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«Taxonomic Revision, Molecular Phylogeny and Zoogeography of the huntsman spider genus Eusparassus (Araneae: Sparassidae) Dissertation for attaining ...»

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– MS distinctly wider than long (Fig. 67f) [Afghanistan to Rajasthan in India]..............kronebergi Results: chapter 3.2: Systematics and zoogeography with revision of Afro-Arabian species Taxonomy & Systematics Family Sparassidae Bertkau, 1872 Subfamily Eusparassinae Järvi, 1912 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.

Results: chapter 3.2: Systematics and zoogeography with revision of Afro-Arabian species 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 chapter 3.1: 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.

Results: chapter 3.2: Systematics and zoogeography with revision of Afro-Arabian species 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 26th European congress of Arachnology) in September 2011.





Specimens were reared in captivity until they reached maturity in August 2012. On the 7th 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).

Results: chapter 3.2: Systematics and zoogeography with revision of Afro-Arabian species Species groups Species groups are recognized by a combination of somatic characters and those of copulatory structures of male and female. Six species groups are proposed: walckenaeri group (3 species), dufouri group (8 species), vestigator group (3 species); jaegeri group (4 species); tuckeri group (2 species) and doriae group [7 species, description of members presented in Moradmand & Jäger (2012a)]. Eusparassus xerxes (Pocock, 1901), E. pontii Caporiacco, 1935 and Cercetius perezi Simon, 1902, could not be placed in any proposed species groups and are listed at the end of the description. In the following section species are listed according to species-groups. For every group the diagnosis, species composition and distribution range are presented. In this paper the representatives from Africa and Arabia are re/described. For description of the species from Europe, the Middle East, Central and South Asia, see Moradmand and Jäger (2012a).

walckenaeri species group Diagnosis. Chelicerae with intermarginal denticles (Figs 1f, 5e, 7e, 10c); ventral opisthosoma lacking any dark marking (Fig. 46d); male palp with ST small in size (compare to jaegeri group) and situated behind EM (Figs 1b, 4a, 7a, 9a); AMLL of epigyne not fused (Figs 2a, 5a, 8a, 10a);

there is no GP, and Gpo situated in a depression on vulva (Figs 2c, 3b, 5c, 8c, 9f, 10e).

Species composition. Three species: Eusparassus walckenaeri (Audouin, 1826), E. laevatus (Simon, 1897) comb. nov. and E. arabicus spec. nov.

Distribution. Eastern Mediterranean to North-Eastern Africa and Arabian Peninsula (Fig. 70a).

–  –  –

Philodromus walckenaerii Audouin, 1826: 390, pl. 6, fig. 1 (description of female, Egypt; no type series designated).

Philodromus linnaei Audouin, 1826: 390, pl. 6, fig. 2 (description of male, Egypt, no type series designated) [synonymy by Simon 1906].

Results: chapter 3.2: Systematics and zoogeography with revision of Afro-Arabian species Drassus civilis Reuss, 1834: 207 (description of juvenile; holotype, immature, examined) [synonymy by Levy 1989].

Sparassus walckenaeri (Audouin). Walckenaer 1837: 585 (transfer); Pavesi 1880: 364; 4; Levy 1989:

132–138, figs 3–18.

Ocypete tersa C. L. Koch, 1837: 83, fig. 305 (description of female; from Greece, type not available) [synonymy by Levy 1989]. C. L. Koch 1845: 39, figs 980–981.

Sparassus cambridgii Simon, 1874: 257 (description of juvenile, from Egypt) [synonymy by Simon 1880].

Sparassus validus Thorell, 1875a: 80 (description of female; holotype female, MZH, examined) [synonymy by Levy 1989]. Thorell 1875b: 124.

Sparassus cognatus O. Pickard-Cambridge, 1876: 588 (description of female; syntypes, one female and 10 immatures, Egypt, not examined) [synonymy by Levy 1989].

Sparassus extensipes Karsch, 1880: 383, pl. 12, fig. 12. (description of male, holotype, male, Egypt:

Cairo, not examined) [synonymy by Simon 1906].

Sparassus walckenaerius (Audouin). Simon 1880: 292.

Sparassus tersa (C. L. Koch). Simon 1880: 291 (in part, material from Greece, MNHN, examined).

