Tetralycosa oraria (L. Koch, 1876 )

Framenau, Volker W. & Hudson, Peter, 2017, Taxonomy, systematics and biology of the Australian halotolerant wolf spider genus Tetralycosa (Araneae: Lycosidae: Artoriinae), European Journal of Taxonomy 335, pp. 1-72 : 17-23

Tetralycosa oraria (L. Koch, 1876)

Figs 1 E–F; 5A–D; 6A–G; 9

Lycosa oraria Koch, 1876: 883–886 , pl. 76, figs 2, 2a, 3, 3a.

Lycosa candicans Koch, 1877: 888–890 , pl. 76, figs 5, 5a, 6, 6a–b. Synonymy established in Framenau et al. (2006).

Lycosa sibyllina Simon, 1909: 188–189 , fig. 7. Synonymy established in McKay (1979b).

Lycosa meracula Simon, 1909: 190–191 . Synonymy established in Framenau et al. (2006).

Lycosa oraria – Simon 1909: 188. — Rainbow 1911: 270. — Bonnet 1957: 2656.

Lycosa candicans – Rainbow 1911: 266. — Hickman 1950: 5. — Bonnet 1957: 2637.

Lycosa sibyllina – Rainbow 1911: 272.

Lycosa meracula – Rainbow 1911: 270. — McKay 1985: 80. — Platnick 1989: 372. — Moritz 1992: 320.

Crocodilosa oraria – Roewer 1955: 238.

Tetralycosa meracula – Roewer 1955: 296; 1960: 949. — Rack 1961: 38.

Hogna sibyllina – Roewer 1955: 253.

Trochosula candicans – Roewer 1955: 304.

Trochosomma oraria – Roewer 1960: 847.

Ocyale oraria – McKay 1973: 380.

Lycorma meracula – McKay 1973: 380.

Trochosa candicans – McKay 1973: 381; 1979b: 293–294, fig. 4e; 1985: 85. — Platnick 1989: 390.

Trochosa oraria – McKay 1979b: 279–282, fig. 1a–h; 1985: 86. — Platnick 1989: 391; 1993: 510.

Tetralycosa oraria – Framenau et al. 2006: 26 –27, fig. 58.

non Lycosa meracula – McKay 1979a: 264, fig. 9a–k: misidentification; not L. meracula but two species in the genus Costacosa Framenau & Leung, 2013 ( Framenau & Leung 2013).

Diagnosis

Tetralycosa oraria is very similar to T. orariola sp. nov. In particular the female genitalia are indistinguishable and separating both species is easiest by the colouration of the carapace. The lateral light bands of T. oraria are wide, although sometimes irregular, and run straight along the carapace margin. In contrast, the light bands in T. orariola sp. nov. are narrow and separated from the carapace margins by narrow dark bands. T. oraria is overall larger than T. orariola sp. nov., but sizes overlap.

The wide and light marginal bands of the carapace are similar to those of T. wundurra comb.nov., however, both species differ in their genital morphology. The embolus of the male pedipalp in T. wundurra comb. nov. is very broad narrowing abruptly at more than three quarters length into a slim tip, whereas it is of similar width along its whole length and twisted apically in T. oraria . The median septum of the epigyne of T. oraria has a distinct bridge with parallel borders, which is not present in T. wundurra comb. nov.

Type material

AUSTRALIA: syntypes of Lycosa oraria L. Koch, 1876 , unknown number of ♂♂ and ♀♀, King George Sound (35°02′ S, 117°56′ E, Western Australia), Bradley Collection (L. Koch, 1876). Considered lost ( Framenau 2005b) (not examined).

AUSTRALIA: holotype of Lycosa sibyllina Simon, 1909 , ♀, Albany (35°01′ S, 117°53′ E, Western Australia), 13/22 Aug. and 10 Oct. 1905, W. Michaelsen and R. Hartmeyer leg., ‘ Hamburger südwestaustralische Forschungsreise’, Station 165 ( ZMB 11102 ) (examined).

