blunt.

Pleonites with sparsely setose pleural margins, rounded; posterior margin of tergites 5–6 setose. Pleuron 1 unarmed. Pleonites 2–6 usually with posterior margin of tergite 5 unarmed and pleura 2–5 unarmed or with 1 or 2 small spines on pleuron 5; pleonite 6 posterior margin unarmed or minutely spinose, posterolateral margin rounded, with or without minute denticle. Pleonites 2–6 of “spinose” specimens with pleura 3–5 (sometimes also pleuron 2) strongly spinose; posterior margin of tergites 5–6 strongly spinose; posterolateral margin produced to distinct spine. Pleonal sternites 3–5 with low, median processes between pleopod bases, distinctly bilobed and widest on sternite 3, bilobed on sternite 4, weakly bilobed and narrowest on sternite 5.

Telson slightly wider than long to longer than wide (usually slightly longer than wide), pentagonal (usually) to sub-linguiform, widest proximally; lateral margins sinuous in dorsal outline, distally subparallel to convergent; transition from lateral to posterior margin obtusely angular; posterior margin angular to rounded, blunt medially; posterior spine row with 19–50 slender, closely spaced spines, (usually) short, evenly graded to distinctly uneven in some “spinose” specimens.

Antennule inner flagellum about 0.2 × body length (21 articles in holotype); article 7 inner margin obtusely angled in adult males, with 1 long, slender clasping spine at proximal corner; outer flagellum 0.4–0.6 × body length (83 articles in holotype) in epigean specimens, 0.4–0.9 × body length in subterranean specimens.Antennal flagellum 0.4–0.6 × body length (62–65 articles in holotype), 0.5–0.9 in subterranean specimens; scaphocerite elongate, ovate, lateral spine slightly distal to midlength; apex reaching almost to midlength of distal peduncular article.

Right mandibular incisor process with proximal tooth distally undivided to trifurcate, usually bifid.

Pleopods 1–5 (rarely 1–4) with endopod in adults. Adult male pleopod 1 distally widened, scoop-like, lateral margins weakly expanded, not obscuring retinacular lobe in lateral view.

Uropodal protopod dorsally unarmed or with 1–3 spines; exopod with 2–4 (usually 3) movable spines on outer margin near position of partial diaeresis; exopod length about 2.5–3.5 times width, as wide as endopod, apex rounded, narrow to relatively broad.

Etymology. Named in honour of Alastair Richardson, for his many contributions to Tasmanian carcinology and limnology.

Measurements. Male (n = 756) 7–37, female (n = 1151) 6–55 mm, indet (n = 16) 5–7 mm.

Remarks. Anaspides richardsoni sp. nov. is characterized by the combination of a single antennular clasping spine in adult males ( Fig. 19D ), the telson posterior margin with a close-set spine row ( Fig. 19B ) and the male pleopod 1 endopod having the retinacular lobe visible in lateral view ( Fig. 20I ). Additionally, the pleopod 5 endopod is present in adults (except in some cave specimens from parts of the Mole Creek and Junee-Florentine systems; see below). Anaspides richardsoni is widespread across central Tasmania ( Fig. 36 ) in a wide arc stretching from the West Coast Range in the northwest, eastwards to Cradle Mountain and as far north as Gunns Plains and Mole Creek, across the Central Plateau, and south to Mt Field. It is primarily an epigean species occurring in springs, creeks and lakes, but also caves.

As presently understood, Anaspides richardsoni is the most morphologically variable species of the genus. Through the central-eastern (Mt Field east to Oatlands and north to Deloraine) and north-western part of its distributions (West Coast Range to Cradle Mountain and Mt Ossa to Gunns Plains), however, A. richardsoni is relatively uniform morphologically. The pleonites are almost always unarmed (or with few spinules on the posterior margin of pleonite 6), and in specimens from Mount Field to the Great Lake-Deloraine area, the telson is about as long as or little longer than wide, with the posterior margin distinctly angular, approximately right-angled ( Fig. 21 ). The telson in specimens from the far north (Gunns Plains, Loongana, Vale of Belvoir, Mole Creek) and western Central Plateau (Walls of Jerusalem area, Western Lakes) tend to be more elongate and often more acutely angular posteriorly ( Figs 21B, F ; 22 ). Specimens from the Mt Field area at the southern extremity of its range generally have more slender uropods than those from the northeast (most evident in large adults), and a usually bifid proximal tooth on the right mandibular incisor process ( Fig. 19H ) compared to the typically trifid condition found in other areas. Populations of A. richardsoni from the peripheries of the main distribution, however, show additional notable variations, particularly those from the far north, the western Central Plateau and the Mole Creek Caves.

