Trochosa hispanica Simon, 1870

Marusik, Yuri M. & Nadolny, Anton A., 2020, On the identity of Trochosa hispanica (Araneae, Lycosidae), with notes on the synonymy of West Palaearctic “ Trochosa ” species, Zootaxa 4859 (1), pp. 56-80 : 58-65

Trochosa hispanica Simon, 1870

Figs 1–2 , 3C , 4–9 , 10 A–B, 11–12, 15

Trochosa hispanica Simon, 1870: 357 (♂ ♀).

Trochosa ruricola rustica Thorell, 1875: 169 (♂ ♀, in part); Thaler et al. 2000: 1079 (S).

Trochosa manicata Thorell, 1875: 170 (♂); Bonnet 1959: 4705 (S).

Lycosa caliginosa Simon, 1876: 287 (♂ ♀); Bonnet 1959: 4705 (S).

Lycosa manicata: Simon 1937: 1112, 1137 , fig. 1744 (♂ ♀, figured ♀ only).

Arctosa nava Roewer, 1955a: 761 , figs 8–10 (♂ ♀). Syn. n.

Geolycosa flavichelis Roewer, 1955a: 765 , fig. 13 (♀). Syn. n.

Loculla austrocaspia Roewer, 1955a: 769 , fig. 20 (♀). Syn. n.

Trochosa hispanica: Hänggi 1989: 168 , figs 1a–b (♂ ♀); Thaler et al. 2000: 1079, figs 37–39, 41 (S); Hepner & Milasowszky 2006: 7, fig. 5.I1a (♀); Moscaliuc 2012: 12, figs 4–5 (♀); Lecigne 2016a: 27, figs 13D–G (♂ ♀); Otto & Japoshvili 2018: 382, figs 46–47 (♂).

Trochosa nava: Mozaffarian & Marusik, 2001: 70 ( Platnick [2002] and subsequent catalogs did not accept the transfer because the types were not examined).

Trochosa austrocaspia: Mozaffarian & Marusik 2001: 70 ( Platnick [2002] and subsequent catalogs did not accept the transfer because the types were not examined).

Trochosa flavichelis: Mozaffarian & Marusik 2001: 70 ( Platnick [2002] and subsequent catalogs did not accept the transfer because the types were not examined).

Types: Trochosa rustica Thorell, 1875 (two males from Ukraine in different tubes, one female from Italy): lectotype ♂ designated here (NHRS) “240/ 1490g, Ekaterinoslaw” =Dnipro City, Ukraine ( Fig. 10 A ), examined; paralecto- type ♀ (NHRS) “239/1490f, Genazzano”, Italy ( Fig. 10 B ), examined. Paralectotype ♂ (NHRS) “240/1490h, Ekat- erinoslaw” belongs to T. terricola , examined.

Arctosa nava Roewer, 1955 ( Fig. 12 A–C): lectotype ♂ and paralectotypes 1♂ 1♀ designated here ( SMF), “Arachn. Coll. Rwr. -Lfd. No.11439, No.701, Typ., Iran: Sabzawaran”, J. Hemsen leg., examined (one palp is missing, maybe in slide mount) ; paralectotypes 1♂ 1♀ ( SMF), “Arachn. Coll. Rwr. -Lfd. No.11440, No.702, Paratyp., Iran: Chorramabad”, F. Starmühlner leg., examined .

Geolycosa flavichelis Roewer, 1955 ( Fig. 12 F–H): lectotype ♀ and paralectotypes 2♀ designated here ( SMF), “Arachn. Coll. Rwr. -Lfd. No.11442, No.704, Typus, Lahidschan”, examined. In the text, 3 localities and 3 different collectors are mentioned ( Roewer 1955a) : 1♀, Lahidschan, Typus, F. Starmuhlner leg.; 1♀, Babol, Paratypoid, J. Hemsen leg.; 1♀, bei Tschalus (Kaspi-Kuste), Paratypoid, H. Loffler leg.

Loculla austrocaspia Roewer, 1955 ( Fig. 12 D–E): ♀ holotype (SMF) “Arachn. Coll. Rwr. -Lfd. No.11366, No.694, Holotyp., Kaspi-Küste,” H. Löffler leg., examined.