AUSTRALIA: lectotype (designated here) of Lycosa meracula Simon, 1909 , ♂, Albany (35°01′ S, 117°53′ E, Western Australia), 13/22 Aug. and 10 Oct. 1905, W. Michaelsen and R. Hartmeyer leg., ‘Hamburger südwest-australische Forschungsreise’, Station 165 ( MHNP 24364) (examined).

AUSTRALIA: syntypes (paralectotypes) of Lycosa meracula Simon, 1909 , 1 immature ♀, Denham (25°55′ S, 113°32′ E, Western Australia), 9–11 Jun.1905, W. Michaelsen and R. Hartmeyer leg., ‘Hamburger südwest-australische Forschungsreise’, Station 65 ( ZMB 11085) (examined); 1 juvenile, same data ( ZMH, Rack (1961) -catalogue 466) (examined); 1 juvenile, same data ( WAM 11/4303) (examined) (None of the three immature syntypes are conspecific with the male type from Albany ( Framenau et al. 2006)).

Other material examined

AUSTRALIA, New South Wales: 1 ♀, with spiderlings, Batemans Bay, 35°42′ S, 150°11′ E ( MV K8157); 1 ♂, Eden, 37°03′ S, 149°54′ E ( WAM T62653); 2 ♂♂, Nadgee Beach, 37°25′ S, 149°58′ E ( AM KS2266, KS50159); 1 ♂, Richmond Beach, 35°42′ S, 150°18′ E ( ANIC). – South Australia: 1 ♀, Baird Bay, 33°06′ S, 134°18′ E ( WAM T65045); 1 ♀, Bald Hill Beach, Port Wakefield, 34°15′ S, 138°10′ E ( SAM NN13934); 1 ♂, Charlton Gully, 34°33′ S, 135°47′ E ( SAM NN13861); 1 ♀, 1 ♀ with egg sac, 3 juvs, Deep Lake, Innes National Park, 35°16′ S, 136°51′ E ( SAM NN13850–1); 2 ♀♀, Edithburg, Yorke Peninsula, 35°05′ S, 137°44′ E ( WAM T56654); 1 ♂, Emu Bay, Kangaroo Island, 35°38′ S, 137°32′ E ( SAM NN13840); 2 ♂♂, 3 ♀♀, Flinders Island, Eyre Peninsula, 33°43′ S, 134°31′ E ( SAM NN13854–7); 1 ♂, 2 ♀♀, 3 juvs, Lake Greenly, 34°20′ S, 135°25′ E ( SAM NN13858–60); 3 ♂♂, Mt Rough, 12.1 km NNE of Mt Rough, Watervalley, 36°15′52″ S, 139°54′55″ E ( SAM NN13440–2); 1 ♂, 2 ♀♀, Policeman Point, Coorong, 36°04′ S, 139°35′ E ( SAM NN13842–4); 1 ♂, 3 juvs, Port Lincoln, 34°43′ S, 135°51′ E ( AM KS85107); 1 ♂, 1 ♀, Port McDonnell, 38°03′ S, 140°42′ E ( SAM NN13841); 1 ♀, Port Victoria, 34°30′ S, 137°29′ E ( SAM NN13852); 1 ♀, Reevesby Island, 34°31′ S, 136°16′ E ( MV K7813); 9 ♂♂, 4 ♀♀, Salt Lagoon, Kangaroo Island, 35°50′24″ S, 137°38′32″ E ( SAM NN13443– 5, NN13830–8); 1 ♀, Sinclairs Gap, salt lake