The stone huntsman spider genus Eusparassus (Araneae: Sparassidae): systematics and zoogeography with revision of the African and Arabian species Author Moradmand, Majid text Zootaxa 2013 2013-06-17 3675 1 1 108 http://dx.doi.org/10.11646/zootaxa.3675.1.1 journal article 110557 10.11646/zootaxa.3675.1.1 253ac560-caad-4129-802d-2094c5cc69de 1175-5326 6422593 7F4D5550-8B85-4694-9482-8A125E9A2650 Genus Eusparassus Simon, 1903 Type species: Eusparassus dufouri Simon, 1932 , subsequent designation by Simon (1932) . The type species was misidentified by Simon (1903) under the name “ E. argelasius ” sensu Latreille, 1818 . The females misidentified by Latreille (1818) under the name “ Micrommata argelasia ” were type specimens which are not available. Thus, the neotype was designated from Montalvão ( Portugal ), re-described and illustrated by Moradmand and Jäger (2012a) [for more details on the nomenclature, see Moradmand and Jäger (2012b) ]. Micrommata Latreille, 1804 [part]. Latreille 1818: 517 ; Dufour 1820: 299 , pl. 2 (misidentification). Sparassus Walckenaer, 1805 [part]. Walckenaer 1830: 108 , pl. 7, fig. 1; Walckenaer 1837: 584 , 585; Simon 1874: 252 ; Simon 1880: 290 ; Simon 1897b: 388 ; Bonnet 1958: 4098 ; Levy 1989: 138 , fig. 20 (misidentification). Olios Walckenaer, 1837 [part]. Pocock 1901: 489–493 ; Lawrence 1927:42 , pls 2, 3, figs 29, 67. Cercetius Simon, 1902: 253 (description of juvenile, holotype examined from Dibba, Persian Gulf). Simon 1903: 1020 , 1023, 1026; Jäger & Kunz 2005: 170 , figs 201–204 (illustration of juvenile holotype ) [see the nomenclatural note in the description of Cercetius perezi Simon, 1902 , below]. Eusparassus Simon, 1903: 1020 , 1023, 1025. Simon 1909: 31 ; Järvi 1912: 57 , 175, fig. 49, pl. 4, figs 9, 10; 1914: 173–175; Reimoser 1919: 200 ; Petrunkevich 1928: 155; Gravely 1931: 238 ; Schenkel 1936: 9 , 283; Roewer 1928: 118 , pl. 2, figs 38–39; 1955a: 775; 1962: 4, figs 82–84; Caporiacco 1935: 216 , pl. 6, fig. 4; 1939: 353; 1941: 109, fig. 40; Denis 1937: 1050 ; 1938: 388 ; 1945: 54 ; 1947: 49 , pl. 2, fig. 12; 1958: 102, f. 30; Barrientos & Urones 1985: 356 , figs 4, 5; Jäger 1999: 1 , 4, 6; 2001: 16, 18, figs 13 a–c, ä, ö; Song et al. 1999: 467 , f. 268H, K; Jäger & Yin 2001: 132 ; Jäger & Kunz 2005: 168 , 169, figs 205–213; Urones 2006: 100 , figs 1–43; Dunlop et al. 2011: 519 , figs 1–3; Deltshev 2011: 28 ; Gabriel 2011: 9–12 , figs 2, 9; Moradmand & Jäger 2012a : figs 1–23. Eusparassus (Doubtful usage). Strand 1906a: 630 ; 1907a: 437 ; 1907b: 671 ; 1908b: 19 . Diagnosis. Eusparassus is easily diagnosable from other members of subfamily Eusparassinae by the presence of two pairs of ventral tibial spines on legs I–IV (three pairs in Pseudomicrommata , Arandisa , Leucorchestris Lawrence, 1962 and Carparachne Lawrence, 1962 ); from Olios (subfamily Sparassinae ) by a combination of characters including the presence of intermarginal denticles in some Eusparassus spp. (absent in Olios spp. ), presence of a single bristle on the anterior margin of cheliceral basal segment below fangs but that number can reach a maximum of five (mostly> 10 in Olios spp. ). However, the best characters to distinguish between these two morphologically closely similar genera are those of the copulatory structures. In Eusparassus spp. the male palp is characterized by embolus and tegulum nearly of the same length arranged as a U-shaped structure, presence of embolus membrane (EM) [EM can be considered a well developed pars pendula, personal communication with C.A. Rheims], lack of any tegulum apophysis ( Fig. 1 ); female epigyne shows two large lateral lobes (LL), and simple straight copulatory ducts leading to a more complex turning loop (TL) ( Fig. 2 ). Description . See Moradmand and Jäger (2012a) . Natural history and habitat preferences. The knowledge on the biology of stone huntsman spiders is quite scanty. They produce large silken papery retreats attached to the underside of stones or in crevices of rocks. They hide during the day in these retreats and also use them to moult in. The excuvia are mostly found within the abandoned retreats (personal observation). Females construct a sealed egg-sac inside the larger retreat and guard it until the spiderlings hatch. In E. walckenaeri ( Audouin, 1826 ) , it took nearly one month from pre-larval stage to hatching stage ( Gabriel 2011 ). Like most Sparassidae , the stone huntsman spiders are nocturnal predators. They are known from semi-arid pine forest in the Atlas Mountains and the borders of the Sahara in Northern Africa to the Wahiba sand dunes and Wadis in Arabia, from the Mediterranean area to Central Asian deserts and the slopes of the Himalayas, and throughout the Eastern and Southern African Savannah to the arid borders of the Namib and Kalahari deserts. They can occur in very high elevations above sea level (e.g., E. pontii up to 3000–4000 m in Himalayas, Moradmand & Jäger 2012a ). Earlier biological notes are restricted to some observations on the species E. walckenaeri by Gerhardt (1928 , 1933 ) who documented his observations on the mating behaviour of this species (sub Sparassus sp. from Greece ). Gabriel (2011) published his observations of the developments of spiderlings and some parasites and predators from Turkey . Copulation. The first photographical documentation of the copulation process of palp and epigyne in the genus Eusparassus is recorded and presented here. Combining knowledge of the morphology of the copulatory structures in Eusparassus spp. and the detailed documentation on how they function in action provide some valuable data on the functional morphology of the pedipalp and epigyne. Juvenile specimens of E. walckenaeri were collected by Dr Peter Jäger in the Negev desert (during the 26 th European congress of Arachnology) in September 2011 . Specimens were reared in captivity until they reached maturity in August 2012 . On the 7 th of August, the female was housed in a glass terrarium ( 30cm diameter x 20cm high) and one day later, the male was introduced into the terrarium. A few minutes later, the male started searching and tracing the female. Suddenly he attacked her and tried to grab her by the legs and chelicerae but the female autotomized one leg and escaped. He fed on the leg of the female and subsequently killed a cricket roaming in the terrarium but did not consume it. The male approached the female again. This time the female did not struggle and the male seized her, face to face, using both his legs and chelicerae. He gently bit the female’s pedicel area between prosoma and opisthosoma and held her with his legs ( Fig. 44a ). They remained in this position for a few seconds until the female was totally subdued and did not move till the end of mating. The male attempted to reach the female’s epigyne, first from her right side using his left palp but without inserting his embolus ( Figs 44b–d ). Then he shifted to the left side of the female. The process of coupling palp and epigyne was initiated by anchoring the RTA (dRTA) into the posterior margin of epigyne between the lateral lobes ( Figs 45a, d ), the male stretched his right palp next, which suddenly expanded and the embolus was inserted into the copulatory opening ( Fig. 45b ). This observation (inserting dRTA into posterior margin between lateral lobes of epigyne) gives some evidence about a similar structure in the vulva which was recently recognized in the species of the genus Sinopoda Jäger, 1999 . This structure was named membranous sac (Msa) and is supposed to hold the dRTA during copulation ( Jäger 2012 ). The Msa can be mistaken for intermediate tissue and muscles around vulva, and is usually removed during vulva preparation since its presence restricts the view on scleriotized vulva structures.. The Msa in Eusparassus species is located medially between the fertilization ducts ( Figs 11d, e ). Another modification in the female copulatory organ might be the following: Eusparassus species with a more robust dRTA have special modifications dorsally of the median septum, from a simple hyaline structure ( Fig. 2b ) to a sclerotized longitudinal band ( Fig. 16b ) and even a complex folded structure ( Fig. 36b ). Identification key to species of Eusparassus In the following key, a combination of the somatic and copulatory characters are used, nevertheless, species identification should be confirmed by checking the detailed diagnoses and descriptions given in the text for each species. The key should be used with special care when identifying females. Species descriptions of the doriae group as well as E. pearsoni ( Pocock, 1901 ) ( vestigator group), E. pontii Caporiacco, 1935 and E. xerxes ( Pocock, 1901 ) (both incertae sedis), E. dufouri Simon, 1932 and E. levantinus Urones, 2006 (both dufouri group) are given in Moradmand and Jäger (2012a) . The character ventral opisthosoma dark marking must be used with special care as preserved specimens could have been faded. Since Cercetius perezi is regarded congeneric (retained usage until ICZN decision on case 3596), this species is included in the Eusparassus key. 1. Cheliceral furrow with intermarginal denticles (e.g. Fig. 1f ).................................................... 2 – Cheliceral furrow without intermarginal denticles (e.g. Fig. 13e )............................................... 16 2. Male [unknown in E. borakalalo spec. nov. ]................................................................ 3 – Female.............................................................................................. 9 3. Palp with enlarged and bulged ST (e.g. Fig. 35a )............................................................. 4 – Palp with small and hidden ST behind T (e.g. Fig. 1a )......................................................... 