Specimens from parts of Cradle Mountain and further north (Black Bluff, Vale of Belvoir, Gunns Plains) have a less angular telson than usual, sometimes being almost rounded. At a relatively small size (18 mm) some Cradle Mountain specimens already have a noticeably shortened telson (slightly wider than long; Fig. 21E ), broad uropods and broad scaphocerite compared to size-matched specimens from elsewhere.

Few specimens are known from the West Coast Range (Mt Murchison), and are the most westerly specimens examined ( Fig. 21A ). All have minimal pleonal spination (at most with few small spines on the posterior margin of pleonite 6). Most known males are juveniles with a single antennular clasping spine, although the largest male has (TMAG G6411) has one antennular clasping spine on the left and two on the right. The right antennule (peduncle and flagellum), however, is shortened overall and appears abnormal, possibly as a result of developmental irregularities. Smith’s (1909a) report of A. tasmaniae from Mt Read (near Mt Murchison, West Coast Range) is probably referable to A. richardsoni .

A number of specimens of A. richardsoni from the northwestern Central Plateau encompassing the Walls of Jerusalem and Clarence Lagoon east to the Ouse River have strongly spinose pereonites as in A. spinulae from Lake St Clair —the A. “ spinulae ” of O’Brien (1990) ( Fig. 22 ). In these specimens, pleonite 4–5 pleura (usually also pleonite 3) and the posterodorsal margins of pleonites 5 and 6 (5 less often so) are prominently spinose and the posterolateral margin of pleonite 6 is usually produced to a prominent spine. The proximal tooth on the right mandibular incisor process is usually trifid. In addition, the telson is often more elongate than usual, with a sometimes somewhat rounded margin on which the posterior spines vary from evenly graded to distinctly uneven in length. Telson spination differs slightly between sampled lakes and the Walls of Jerusalem. Specimens from Lakes Fox and Johnny ( Fig. 22N–S ) have relatively uneven telson spination compared to those from Clarence Lagoon ( Fig. 22A–I ), Lake Halkyard ( Fig. 22T,U ), Jacks Lagoon and most specimens from the Walls of Jerusalem ( Fig. 22J–M ), which have more evenly graded telson spines like those of typical epigean forms. It should be noted, however, that most specimens found throughout this area are non-spiny or less-spiny than “ spinulae ” (in which only pleura 4–5 and the dorsal margin of pleonite 6 may be spinose, e.g., Jacks Lagoon, QVM 10:13977). Both spiny and non-spiny forms were found together in tarns and runnels in the Walls of Jerusalem (TMAG G6396, TMAG G6379), the vicinity of Lake Fox ( Fig. 21F ), the Ouse River (TMAG G6359) and Clarence Lagoon (TMAG G6359) ( Fig. 22A–I ). The strong similarity in pleonal spination of the most spinose specimens of A. richardsoni to A. spinulae is remarkable, suggesting similar underlying developmental processes. O’Brien (1990), noting a correlation between spininess and lake habitats, suggested that A. spinulae might be a separate lacustrine species or lacustrine ecomorph, given that non-spiny specimens seemed to be from creeks, runnels and tarns. If so, however, it would remain to be explained why A. richardsoni from other lakes, such as Pine Tier Lagoon and Pine Lake are non-spiny, and why spiny forms can also be found together with non-spiny forms in tarns, small creeks, and pools. Both non-spiny and spiny forms are recorded from Clarence Lagoon ( Fig. 22A–I ). Additionally, the “ spinulae ” form of A. richardsoni occurs only on the Central Plateau to the east and northeast of the similarly spinose A. spinulae from Lake St Clair, but not in other lake populations of A. richardsoni (e.g., lakes at Mt Field and the West Coast Range such as Lakes Tyndall and Sandra), let alone lake populations of other species of Anaspides such as A. jarmani and A. swaini . The presence of predatory fish, such as trout and Galaxias might be posited to induce a spiny “defensive” phenotype in Anaspides as has been observed in Daphnia (Adler & Harvell, 1990) . Trout are certainly present in Lake St Clair ( A. spinulae sensu stricto), Clarence Lagoon, Lakes Fox, Halkyard and Johnny, but they are also present in more easterly water bodies such as Pine Tier Lagoon, Little Pine Lagoon and Pine Lake in which A. richardsoni shows no such defensive spination. Instead, the distribution of the “ spinulae ” form of A. richardsoni correlates more strongly with geography than ecology, and might reflect phylogeographic genotypic variation. In particular, the relationship between these spinose A. richardsoni and A. spinulae from Lake St Clair warrants further investigation, with A. spinulae differing most fundamentally in having two rather than one antennular clasping spines. Curiously, specimens to the immediate south and west of Lake St Clair, all referrable to A. swaini , do not have the spinose pleon of A. spinulae . Specimens to the immediate east of Lake St Clair from the eastern side of the Traveller Range are non- or minimally spinose as in typical specimens of A. richardsoni . In addition, the telsons of both spiny and non-spiny specimens from the Walls of Jerusalem to the northern tiers (as well as Mole Creek) tend to be more elongated than specimens from elsewhere. More detailed population level studies, currently underway in collaboration with S. Richter and colleagues, are required to further understand these morphological patterns.