nearby, 33°07′ S, 137°03′ E ( SAM NN13928); 2 ♂♂, 2 ♀♀, 1 ♀ with spiderlings, St Kilda Boat Barbour, Adelaide, 34°44′ S, 138°32′ E ( QM S61135 –6); 1 ♂, 1 ♀, 1 juv., Stokes Bay, Kangaroo Island, 35°37′ S, 137°12′ E ( SAM NN13839, QM S66329 ); 1 ♀, Thistle Island, 34°58′ S, 136°07′ E ( QM S61137 ); 1 ♀, Thistle Island, Eyre Peninsula, 35°00′ S, 136°10′ E ( SAM NN13853); 1 ♀, 1 juv., Troubridge Point, 35°09′30″ S, 137°40′00″ E ( SAM NN13849); 3 ♂♂, 1 ♀, Victor Harbour, W of jetty to Granite Island, 35°33′ S, 138°38′ E ( SAM NN13845–8); 1 ♀, Whyalla, sewerage ponds, 33°03′ S, 137°35′ E ( SAM NN13865). – Tasmania: 4 ♂♂, 1 ♀, 5 juvs, Adventure Bay, Bruny Island, 43°17′ S, 147°17′ E ( QM S66330 ); 3 ♀♀, 2 juvs, Brooks Creek, 41°18′ S, 144°44′ E ( QVMAG 13:44304–5); 2 ♀♀, Chain of Lagoons, East Tasmania, 41°40′ S, 148°17′ E ( QVMAG 13:11118); 1 ♂, 2 ♀♀, Clarke Island, 40°32′ S, 148°10′ E ( QVMAG 13:25911, 13:25923); 1 ♀, 6 juvs, Darlington Beach, Maria Island, 42°37′ S, 148°05′ E ( TMAG J3513); 2 ♂♂, Deal Island, 39°28′ S, 147°19′ E ( MV K7810); 3 ♂♂, 1 ♀, 3 juvs, Eaglehawk Neck, 43°01′ S, 147°55′ E ( QM S66331 –2); 1 ♀, Flinders Island, Whitemark, 40°07′S, 148°01′ E ( TMAG J650); 1 ♂, 1 ♀, 2 juvs, Hobbs Lagoon, 42°15′ S, 147°40′ E ( QM S66327 ); 1 ♂, 1 ♀, Lauderdale, 42°54′ S, 147°29′ E ( WAM T53669); 1 ♀, Pass River, King Island, 39°48′ S, 143°52′ E ( QVMAG 13:44306); 2 ♂♂, Preservation Island, Furneaux Group, 40°28′ S, 148°03′ E ( TMAG J1141); 2 ♀♀, Sandspit Point, Schouten Island, 42°17′ S, 148°15′ E ( TMAG J1660–1); 1 ♀, Strahan, 42°09′ S, 145°19′ E ( AM KS85104); 3 ♂♂, 3 ♀♀, Wynyard, Table Cape, 40°59′ S, 145°43′ E ( QM S66328 ). – Victoria: 1 ♂, no exact location ( MV K7809); 1 ♂, Cockatoo, 37°57′ S, 145°29′ E ( MV K8133); 1 ♂, Grey River, Otway Ranges, 38°41′ S, 143°50′ E ( AM KS85106); 1 ♀, Mornington, 38°13′ S, 145°02′ E ( MV K8154); 1 ♂, 2 ♀♀, Point Cook, 37°54′ S, 144°45′ E ( MV K9088); 1 ♀, Werribee, 37°59′ S, 144°33′ E ( MV K9104). – Western Australia: 1 ♀, Anvil Island, Recherche Archipelago, 33°30′ S, 123°58′ E ( WAM T62359); 2 ♂♂, 1 ♀, Australind, 33°16′ S, 115°43′E ( WAM 71/360–2); 1 ♀, Busselton, 33°38′ S, 115°20′ E ( AM KS85105); 1 ♂, City