6 4. dRTA bifurcated at its tip ( Figs 35a, b ) [ Zimbabwe ]............................................ jocquei spec. nov. – dRTA pointed and not bifurcated at its tip.................................................................. 5 5. ET triangular and flattened proximally and pointed distally ( Fig. 31c ) [ South Africa : Northern Cape Province ]............................................................................................... schoemanae spec. nov. – ET slender and curved at its distal end ( Fig. 29c ) [ South Africa ].................................... jaegeri spec. nov. 6. dRTA bent toward cymbium and pointed disto-ventrad ( Fig. 4a ) [Horn of Africa to Arabia]..................... laevatus – dRTA directed distad................................................................................... 7 7. Ventral opisthosoma with large solid black marking ( Fig. 57b ), ET directed distad ( Fig. 42c, f ) [Arabia and Horn of Africa].......................................................................................... Cercetius perezi – Ventral opisthosoma pale, ET directed retrolaterad (e.g. Fig. 1d )................................................ 8 8. Palp and dRTA robust, PE and AE roughly subequal ( Figs 1a–e ) [Eastern Mediterranean to Egypt and Algeria ].. walckenaeri – Palp and dRTA elongated and slender, PE distinctly larger than AE ( Figs 7a–d ) [Arabian Peninsula]..... arabicus spec. nov. 9. Epigyne with AMLL fused together anteriorly (e.g. Fig. 32a ).................................................. 10 – Epigyne with AMLL not fused together anteriorly (e.g. Fig. 2a )................................................ 14 10. Epigyne with MS clearly visible posteriorly ( Fig. 36a ) [ Zimbabwe ]................................ jocquei spec. nov. – Epigyne with MS not visible posteriorly (LL are in contact)................................................... 11 11. Vulva composed of several bulbous parts in TL ( Figs 43c, d ) [Horn of Africa to Arabia]................. Cercetius perezi – Vulva different (with single large TL)..................................................................... 12 12. MS as long as wide, CD and MS partially to fully sclerotized ( Figs 30a–d ) [ South Africa ]............... jaegeri spec. nov. – MS longer than wide and membranous, CD hyaline......................................................... 13 13. EF longer than wide ( Figs 32a , 33a ) [ South Africa : Northern Cape Province ].................... schoemanae spec. nov. – EF wider than long ( Figs 34a, f ) [ South Africa ]............................................. borakalalo spec. nov. 14. PE distinctly larger than AE ( Fig. 7d ), EF bridge present, ( Fig. 8a ) [Arabian Peninsula]............... arabicus spec. nov. – PE and AE nearly equal, EF bridge mostly absent........................................................... 15 15. MS as wide as long, MS length ¼ EF length, vulva with Gpo situated in a depression in connection with collar form a continuous ridge ( Figs 5a–c ) [Horn of Africa to Arabia]....................................................... laevatus – MS mostly longer than wide, MS length ½ of EF length, vulva with Gpo situated in a depression separated from collar part ( Figs 2a–c ) [Eastern Mediterranean to Egypt and Algeria ]............................................ walckenaeri 16. Male [unknown in syrticus , pearsoni , maynardi , pontii ]...................................................... 17 – Female............................................................................................. 35 17. Ventral opisthosoma with distinct dark marking............................................................. 18 – Ventral opisthosoma lacking distinct dark marking.......................................................... 28 18. vRTA well developed: as long as one-third of dRTA (e.g. Fig. 25a )............................................. 19 – vRTA not well developed: less than one-third of dRTA....................................................... 20 19. ET flat and wide with a pointed triangular process, dRTA robust and flattened dorso-ventrally ( Figs 27a–c ) [ Burkina Faso and Nigeria ]............................................................................. reverentia spec. nov. – ET and dRTA different ( Figs 25a–c ) [Eastern Africa: Ethiopia , Kenya and Tanzania ]......................... vestigator 20. EM with projecting bulge covering proximal end of ET in ventral view ( Figs 20a–c ) [Eastern Morocco ]........... fritschi – EM without any projecting bulge........................................................................ 21 21. ET directed proximad ( Fig. 11a )......................................................................... 22 – ET pointing in differentdirection......................................................................... 