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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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Spination. Palp 131, 001, 1111; Legs: Femur I–III 323, IV 321; Patella I–IV 000(1)/101; Tibia I– IV 2224; Metatarsus I–III 2(1)024, IV 3034/30(1)36.

Results: chapter 3.2: Systematics and zoogeography with revision of Afro-Arabian species Palp. As in diagnosis with cymbium longer than tibia; tegulum shorter than embolus; dRTA long and slender with slight median bent, vRTA weakly developed (Figs 42a, b); tip of embolus

FIGURE 43. Cercetius perezi Simon, 1902: (a–c) female from Oman: Mudhaybi; (d) female from Oman:

Fallah. (a) epigyne, ventral; (b) vulva, dorsal; (c–d) left vulva, anterio-dorso-lateral.

Results: chapter 3.2: Systematics and zoogeography with revision of Afro-Arabian species hyaline and worm-like, embolic membrane consisting of folded layers distally; conductor elongated (Fig. 42c).

Female (ranges: n=7, single measurement: MM 30):

Measurements. Medium to large sized; total length 21.5–28.0, prosoma length 9.5–12.5, prosoma width 8.6–11.1, anterior width of prosoma 5.7–7.5, opisthosoma length 12.0–15.5, opisthosoma

width 7.8–11.0. Eye diameters: AME 0.81, ALE 0.84, PME 0.70, PLE 0.88; eye interdistances:

AME-AME 0.33, AME-ALE 0.20, PME-PME 0.85, PME-PLE 0.95, AME-PME 0.90, ALE-PLE 0.75, clypeus height at AME 0.65, clypeus height at ALE 0.70.

Chelicerae. Chelicerae with 2 anterior and 3 or 4 posterior teeth, Cheliceral furrow with 1 or 2 intermarginal denticles close to anterior teeth or without denticles. Basal segment of chelicerae at distal end retro-marginally with a single bristle.

Legs. Leg formula: II I IV III. Measurements of palp and legs: Palp 16.3 [5.0, 2.5, 3.1, 5.7], I 46.4 [13.5, 6.5, 11.4, 11.8, 3.2], II 51.9 [14.9, 6.6, 14.0, 13.2, 3.2], III 44.4 [13.7, 6.0, 11.1, 10.8, 2.8], IV 45.6 [14.3, 5.8, 10.9, 11.4, 3.2].

Spination. Palp 131, 001, 1111, 1013; Legs: Femur I–III 323, IV 321; Patella I–IV 101; Tibia I– IV 22(1)24; Metatarsus I–III 2(1)024, IV 3034/3036.

Epigyne/vulva. As in diagnosis with epigyne composed of two large triangular lateral lobes, epigynal field slightly longer than wide, anterior margin of lateral lobes fused together and encircling MS entirely, epigynal field bridge (EFB) present and not separated from anterior margin of lateral lobes (Figs 43a, b).

Colouration. A freshly collected single male was obtained from Somalia whose prosoma and dorsal opisthosoma is creamy-white with shiny white hairs dorsally on legs (Fig. 57a); preserved specimens are reddish-brown spiders with darker scopula hairs on metatarsus and tarsus; prosoma margins, anterior part of prosoma around eyes, chelicerae and dorsal side of femora covered with dense white hairs (Figs 57a, c), in contrast, sternum, coxae of legs and basal segment of chelicerae covered with dense black hairs, ventral opisthosoma with large black marking posterior to and around epigastric furrow (Fig. 57b).

Systematic position. The somatic features and the copulatory organs of Cercetius perezi correspond well with the Eusparassus delimitation as given in Chapter 3.1 [Moradmand and Jäger (2012a)] and in this paper. The presence of intermarginal denticles of chelicerae, eyes arrangement, leg formula, spination pattern and the presence of dark marking at ventral opisthosoma are all somatic characters which are present in the immature holotype and the newly Results: chapter 3.2: Systematics and zoogeography with revision of Afro-Arabian species discovered adult specimens from the type locality. However, C. perezi cannot be affiliated with any of the known species groups. This species embodies some synapomorphies of the dufouri group (e.g. dark marking of ventral opisthosoma, epigyne with AMLL encircling MS entirely) and also some of the jaegeri and walckenaeri groups (presence of intermarginal denticles in some specimens). Cercetius perezi is likely to belong to an intermediate lineage among the noted Eusparassus species groups. The geographical distribution between the three groups mentioned above supports this hypothesis.