Beach, near rifle range, 31°56′ S, 115°45′ E ( WAM T53654); 1 ♂, E of Culham Inlet, 33°59′ S, 120°03′ E ( WAM 98/2150); 4 ♂♂, 1 ♀, Dingo Beach, Torbay, 35°05′ S, 117°38′ E ( QM W5649, W5651); 1 ♀, Dirk Hartog Island, 25°44′ S, 113°01′ E ( WAM T53817); 15 ♂♂, 9 ♀♀, Dongara, 29°14′ S, 114°55′ E ( WAM 71/ 1666–75, 71/1697–709, 71/1801); 1 ♀, 1 juv., Drummond Cove, via Geraldton, 28°39′ S, 114°36′ E ( QM W5643); 1 ♀, Esperance District, 33°51′ S, 121°53′ E ( MV K8145); 1 ♂, 4 ♀♀, 10 juvs, Fitzgerald River Inlet, 34°08′ S, 119°24′ E ( WAM 70/3800–14, 71/642); 1 ♀, Fremantle, North Mole, 32°03′ S, 115°44′ E ( WAM 70/217); 1 ♂, Fremantle, Obelisk Hill, 32°03′ S, 115°44′ E ( ZMB 10583); 1 ♀, Garden Island, 32°12′ S, 115°40′ E ( WAM 69/844); 3 ♂♂, 2 ♀♀, 1 juv., Geraldton, 28°46′ S, 114°37′ E ( WAM 71/1690–5); 1 ♂, Goode Beach, Albany, 35°04′ S, 117°56′ E ( WAM T53653); 3 ♂♂, 3 ♀♀, Guilderton, 31°21′04″ S, 115°29′52″ E ( WAM T53393–7); 1 ♀ with spiderlings, Guilderton, Moore River, 31°20′ S, 115°30′ E ( WAM 71/1993); 1 ♀, 1 juv., Gun Island, South Group, Houtman Abrolhos, 28°53′ S, 113°51′ E ( WAM 71/1667–7); 3 ♀♀, Lancelin Island, 31°0′ S, 115°19′ E ( WAM 71/771–3); 1 ♂, Leeman Swamp, 29°56′27″ S, 114°59′50″ E ( WAM T93541); 1 ♀, Mettams Pool, 31°52′ S, 115°45′ E ( WAM T56531); 1 ♂, 1 ♀, Middle Island, Recherche Archipelago, 34°06′ S, 123°11′ E ( WAM T69946); 4 ♂♂, 7 ♀♀, 1 juv., North Island, Houtman Abrolhos, 28°43′ S, 113°47′ E ( WAM 71/ 1680–85, 71/1802–5, 71/1991–2, 71/1994); 1 ♀, Point Malcolm, 33°47′ S, 123°45′ E ( WAM T51432); 2 ♂♂, Post Office Island, South Group, Houtman Abrolhos, 28°52′ S, 113°58′ E ( WAM 71/1678–9); 2 ♂♂, 1 ♀, Quarantine Bay, 37°03′ S, 149°52′ E ( WAM T64979); 1 ♂, 1 ♀, Rossiter Bay, Cape Le Grand National Park, 33°58′21″ S, 122°16′13″ E ( WAM T93994); 1 ♀ with egg sac, Rottnest Island, 32°00′ S, 115°36′ E ( WAM T47247); 1 ♂, Safety Bay Foreshore Reserve, 32°18′ S, 115°43′ E ( WAM T65111); 1 ♂, Shelly Beach, 35°02′ S, 116°44′ E ( WAM T53495); 3 ♂♂, South Cottesloe Beach, 31°59′ S, 115°45′ E ( WAM T65606); 2 ♀♀, Two Peoples Bay, 34°57′ S, 118°11′ E ( WAM T51431, T53692); 1 ♀, Windy Harbour, back beach, 34°50′ S, 116°01′ E ( WAM 72/251).