Anaspides richardsoni has entered subterranean habitats in at least three parts of the periphery of its range ( Fig. 23 ): Mole Creek karsts and Great Western Cave (Gunns Plain) in the north, and the Junee-Florentine system in the south. The single known specimen from Great Western Cave (AM P99296; Fig. 23J ) is a juvenile male that agrees well with epigean juveniles (e.g., TMAG G6169); it also represents the northernmost record of Anaspides . Aside from loss of body pigmentation, the specimen exhibits no obvious troglobitic adaptations and has well-developed, pigmented eyes. At Mole Creek, three subterranean forms occur, differing between cave systems.

• Form 1 occurs in the Sassafras (Prohibition Cave) and Marakoopa systems (Marakoopa I and Marakoopa II caves) ( Figs 23A–C , 35F ). Apart from reduced body pigmentation, it closely resembles epigean specimens, having well-developed eyes, the outer antennular flagellum 0.4–0.6 (usually 0.5–0.6) body length, unarmed pleonites except for scattered denticles along the upper posterior margin of pleonite 6, and in having a finelygraded spine row on the posterior margin of the telson.

• Form 2 is found in caves in the Mole-Lobster system (Honeycomb Cave, Wet Cave, Herberts Pot and Kellys Pot) ( Figs 23D , 35E ). It resembles Form 1 in telson structure, but has reduced eyes, outer antennular flagella 0.5–0.8 (usually 0.6) body length, and more extensive pleonal spination, with denticles on the pleura of pleonites 4–5 as well as the dorsal posterior margin of pleonite 6. Form 2, reaches the greatest size of the three forms (> 50 mm), the largest being from Honeycomb Cave.

• Form 3 occurs in the Kubla Khan system (Kubla Khan Cave) and an unnamed cave in the Sassafras system ( Fig. 23E,F ). It resembles Form 1 in eye development and minimal pleonal spination, but differs from both Forms 1 and 2 in consistently having much more elongate antennular flagella (0.7–0.9, usually 0.8) and a more slender telson with longer, more robust spines, somewhat approaching that of A. richardsoni from the Walls of Jerusalem and localities close to Mole Creek such as Western Bluff, and Ironstone Mountain. Additionally, the pleopod 5 endopod is frequently absent in specimens from the Kubla Khan system. One series corresponding to Form 3 (USNM 1277683; Fig. 23F ) was collected from a deep, water filled sink hole in the Mayberry area (near Mole creek), believed to be hydrologically connected to the Kubla Khan system ( Iliffe, 1988); the specimens differ from the Kubla Khan specimens only in having more extensive body pigmentation typical of epigean forms, and a slightly wider cornea.