Material examined: PORTUGAL: 4♂ 1♀ ( CRB), Algarve , 02.2006, 05.2007 ( R. Bosmans) . SPAIN: 1♂ 1♀ ( CRB), Cadize , Tarifa, 04.1991 ( R. Bosmans) ; 1♂ 1♀ ( CRB), Caceres, Monfrague , 3.04.1994 ( R. Bosmans) . FRANCE: 7♂ 5♀ ( TNU), Villeneuve-les-Maguelone , salt marshes, 25.05– 10.06.2011 (J. Pages) . GREECE: Corfu : 1♀ ( MMUE G7572.9628 ), Agios , 2– 9.04.1983 (J. & F. Murphy) ; 1♂ ( MMUE G7572.9662 ), Peroulades , 3.04.1983 (J. & F. Murphy); Halkidiki : 3♂ 2♀ ( MMUE G7572.3787 , G7572.3841 , G7572.4884 ), reedbed, 13– 19.04.1978 (J. & F. Murphy); Crete : 1♂ ( MMUE G7572.16293 ), Mallia , marsh, 9.04.1972 (J. & F. Murphy) ; 1♂ ( MMUE G7572.6286 ), 12.04.1979 (J. & F. Murphy) ; 2♀ ( MMUE G7572.8480 ), Georgioupolis , sand dunes, 15.04.1981 (J. & F. Murphy) ; 2♂ 1♀ ( MMUE G7572.8441 ), Akrotiri, Kalathas pond, reedbed, 19.04.1981 (J. & F. Murphy) ; 3♂ 1♀ ( MMUE G7557.260 ), edge of Kournas Lake, near Georgioupolis , on and under stones in dense shrubs, 28.05.1993 (J. & F. Murphy) ; 1♂ ( MMUE G7557.259 ), Georgioupoli , 29.05.1993 (J. & F. Murphy) ; 1♀ ( MMUE G7557.261 ), near Lefkogia , between stones by stream edge, 4– 5.06.1993 (J. & F. Murphy) ; 1♂ ( MMUE G7557.269 ), 7.06.1993 (J. & F. Murphy) . BULGARIA: 10♂ 2♀ ( TNU), Zemen Gorge, Rhuzhdavitsa Vill. , 6.07.1984 (G. Blagoev). TUR- KEY : 1♂ ( ZMMU), Bursa Prov., Nilufer Distr. , 44°07.466′N 28°42.105′E, 570 m, 2.06.2009 (unknown collector). AZERBAIJAN: Lenkoran : 7♂ 4♀ ( ZMUT), Hyrcan Reserve, env. of Mamusta Vill. , 38°38’N 48°47’E, lowland forest, 27.05.2003 (Y.M. Marusik) ; 1♀ ( ZMMU), 25th km of Lenkoran-Lerik Hwy , 38°44.385’N 48°37.024’E, 254 m, tea plantation, 26.05.2003 (Y.M. Marusik) ; 1♂ 2♀ ( ZMMU), Hyrcan Reserve, env. of Apo Vill. , 38°38’N 48°47’E, 28.05.2003 (Y.M. Marusik) ; 2♂ ( ZMMU), Hyrcan Reserve , 38°38.5’N 48°47.5’E, 23.05.2003 (Y.M. Marusik) ; 7♂ 1♀ ( ZMMU), environs of Aurora Vill. , 38°41’N 48°17’E, 36 m, 21– 29.05.2003 (Y.M. Marusik). Lerik : 1♀ ( ZMMU), ca 3 km E of Divagatch Vill. , 38°41.5’N 48°23’E, 1400m, meadow with stones along riv- er, 26.05.2003 (Y.M. Marusik) . IRAN: Mazandaran Prov.: 2♂ 5♀ ( ZMMU), Khorran-abad Vil. env., 50°48’E 36°43’N, 8– 10.06.2000 (Y.M. Marusik) ; 2♂ 3♀ ( MMUE), N of Javaher-Deh Vil. , 50.467°E 36.872°N, 9.06.2000 (Y.M. Marusik) ; 1♀ ( MMUE), Javaher-Deh Vil. , 50.467°E 36.867°N, 9.06.2000 (Y.M. Marusik) ; 3♀ ( ZMMU), Nashtarood-Khoshkadaran , 51.033°E 36.750°N, 9– 10.06.2000 (Y.M. Marusik); Tehran Prov. : 4♂ 1♀ ( ZMMU), NW of Tehran, Sardor area , 10km N of Karaj, 51°05’E 35°50’N, 13.06.2000 (Y.M. Marusik) ; 1♀ ( MMUE), Tehran , Plant-Protection-Organization Pk , 51.414°E 35.673°N, 7– 22.06.2000 (Y.M. Marusik). CRIMEA : 1♂ 3♀ ( MMUE), Sevastopol, Balaklava, Asketi Mt. , 2019 (A.A. Nadolny) ; 132♂ 23♀ ( MMUE & TNU), Yalta, Massandra park, 2000–2001 (M.M. Kovblyuk) ; 4♂ 2♀ ( TNU), Yalta Distr., Martyan Cape Reserve , 2000–2001 (M.M. Kovblyuk) ; 2♀ ( TNU) Alushta Distr., Alaka River bank, 2000 (G.A. Prokopov) ; 1♂ ( TNU), Sudak Distr., Mezhdurech’e Vill. , 2010 (M.K. Yusufova) ; 118♂ 68♀ ( MMUE & TNU), Feodosiya Distr., Karadag Nature Reserve , 2003–2011 (M.M. Kovblyuk et al.) ; 2♂ ( TNU), Lenino Distr., Kerch Peninsula, Kazantip Nature Reserve , 2015–2016 (M.M. Kovbly- uk et al.) .