Description

MEASUREMENTS. ♂ WAM T53697 (♀ WAM T53393): TL 8.25 (12.30), CL 4.50 (5.10), CW 3.45 (4.13). Eyes: AME 0.21 (0.21), ALE 0.17 (0.18), PME 0.38 (0.41), PLE 0.33 (0.37). Row of eyes: AE 0.93 (1.05), PME 0.93 (1.05), PLE 1.23 (1.54). Sternum length/width: 1.95/1.65 (2.25/1.88). Labium length/ width: 0.58/0.60 (0.72/0.79). AL 2.85 (6.75), AW 4.2 (5.1). Legs: Lengths of segments (femur + patella/ tibia + metatarsus + tarsus = total length): Pedipalp 1.95+1.65+ – +1.25 = 4.85, I 3.60+4.50+3.30+1.80 = 13.20, II 3.75+4.65+3.75+2.25 = 14.4, III 3.38+3.83+3.60+1.65 = 12.46, IV 4.20+4.95+4.65+1.95 = 15.75 (Pedipalp 2.25+2.10+ – +1.35= 5.70, I 3.60+4.35+2.70+1.65 = 12.30, II 3.60+4.20+2.85+1.65 = 12.30, III 3.45+3.90+3.30+1.58 = 12.23, IV 4.50+5.10+4.35+1.95 = 19.90).

VARIATION. ♂ (♀) (range, mean ± SD): TL 6.75–9.45, 8.35 ± 0.82; CL 3.75–4.95, 4.55 ± 0.39; CW 2.85– 4.05, 3.55 ± 0.46; n = 9 (TL 9.75–14.25, 11.26 ± 1.35; CL 4.05–6.45, 5.04 ± 0.77; CW 3.15– 5.03, 3.98 ± 0.64; n = 13). Tetralycosa oraria is very variable in colouration. Some specimens are extremely light without a distinct colour pattern; others are dark olive-grey with light bands ( Fig. 1 E–F). It appears that the particular microhabitat (for example dark seaweed vs plain beach) may explain the colour patterns of individual specimens (see also McKay 1979b). There also appears to be some intraspecific morphological variation in leg length and eye position, as McKay (1979b) reported a different leg formula (IV>I>II>III) and relative length of the anterior row of eyes and posterior median eyes.

Male (based on WAM T53397)

CARAPACE ( Fig. 5A ). Dorsal line straight in lateral view ( Fig. 6A ); brown with indistinct and irregular light colouration medially and broad, irregular light marginal bands; indistinct darker radial pattern; covered with mainly brown setae, but white setae in marginal bands; very few brown bristles in eye region mainly between PME; four long bristles below AE; one long bristle between AME.

EYES ( Fig. 6B ). Row of AE as wide as row of PME; row of AE slightly procurved ( Fig. 6B ).

CHELICERAE. Brown, darker apically; covered with long white setae and few brown bristles medially; three promarginal teeth with the median largest; three retromarginal teeth of similar size.

STERNUM ( Fig. 5B ). Yellow-brown, with dark pigmentation towards the margins; covered with few white setae and longer brown macrosetae that are longer towards margins.

LABIUM. Brown, basally dark brown; front end truncate and white.

PEDIPALPS ( Fig. 6 C–E). Tegular apophysis broad with a basally pointing tip ( Fig. 6C ); terminal apophysis forms sheath for the resting embolus; embolus long and thin with curved tip ( Fig. 6E ).

ABDOMEN. Dorsally dark olive-brown with light lanceolate heart mark in anterior half; few irregular light patches formed by fields of lighter setae; otherwise covered in brown setae ( Fig. 5A ). Ventrally yellowbrown with some mottled olive-grey discolouration and with white and fewer brown setae. Spinnerets yellow-brown ( Fig. 5B ).

LEGS. Leg formula IV>II>I>III. Femora yellow, distal segments darker; weak scopula on tarsus I. Spination of leg I: femur: three dorsal, one apicoprolateral; patella: one prolateral; tibia: three ventral pairs, two prolateral, two retrolateral; metatarsus: three ventral pairs, two prolateral, two retrolateral, one apicoventral, one apicoprolateral.

Female (based on WAM T53393)

CARAPACE ( Fig. 5C ). Brown, indistinct narrow light median band and wide marginal bands; covered with white setae and few black setae anteriorly of fovea; brown bristles in and around eye region; four long bristles below AME, one long bristle between AME.

EYES. Row of AE as wide as row of PME; row of AE slightly procurved.