Although most common on the surface around Mt Field, A. richardsoni is recorded from the Junee-Florentine system (Rift Cave and Growling Swallet; Fig. 23G–I ) on the basis of two juveniles and an adult female. These specimens agree well with epigean A. richardsoni , including pigmentation (albeit seemingly somewhat reduced) and well-developed eyes, but differ in lacking the pleopod 5 endopod. The right eye of the Rift Cave specimen is deformed, seemingly having been damaged ( Fig. 23I ). Two other species of Anaspides occur in caves in the Mt Field area: A. eberhardi (Junee-Florentine karsts including Growling Swallet), which is closely related to A. richardsoni , and A. swaini (Junee- Florentine and Risby’s Basin karsts), which also sometimes occurs on the surface at Mt Field, but is the dominant epigean species to the south and west of the area.

Anaspides richardsoni features well-developed secondary sexual characteristics by 18–25 mm (usually 20–22 mm). The pleopod 5 endopod is the last of the endopods to develop, appearing in juveniles as small as 11 mm to as large as 22 mm, but usually by 15–18 mm. The presence of a single male antennular clasping spine is highly consistent, with, in rare cases, a second spine developing on one side, often associated with possible regeneration after damage. The male from Wet Cave, however, is highly aberrant in having two clasping spines on one side, three on the other. In both cases, the normal single clasping spine is present on the proximomedial corner of flagellar article 7, but the additional abnormally and asymmetrically developed spines arise on the anteromedial corner. The abnormal positions and asymmetrical development of the clasping spines in the Wet Cave male indicate that it is aberrant.

Anaspides richardsoni is the most taxonomically challenging species of the genus given the range of forms encountered, especially those from the peripheries of its range with distinctive morphologies. These may carry the signature of populations from areas untouched by the Pleistocene glaciations, which otherwise dominated the Central Plateau at the time, or may indicate ongoing differentiation ( Andrew, 2005). Further sampling is required to better understand these peripheral populations, especially since some localities, such as Gunns Plain, the West Coast Range and Sandbanks Tier are represented by juveniles or only few specimens. Further sampling is also required in the eastern Great Western Tiers in the vicinity of Deloraine, Great Lake, and Lake Sorell down to the Oatlands area to determine the current extent of occurrence. The specimens from the Oatlands area ( Fig. 21H ) are the easternmost records of the genus, although O’Brien (1990) speculated their provenance to be from surrounding hills slightly to the west. Likewise, specimens from Deloraine were speculated to originate from hills slightly to the south ( O’Brien, 1990). The known distribution A. richardsoni is discontinuous between the West Coast Range and Cradle Mountain area, and between the Western Lakes and Mt Field. Whether these “gaps” are real or owe to lack of sampling remains to be determined. Despite the wide morphological diversity, A. richardsoni is readily diagnosed by the combination of the single antennular clasping spine in adult males and the close-set spine row on the posterior margin of the telson.

Distribution. Wide ranging in central Tasmania, from the West Coast Range and Cradle Mountain in the west to the Central Plateau from Mt Ossa and localities east of Lake St Clair, to Mole Creek and the vicinity of Great Lake, south to Mt Field and as far east as the Oatlands area; 470–1455 m asl (epigean), 109–520 m asl (subterranean).

Figures

Figure 18.Anaspides richardsoni sp. nov., holotype male,33 mm, Mt Field,AMP72839. Habitus, right lateral view.Scale 4.0 mm.

Figure 19. Anaspides richardsoni sp. nov., holotype male, 33 mm, Mt Field, AM P72839. (A) cephalothorax, dorsal view; (B) pleonite 6, telson and left uropod; (C) pleonites 4–6 pleura, right lateral view; (D) right antennule; (E) left antenna; (F) labrum, anterior view; (G) right mandible; (H) right mandible incisor process; (I) paragnaths, anterior view; (J) right maxillule; (K) right maxilla. Scale: A–E = 1.0 mm; F–K = 0.7 mm.