Diagnosis. Males of T. hispanica can be easily distinguished from all congeners by: 1) the combination of a coiled embolus and a fang with a basal tooth ( Figs 2I , 4 A–F, these characters never occur together in other species); 2) a unique coloration of leg I (see Fig. 3C ), with a blackish patella, and tibia and proximal half of metatarsus black (vs. patella not black); 3) the dorsal stripe or spots of white setae on tibia I ( Fig. 2 B–C, K) (present in some specimens, vs. white pubescence absent). Females of T. hispanica differ from congeners (except for T. cachetiensis and some aberrant specimens of T. ruricola ) by having the epigynal hoods close together, often touching each other ( Fig. 6 ). Females of T. cachetiensis , T. hispanica , and some T. ruricola are indistinguishable due to the considerable variation in the epigyne.

Description. Male ( Figs 1 A–B, 2, 3C, 4–5, 8–9, 10A, 11A, 12A–B). Length of carapace 3.5–4.6. General pattern as in other species ( Fig. 1 A–B). Fang with basal tooth ( Fig. 2I ). Leg I with characteristic pattern ( Fig. 2 E–H): patella, tibia and proximal part of metatarsus black; femur and distal part of metatarsus and tarsus light yellow. Tibia I often with white dorsal setae ( Figs 2K ; 8C, F, I ) forming 3 types of dorsal pattern: 1) longitudinal white stripe ( Figs 2 B–C, K; 8A), 2) series of white spots ( Thaler et al., 2000: fig. 37) or 3) uniformly dark with several distal white, nearly indistinct setae ( Figs 1A ; 2A, D, J ; 8B, E, H ). Microstructure of white setae similar to that of black setae, but the white setae are slightly bent and have longer barbs ( Figs 9E, F ). Specimens studied with SEM microscope have different density of fine setae on the tibia I: specimen with a white pubescence has 40–45 setae per 0.04 mm 2, specimens without a white pubescence—25–35 setae per 0.04 mm 2. Tibia I with a row of elongate, glabrous cuticular spots (Cs) ( Fig. 9 A–C, G), undocumented in other lycosids. Palp ( Figs 4–5 , 10A , 12A ): cymbium with one claw; tip of embolus coiled, forming one loop.

Female ( Figs 1 C–D, 6–7, 10B, 11B, 12C–H). Length of carapace 4.1–6.2. Epigynal hoods (Eh) located near each other and touching each other in most specimens. Shape of hoods variable ( Figs 6–7 ). In almost all specimens, the hoods are filled with a substance that resembles a mating plug ( Fig. 7 D–G). It seems that the hoods together have a furrow along the lateral edge of the fovea (Ff) ( Figs 6–7 ) that serves as guide for one of the bulbal apophyses. Receptacles globular, 1.5 times wider than stalks, receptacle supplied with gland. Position of receptacles and angle of inclination toward the epigynal plate variable, and the angle of inclination can differ between the right and left receptacle of the same specimen.