CHELICERAE. Colour, setae and dentition as male.

STERNUM ( Fig. 5D ). Yellow-brown and somewhat darker marginally; setae as male.

LABIUM. As male.

ABDOMEN. Dorsally yellow-brown with indistinct light lanceolate heart mark; setae as male but with grey setae on dark patches ( Fig. 5C ). Venter and spinnerets yellow-brown with little darker pigmentation ( Fig. 5D ).

EPIGYNE. Ventral view ( Fig. 6F ): ovoid median septum with curved posterior border. Dorsal view ( Fig. 6G ): small spermathecal heads, convoluted spermathecal stalks attach posteriorly.

LEGS. Leg formula IV>I=II>III. Colour pattern as male. Spination of leg I: femur: two dorsal, one apicoprolateral; tibia: three ventral pairs, two prolateral (only one on left leg); metatarsus: three ventral pairs, two prolateral; one retrolateral (only on left leg), one apicoventral.

Remarks

An adult male syntype of Lycosa meracula was only recently discovered in the MHNP resulting in reevaluation of the juvenile type material of this species that was examined by McKay (1979a) ( Framenau et al. 2006). The species illustrated as L. meracula by McKay ( McKay 1979a) is not conspecific with the male syntype of L. meracula from Albany. The material included two closely related coastal species in the genus Costacosa ( Framenau & Leung 2013) . To fix the taxonomic concept of T. meracula , the male from Albany (MHNP 24364) is here designated as lectotype.

Life history and habitat preferences

Tetralycosa oraria is an exclusively coastal species inhabiting sandy beaches and the base of sand dunes on the foreshore. Adult spiders of T. oraria can be found all year round and do not seem to have a clear seasonal pattern for reproduction, as females with egg sacs and spiderlings have been found from December to February, but also in July and October.

Distribution

Coastal mainland Australia south of approximately 25°S latitude, and Tasmania, although a few records are from further inland, if saline wetlands are found near the beach ( Fig. 9 ).

Figures

Fig. 1. Typical habitats and live images of Tetralycosa Roewer, 1960. A. Salt lake (southern Lake Lefroy, Western Australia), where T. alteripa (McKay, 1976) and T. baudinettei sp. nov. can be found. B. T. alteripa (McKay, 1976), burrow on Lake Lefroy. C. T. alteripa (McKay, 1976), ♀ from Lake Lefroy. D. Sea shore (Cottesloe Beach, Western Australia), where T. oraria (L. Koch, 1876) can be found. E. T. oraria (L. Koch, 1876), ♂ from Cottesloe Beach (WAM T65606). F. T. oraria (L. Koch, 1876), ♂ from Safety Beach (Western Australia) (WAM T65111).

Fig. 5. Tetralycosa Roewer, 1960, microscopic photographs showing dorsal and ventral views. A–B. T. oraria (L. Koch, 1876), ♂ (WAM T93541). C–D. T. oraria (L. Koch, 1876), ♀ (WAM T93944). E–F. T. arabanae (Framenau, Gotch & Austin, 2006), ♂ (WAM T47298). G–H. T. arabanae (Framenau, Gotch & Austin, 2006), ♀ (WAM T47298). I–J. T. caudex sp. nov., ♀ (WAM T55576). Scale bars: A–J = 2.00 mm.

Fig. 6. Tetralycosa oraria (L. Koch, 1876), ♂ (WAM 71/360) and female (WAM 71/361). A–B. Male carapace, lateral and frontal view. C–D. Left male pedipalp, ventral and retrolateral view. E. Left male pedipalp, palea section of bulbus, ventral view. F–G. Female epigyne, dorsal and ventral view. Scale bar: A = 4.42 mm; B = 1.88 mm; C–D = 1.28 mm; E = 0.56 mm; F–G = 0.89 mm.

Fig. 9. Tetralycosa oraria (L. Koch, 1876), T. arabanae (Framenau, Gotch &Austin, 1976) and T. caudex sp. nov., distribution records in Australia.