Figure 20. Anaspides richardsoni sp. nov. (A–P) holotype male, 33 mm, Mt Field, AM P72839; (Q–R) paratype female, 32 mm, Mt Field, AM P72840;(A) right thoracopod 1 (maxilliped); (B–H) right thoracopods 2–8; (I–J) right pleopod 1 endopod, lateral and ventral views; (K–L) right pleopod 2 endopod, lateral and ventral views; (M) right pleopod 5, anterior view; (N–P) pleonites 3–5 median sternal processes; (Q–R) female gonopore, right lateral and ventral views. Scale A–M = 2.0 mm; N–R = 1.0 mm.

Figure 21. Anaspides richardsoni sp. nov., selected features. (A) male, 23 mm, S of Lake Tyndall, TMAG G6312; (B) female, 23 mm, Loongana, QVM 10:13975; (C) male, 22 mm, Deloraine, TMAG G135; (D) female, 26 mm, Little Pine Lagoon, QVM 10:13981; (E) juv. male, 18 mm, Cradle Mountain, SAM C6303; (F) female, 28 mm, W of Lake Fox, QVM 10:49165; (G) female, 27 mm, Mt Ossa, AM P82858; (H) female, 36 mm, Oatlands, TMAG G6360. Scale = 1.0 mm.

Figure 22. Anaspides richardsoni sp. nov., selected features. (A–D) female, 30 mm, Clarence Lagoon, TMAG G6359; (E) juv. female, 13 mm, Clarence Lagoon, TMAG G6359; (F–I) male, 23 mm, Clarence Lagoon, TMAG G6359; (J–M) male, 30 mm, Pool of Bethesda, TMAG G6434; (N–P) male, 23 mm, Lake Johnny, QVM 10:49162; (Q–S) Lake Fox, QVM 10:49164, female (29 mm), female (20 mm), female (30 mm); (T–U) Lake Halkyard, QVM 10:49163, male (18 mm), female (27 mm). Scale: A–D, F–U = 1.0 mm; E = 0.5 mm.

Figure 23. Anaspides richardsoni sp. nov., selected features. (A–C) Marakoopa Cave, SAM BS0464, female (31 mm), juv. female (21 mm), juv. female (9 mm); (D) male, 37 mm, Honeycomb Cave, AM P73038; (E) male, 25 mm, Kubla Khan Cave, USNM 1277679; (F) female, 28 mm, Mayberry area, sink hole, Mole Creek, USNM 1277683; (G–H) Growling Swallet, female 41 mm (TMAG G6473), juv. female 12 mm (TMAG G6481); (I) juv. female, 20 mm, Rift Cave, QVM 10:12158; (J) juv. female, 12 mm, Great Western Cave, Gunns Plain, AM P99296. Scale: A, B, D–G = 1.0 mm; C, H–J = 0.5 mm.

Figure 36. Distribution of Anaspides. Dubious Orford record of A. swaini indicated by “?”.

Figure35.(A) Anaspides tasmaniae (Thomson, 1893), female,Mt Wellington,specimen not preserved;(B) A.clarkeiAhyong, 2015, female, 24 mm, Eastern Passage streamway of Little Grunt Cave and Exit Cave, QVM 10:49168; (C) A. clarkeiAhyong, 2015, male, Lake Pluto, Wolf Hole, specimen not examined; (D) A. richardsoni sp. nov., female, Tarn Shelf, Mt Field, specimen not preserved; (E) A. richardsoni sp. nov., female, 32 mm, Kellys Pot, Mole Creek karst,TMAG G6463;(F) A. richardsoni sp.nov., female,35 mm, Marakoopa Cave, Mole Creek karst, specimen not preserved. Photo credits: A, Kristi Ellingsen, used under a Creative Commons Attribution-NonCommercial license (http://creativecommons.org/licenses/by-nc/2.0/); B, E, ©Stefan Eberhard; C, F, ©Arthur Clarke; D, ©Simon Grove, TMAG.