Sparassus linnaei (Audouin) Kulczyn'ski. 1901: 43 (transfer) (one male examined from Cairo in MIZ).

Eusparassus walckenaeri (Audouin). Simon, 1906: 1168; Strand 1908b: 24; Denis 1947: 50, pl. 2, figs 14–16.); Deltshev 2011: 28; Gabriel 2011: 9–12, figs 2, 9; Moradmand and Jäger 2012a: 2453, figs 1B, 5– 6, 23A (designation of neotype, neotype male from EGYPT examined).

Eusparassus tersa (C. L. Koch). Järvi 1912: 57, fig. 48, pl. 4, figs 4–8 (transfer); Järvi 1914: 173.

Heteropoda civilis (Reuss). Strand 1916: 36 (unjustified combination).

Type material. Neotype of E. walckenaeri (subsequent designation by Moradmand and Jäger 2012a): male, EGYPT: Muhafazat al Qahirah: Cairo [N 30° 3', E 31° 15'], 1971 (SNSD 52);

Holotype of Drassus civilis (designated by Reuss 1834): immatere, EGYPT: Sinai: Tor, 1827 Rüppell leg. (SMF 4575); Holotype of Sparassus validus (designated by Thorell, 1875a): female, TURKAY: Taurus Mountains, (label: Taur. Merid., Ent.etikett nr=232), Nordmann leg. (MZH 20.492).

Other material examined. (16♂♂, 18♀♀): EGYPT: 1♂, 1♀ (MM 121), with same data as for neotype (SNSD); Muhafazat al Qahirah: 1♂, Helwan, March 1901, in house (NHM); 1♂, Fayid,

gravel area, 15 April 1947, J.H. Graham leg. (NHM 1948.10.11.7); 1♂, Cairo, with label:

―Sparassus linnaei, Cairo, det. Kulczyński, F.1691‖ (MIZ 212984); Muhafazat al Qina: 1♂ (MM3), 2♀♀ (MM17, MM206), Luxor (Al Uqsur), Thebes (SMF 5557); Muhafazat al Jizah:

Results: chapter 3.2: Systematics and zoogeography with revision of Afro-Arabian species 1♂ (MM7), Al Jizah (=Gizeh) (SMF 5576); Muhafazat al Suways: 1♀, Djebel Genaifa, W of Suez Canal, 9 June 1947, G. Konieczny leg. (SMF); 3♂♂, 7♀♀, 1 juv., ―Prof. J.

Omer-Cooper SIWA Expedition 1935‖, Libyan Desert, Siwa Oasis, 20–30 May (2♀♀, 1 juv.: NHM 1936.2.12.94–96, 2♂♂: NHM 1936.2.12.97–98), 29 April (1♂: NHM 1936.7.10.6), 22 July (1♀:

NHM 1936.2.12.176), August (1♀: NHM 1936.2.12.157), 22 August (1♀: NHM 1936.2.12.177), 30 August (1♀: NHM 1936.2.12.560); 1♂, 2♀♀, 18 October 1985, Sörensen & Kollend leg.

(ZMUC); LIBYA: Baladiyat al Kufrah: 1♂, Jebel Uweinat, Karkur Talh [N 21° 54’, E 024° 58’], ―Mission Scientifique Belge‖, 25 October 1968 (MRAC 135886); 1♀, Baladiyat Shahhat, Susa (=Soussa) (MNHN 227.61); SUDAN: Wilayat al Khartoum:1♂, Khartoum, July 1909, S.S.

Floman leg. (NHM 09.10.13.1048); 1♀, 1 juv., Khartoum, October 1979, El Hamin El Rayal leg.

(MRAC 152086–87); Wilayat al Bahar al Ahmar: 1♀, Gabet al Maadin, 20 km S of Mohammed

Qul., 21 September 1960, Prof. J. C. Thompson leg. (NHM); ALGERIA: Wilaya d' El Oued:

1♂, 1♀, El Oued, C.I.E. coll: 13593-1559, 1953, L. Past al Ag leg. (NHM); Wilaya d' Illizi: 1♂, Fort Polignac, C.I.E. coll: 13593-3585, September 1953, L. Pastal Ag leg. (NHM); Wilaya d' Tamanghasset: 1♀, 1 juv., Tamarnaset, C.I.E. coll: 13593, 1953, L. Past al Ag leg. (NHM);



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