Currently known distribution and habitats. Eastern and southern Arabian Peninsula in the United Arab Emirates (type locality), Oman (new country record) and Yemen (new country record), horn of Africa in Somalia and Djibouti (new country record) (Fig. 72b). Specimens were collected in wadis, sandy substrates, gravel plains and from under stones in deserts.

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Sparassus bicorniger Pocock, 1898: 519, pl. 41, fig. 9 (description and illustration of male), [holotype male, label: type, Ndi, Weiss Rd. Camp, by Steuart Betton/ Nd (Weiss Rd Camp), Aug 4–25, 1897, NHM, examined].

Eusparassus biocorniger (Pocock) Strand 1908b: 22 (unjustified combination).

Remarks. The species cannot be categorized in any known Sparassidae genus, and it certainly does not belong to the genus Eusparassus. More likely it is a member of an undescribed genus which would be grouped within the subfamily Sparassinae, when more material (especially the conspecific female) from the type locality is recovered. The holotype was most likely collected between Mombasa and Lake Victoria in Kenya, formerly known as ―British East Africa‖.

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

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Eusparassus laterifuscus Strand, 1908a: 5 (description of juvenile) [subadult male holotype, Madagascar, 18 Decemper 1885, A. Stumpff leg., SMF 4571, examined].

Remarks. The subadult male holotype was collected from Madagascar. The combination of somatic characters clearly distinguishes it from Eusparassus: three anterior teeth in chelicerae (two in Eusparassus), a patch of several intermarginal denticles close to anterior teeth (scattered or in a line in Eusparassus, if present) and the number of ventral tibial spines: I–II 8, III–IV 6 (I– IV 4 in Eusparassus); the species is tentatively classified in “Rhitymna” saccata group and is actually an undescribed genus endemic to Madagascar (Peter Jäger, unpublished data). There is no other record of the genus Eusparassus from Madagascar.

“Eusparassus” ubae Strand, 1906 nomen dubium Eusparassus ubae Strand, 1906a: 684 (description of female) [type from Uba in East Africa, lost, see Renner 1988: 322]; Strand 1908c: 41, pl. 2, fig. 8 (redescription and illustration of epigyne).

Remarks. The sketchy illustration of the epigyne by Strand (1908c) reveals that this species does not belong to the genus Eusparassus, as it lacks the diagnostic triangular lateral lobes and the median septum is visible throughout median line posteriorly. However, it cannot be affiliated with any sparassid genus.

“Eusparassus” palystiformis Strand, 1907 nomen dubium Eusparassus palystiformis Strand, 1907a: 541 (description of female) [type from Capeland in South Africa, Museum Lübeck, lost]; Strand 1907b: 671.

Remarks. According to the original description, this species could not be placed in the genus Eusparassus, as its legs have three pairs of ventral tibial spines (two pairs in Eusparassus).

Results: chapter 3.2: Systematics and zoogeography with revision of Afro-Arabian species Strand (1907a) expressed doubts on placing his new species in Eusparassus by using a question mark.

Remarks on Ethiopian types of Strand. The type specimens of the following four species were unfortunately destroyed during World War II (Renner 1988). The excursion of the author to Ethiopia (June 2011) as well as investigations in major spider collections in Europe resulted in the recognition of two known valid species of Eusparassus distributed in the country, namely E.

laevatus comb. nov. (East and North-East), and E. vestigator comb. nov. (South). These two species were described before those of Strand. Therefore, Strand’s species are most likely junior synonyms of E. laevatus comb. nov. and/or E. vestigator comb. nov..

“Eusparassus” cornipalpis Strand, 1906 nomen dubium Eusparassus cornipalpis Strand, 1906a: 631 (description of male), [type from Mane River in Ethiopia, lost, see Renner 1988: 322].

“Eusparassus” nigrichelis Strand, 1906 nomen dubium Eusparassus nigrichelis Strand, 1906a: 631 (description of female), [type from Mane River in Ethiopia, lost, see Renner 1988: 322].