23 22. ET robust and flat, dRTA sickle-like ( Figs 11a, b ), ventral opisthosoma with V-shaped marking ( Fig. 48b ) [Western Iberian Peninsula]...................................................................................... dufouri – ET slim, dRTA more straight ( Fig. 60c ), V-shaped marking with additional median band ( Fig. 48d ) [Eastern Iberian Peninsula] levantinus 23. AE larger than or subequal as PE....................................................................... 24 – PE generally larger than AE, PLE largest.................................................................. 27 24. ET directed retrolaterad ( Figs 12a, c ) [ Morocco ]...................................................... atlanticus – ET directed ventrad ( Fig. 22c ), PLE subequal to PME........................................................ 25 25. ET flattened, dRTA bent toward cymbium, directed ventrad ( Figs 15a–c ) [Northern Algeria ]................... barbarus – ET slim, dRTA directed distad ( Figs 22a–c )................................................................ 26 26. Small to medium Eusparassus species ( 16 to 18 mm ) with ventral opisthosoma marking more solid in fresh samples and Vshaped in preserved ones (lines of marking are bold dark) ( Fig. 49f ) [North-Eastern Algeria ]................. letourneuxi – Large Eusparassus species ( 21 to 25 mm ), a vase-like black marking on ventral opisthosoma ( Fig. 56d ) [ Iran to Pakistan ].................................................................................................... xerxes 27. ET directed retrolaterad, vRTA pointed and triangular in ventral view ( Figs 17a–c , 62a ) [ Algeria to Morocco ].... oraniensis – ET directed distad, vRTA broad and not pointed ( Figs 42a–c , 66a, e )................................. Cercetius perezi 28. Embolus long and ET slender (e.g. Figs 37a–c )............................................................. 29 – ET short and robust................................................................................... 30 29. Palpal structures strongly elongated, embolus covered by slender embolus membrane ( Figs 40a–c ) [Southern Namibia ]........................................................................................... educatus spec. nov. – Embolus membrane projected into a folded part close to ET ( Figs 37a–c ) [Northern Namibia , Angola ]............. tuckeri 30. AME strikingly larger (~1.5 times) than other eyes ( Fig. 58e ) [Central Asia]................. oculatus (Kroneberg, 1875) – AME subequal to or <1.5 times larger than others........................................................... 31 31. ET proximad, long and robust ( Fig. 67c ) [ Afghanistan ]..................................... fuscimanus Denis, 1958 – ET shorter and directed in different orientations............................................................. 32 32. vRTA rounded and not well developed (e.g. Fig. 68a )........................................................ 33 – vRTA pointed and clearly triangular (e.g. Fig. 68e , see Moradmand & Jäger, 2012a : Fig 17C )........................ 34 33. ET slim ( Fig. 67e ) [ Afghanistan to Rajasthan in India ]...................................... kronebergi Denis, 1958 – ET robust ( Fig. 68a ) [ Iran , Iraq and Turkey ]............................... mesopotamicus Moradmand & Jäger, 2012 34. dRTA straight and beak-like, distal end of ET pointing distad ( Fig. 68e ) [ China : Xinjiang Uyghur ]... potanini (Simon, 1895) – dRTA with a slight bend in proximal half, ET leaf-like, distal end of ET pointing proximad ( Fig. 67a ) [Central Iran ]............................................................................................ doriae ( Simon, 1874 ) 35. Ventral opisthosoma with distinct dark marking............................................................. 36 – Ventral opisthosoma lacking distinct dark marking.......................................................... 48 36. MS widened (approximately as wide as EF), fully sclerotized and prominent (e.g. Figs 63b, d, f ), chelicerae usually with more than two thick bristles (max. five bristles) at ventral base of fangs.............................................. 37 – MS small, hyaline to partially sclerotized, chelicerae mostly with one thick bristle (max. two bristles).................. 39 37. MS heart-shaped ( Fig. 63f ), femur spination 323 [ India : Western Ghats].................................... pearsoni – MS quadrangular ( Fig. 63b ), femur spination 424........................................................... 38 38. GP separated from CD by most of its entire length ( Figs 26b, c ) [East Africa: Tanzania to Ethiopia ]............. vestigator – GP attached to CD by most of its entire length ( Figs 28b, c ) [ Burkina Faso , Nigeria ]................ reverentia spec. nov. 39. PE generally larger than AE, PLE largest..................................................................40 – AE larger than or subequal to PE........................................................................ 42 40. Vulva with several bulbous parts in the turning loop ( Figs 43b–d ) [Arabia to Horn of Africa].............. Cercetius perezi – Vulva different....................................................................................... 41 41. TL extending CD laterally (at least slightly) in dorsal view, GP small ( Figs 18b , 19b ) [ Algeria to Morocco ]...... oraniensis – TL invisible and covered by CD in dorsal view, GP enlarged ( Figs 24b, c ) [ Tunisia ]............................ syrticus 42. AMLL not encircling MS entirely, ventral opisthosoma with a vase-like black marking [ Iran to Pakistan ]............ xerxes – AMLL encircling MS entirely, ventral opisthosoma with a V-shaped or solid black marking.......................... 43 43. EF quadrangular (e.g. Fig. 23a ).......................................................................... 44 – EF rather triangular (e.g. Fig. 13a )....................................................................... 45 44. MS semicircular ( Figs 23a, f , 61f ) [North-Eastern Algeria ]............................................ letourneuxi – MS triangular ( Figs 16a ) [North-Western Algeria ]..................................................... barbarus 45. PLE very small (PLE ~1.4 times smaller than AME) ( Fig. 21d ); MS and EF mostly as long as wide ( Fig. 21a ) [ Morocco ].................................................................................................. fritschi – Eyes different; EF distinctly longer than wide.............................................................. 46 46. Lacking a sclerotized longitudinal strip on dorsal MS ( Fig. 13b ), ventral opisthosoma with a solid dark marking ( Fig. 49b ) [ Morocco ].................................................................................... atlanticus – A sclerotized longitudinal strip on MS present in dorsal view (e.g. Fig. 16b ), ventral opisthosoma with a V-shaped dark marking................................................................................................ 47 47. GP located on a continuous part distinguishable from turning loop; ventral opisthosoma with a clear V-shaped marking ( Fig. 48b ) [Western Iberia ].............................................................................. dufouri – GP situated on a semicircular process which is fused to entire body of vulva; V-shaped marking with an additional median band ( Figs 48d, f ) [Eastern Iberia ]................................................................ levantinus 48. AMLL fused together and encircling MS entirely (e.g. Fig. 65b )............................................... 49 – AMLL not fused together and not encircling MS entirely (e.g. Fig. 67d )......................................... 50 49. Vulva ducts and TL coiled and twisted ( Fig. 41c ) [ Namibia ]..................................... educatus spec. nov. – Vulva ducts and TL simple, straight and spherical, respectively ( Fig. 38c ) [ Namibia ]........................... tuckeri 50. EF bridge absent (e.g. Fig. 68d )......................................................................... 51 – EF bridge present (e.g. Fig. 68f )......................................................................... 52 51. EF distinctly longer than wide, AMLL strongly developed ( Fig. 66f ), eyes subequal [ Pakistan : Baluchistan ]............................................................................................... maynardi ( Pocock, 1901 ) – EF nearly as long as wide, AMLL not developed ( Fig. 68d ), AME strikingly largest ( Fig. 58e ) [Central Asia]...... oculatus 52. EF bridge distinctly separated from AMLL and not bordering MS ( Fig. 68b ) (see also Moradmand and Jäger 2012a : fig. 10A) ................................................................................................... 53 – EF bridge fused to AMLL and bordering MS............................................................... 55 53. EF as long as wide, AMLL not extended anteriorly ( Fig. 67b ) [Central Iran ]................................... doriae – EF longer than wide, AMLL extended anteriorly (e.g. Fig. 68b )................................................ 54 54. EF bridge distinctly separated from AMLL, approximately as long as MS length ( Fig. 68f ) [ China : Xinjiang Uyghur ] potanini – EF bridge separated from AMLL, but less than MS half length ( Fig. 68b ) [ Iran , Iraq and Turkey ].......... mesopotamicus 55. EF as wide as long or slightly wider than long ( Fig. 67d ) [ Afghanistan ].................................. fuscimanus – EF clearly longer than wide............................................................................ 56 56. MS as wide as long ( Fig. 66g ) [ Pakistan : Karakoram; India : Ladakh]......................................... pontii – MS distinctly wider than long ( Fig. 67f ) [ Afghanistan to Rajasthan in India ]............................... kronebergi