Note. Mating plugs are reported for some other lycosids (see Uhl et al. 2010), but the specific type of plug was not mentioned. Mating plugs have been documented in detail for Wadicosa oncka ( Lawrence, 1927) ( Kronestedt 1987) and Alopecosa psammophila Buchar, 2001 ( Szinetár et al. 2005). In both cases, the plug occupies the entire fovea (atrium). However, in T. hispanica it blocks the pathway for one of the bulbal apophyses (furrow, Figs 6–7 ) and looks like a dark stripe ( Fig. 7D, G ).

Variation. Examination of T. hispanica collected from localities found throughout the entire range reveals variation in several characters, as listed below.

1) We have found out that one of the most distinctive characters allowing the identification of this species, a white pubescence on males’ tibia I, can be present or absent in specimens collected in different localities ( Fig. 2 A–B) and collected together from the same locality ( Figs 2 C–D, J–K; 8A–D). Distant populations have different proportions of males with this character—ranging from 0 to 100 % ( Fig. 15A ). In specimens from the Mediterranean, the white pubescence was only found in a few males: in single males from Corfu and France. The white pubescence is found in a few males from Iran, including the syntypes of Arctosa nava ( Fig. 12B ), and it

is found in all males examined from Azerbaijan. Occasionally, there are spots of white setae on the tibia of males from the Caucasus and Greece ( Thaler et al. 2000: fig. 37; present data). All specimens studied from the Crimea, Crete, Halkidiki, Turkey, and Bulgaria lack the white pubescence. Close examination of numerous males ( Fig. 15A ) from the Crimea and males from the Mediterranean and Iran that lack the white pubescence revealed that there are about 10 white setae on the distalmost part of the tibia, although they are difficult to see ( Fig. 8 ).

2) The male palp is very similar within and between distant populations ( Figs 4–5 ); however, specimens from Iran have a smaller bulb and shorter cymbium tip.

3) The epigyne displays high variation, even among specimens collected from the same locality simultaneously: the edges of the hoods can be rounded ( Fig. 6 A–G), straight ( Fig. 6H ) or bent toward the septum ( Fig. 6I ). The shape of the hoods also varies ( Fig. 7 ).

4) The size of males and females within and between populations is highly variable, but there is overlap. In general, specimens from southern and southwestern populations are smaller than those from Crimea. The exception is specimens from Crete, which are larger than Crimean specimens ( Fig. 11 ).

Biology. In the Crimea, T. hispanica occurs from sea level up to 300 m and inhabits shrublands on the Southern Coast and Kerch Peninsula ( Fig. 15B ). Adult males and females occur from March to October with peak activity in May. Females with cocoons ( Fig. 1D ) were observed from July to September. In the Crimea, T. hispanica is abun dant on the Southern Coast of Crimea (with a dynamic density of up to 5 adults in 100 pitfall-traps per day), but T. robusta ( Simon, 1876) and T. ruricola are rare in the south (up to 0.1 adults in 100 pitfall-traps per day) and are common in the mountains, semi-desert and true steppes.

Distribution. The known range of T. hispanica stretches from the Iberian Peninsula to Tajikistan (World Spi- der Catalog 2020), north to the Dnipropetrovsk Area in Ukraine (present data), and south to Algeria and Morocco ( Nentwig et al. 2020). The actual distribution proven by the material is smaller, from the Iberian Peninsula to Mazandaran Province, Iran ( Fig. 15 ). A record of a single female from northern Tajikistan ( Charitonov 1951, sub T. rustica Thorell, 1875 ) most likely refers to another species because identification of females cannot be made with certainty, especially in a time when this species was not illustrated. This specimen was meant to be deposited in Perm University collections but could not be located (Esyunin, pers. comm.). In addition, all Trochosa specimens we examined from Central Asia belong to T. ruricola , a species known from all over Central Asia ( Mikhailov 2013). It is likely that southeasternmost record of T. hispanica ( Fig. 15C ) in Iran is based on mislabeling. There are no recent records of this species in the continental part of Ukraine. The single record refers to specimen collected in 19 th century ( Fig. 15B ).

In order to allow for easier identification of Trochosa species occurring in the West Palaearctic, a key for the males is provided.

Figures

FIGURE 1. Males (A–B) and females (C–D) of Trochosa hispanica from the Crimea.A, C–D living and B preserved in ethanol. Scale bar 2 mm

FIGURE 2. Males of Trochosa hispanica from France (A, E), Azerbaijan (B), Iran (C–D, F–G, J–K), and Crimea (H–I). A–B prosoma, dorso-lateral; C–D general appearance, dorsal; E–H leg I, retrolateral; I chelicera, posterior view (arrow indicates position of tooth); J–K tibia I, dorsal-retrolateral. Scale bars A–H 1 mm; I–K 0.5 mm

FIGURE 3. Leg I of male Trochosa robusta (A), T. ruricola (B), T. hispanica (C), T. spinipalpis (D), T. terricola (E), and T. urbana (F). A–C from Crimea; D from Finland; E from Adygea Republic; F Seychelles. Scale bars 1 mm

FIGURE 4. Male palps of Trochosa hispanica from France (A, E, H–I), Crete (B), Iran (C), and Crimea (D, F–G, J). A–F ventral; G–H anterior; I–J retrolateral. Abbreviations: Em embolus, Se synembolus, Ta tegular apophysis. Scale bars A–H 0.2 mm; I–J 0.5 mm

FIGURE 5. Embolic divisions of Trochosa hispanica from France (A–B) and Crimea (C–D). A, C ventral; B, D posterior. Abbreviations: Em embolus, Se synembolus. Scale bar 0.1 mm

FIGURE 6. Ventral view of epigynes of Trochosa hispanica from France (A–B), Crete (C), Iran (D–E), Bulgaria (F), and Crimea (G–I). Abbreviations: Eh epigynal hoods, Ff furrow, Mp remnants of mating plug. Scale bars 0.2 mm

FIGURE 7. Dorsal view of endogynes of Trochosa hispanica from France (A–B), Crete (C), Iran (D–E), Bulgaria (F), and Crimea (G–I). Abbreviations: Eh epigynal hood, Ff furrow, Mp remnants of mating plug. Scale bars 0.2 mm

FIGURE 8. Dorsal view of male tibiae I of Trochosa hispanica from Iran (A–D), France (E–G), and Crimea (H–J). A tibia with a white stripe; B, E, H tibiae without a white stripe; C white setae; D, G, J black setae; F, I white setae on the distalmost part of the tibia. Scale bars A, B, E, H 0.5 mm; C–D, F–G, I–J 0.01 mm

FIGURE 9. Details of male tibiae I, dorsal view, of Trochosa hispanica from Crimea (A, F), France (B), and Iran (C–E, G).A–C tibiae without a white stripe; D tibia with a white stripe; E white setae; F black setae; G glabrous spots. Abbreviations: Bs and Ws black and white setae; Cs glabrous cuticular spots. To show the cuticle surface, part of setae were removed in A–D

FIGURE 10. Trochosa hispanica (types of Trochosa rustica, A–B) and T. adjacens (C). A palp of lectotype, ventral; B epigyne of paralectotype, ventral; C epigyne of holotype, ventral. Scale bar C 0.2 mm. A–B photo: Torbjörn Kronestedt

FIGURE 11. Carapace length/femur I length ratio in males (A) and females (B) of Trochosa hispanica from different localities

FIGURE 12. Trochosa hispanica: types of Arctosa nava (A–C), Loculla austrocaspia (D–E) and Geolycosa flavichellis (F–H). A male palp, ventral; B male prosoma, lateral showing white setae on tibia I; C, E–H epigyne, ventral; D female, dorsal. Letters marked with ’ refer to original figures by Roewer (1955a)

FIGURE 15. A—map showing the proportion of males of Trochosa hispanica with (white sectors) or without (black sectors) white pubescence on tibiae I among studied material (number in circles indicates how many males were examined). The broken line refers to the general distribution. B—map showing distribution of T. hispanica in Ukraine, black dots refer to recently collected material, and white dot refers to specimens collected in the 19th century (type locality of T. rustica). C—map showing distribution of T. hispanica in Iran, black dots refer to recently collected material, and white dots refer to Roewer’s types, synonyms of T. hispanica