“Eusparassus” fulviclypeus Strand, 1906 nomen dubium Eusparassus fulviclypeus Strand, 1906a: 630 (description of male and female), [types from Mane River and Ginir-Daua in Ethiopia, lost, see Renner 1988: 322].

Olios fulviclypeus (Strand). Caporiacco 1940: 843 (transfer).

“Eusparassus” subadultus Strand, 1906 nomen dubium Eusparassus subadultus Strand, 1906a: 631 (description of subadult female), [type from Maki-Abassa Lake in Ethiopia, lost, see Renner 1988: 322].

Results: chapter 3.2: Systematics and zoogeography with revision of Afro-Arabian species Remarks on West-African types of Strand. The descriptions of the following two species by Strand (1906b) are based on a highly variable character, namely the number of cheliceral retromarginal teeth. Eusparassus spp. show intra and interspecific variation in this character [three to six teeth (three larger and one to three smaller ones)]. Unfortunately, both type specimens were destroyed in Stuttgart (Renner 1988).

“Eusparassus” quinquedentatus Strand, 1906 nomen dubium Eusparassus 5-dentatus Strand, 1906b: 71–73 (description of female from Ghana [sub Gold Coast], lost, original incorrect spelling).

Eusparassus quinquedentatus (Strand) Roewer 1954: 675 (justified emendation).

“Eusparassus” sexdentatus Strand, 1906 nomen dubium Eusparassus 6-dentatus Strand, 1906b: 73 (description of juvenile, from Togo: Lome, lost, original incorrect spelling).

Eusparassus sexdentatus (Strand) Roewer 1954: 675 (justified emendation).

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Replacement name for Eusparassus concolor Caporiacco, 1939: 353 (description of subadult male) [subadult male holotype, Ethiopia: Moyale, 13 May 1973, Prof. Zavattari leg., MZUF 212, examined] (preoccupied by Olios concolor Keyserling, 1884: 682, pl. 21, fig. 29).

Etymology. The largest Sparassidae genus in terms of species number, Olios, needs a comprehensive revision to uncover its hidden diversity and several misplacements. ―Quesitio‖ is the Latin translation for the term ―investigation‖, referring to the need for taxonomic revision of Olios spp. Noun in apposition.

Remarks. The subadult male holotype was collected by Prof Edoardo Zavattari from Borana region in Southern Ethiopia. The combination of somatic characters revealed that the specimen Results: chapter 3.2: Systematics and zoogeography with revision of Afro-Arabian species belongs to the genus Olios based on eye arrangement, equal length and width of prosoma, absence of intermarginal denticles, presence of two to five thick bristles at retromarginal side of chelicerae basal segment and spotted legs. The new combination is a secondary homonym of Olios concolor Keyserling, 1884 (currently considered a junior synonym of O. giganteus Keyserling, 1884), therefore a replacement name is proposed here.

Systematics and zoogeography

With completion of this revision, 30 species of the stone huntsman spiders, genus Eusparassus are known (including C. perezi), of which 27 species are classified into six species groups and the rest three species are listed as incertae sedis. All species groups show a continuous range of distribution, fully to partially separated from nearby groups. The intermarginal denticles of chelicerae are present in just two species groups namely walckenaeri and jaegeri (present also in some specimens of C. perezi, but in different pattern). Thus, it could be assumed that these two groups are phylogenitically closely related. Consequently, the recently discovered Eusparassus fossil (amber), E. crassipes, is probably allied to one of these groups, since its chelicerae have distinct intermarginal denticles (Dunlop et al. 2011: figs 2e–f). The jaegeri group is endemic to Southern Africa (Fig.71b) which is far from the locality of E. crassipes in Northern Europe.

Thus, E. crassipes is probably closely related to the walckenaeri group whose distribution range extends into the Eastern Mediterranean region (Fig.70a). The remaining four species groups lack any intermarginal denticles on their chelicerae. Nevertheless, the presence of intermarginal denticles could be also the result of homoplasy and gain and loss of the character might have been taken place several times during the evolution of the members. The dufouri and vestigator groups are related in terms of the presence of the dark marking ventrally on the opisthosoma and the spination of the legs femora (I–IV 424, exception E. pearsoni 323). The disjunctive distribution of the isolated member of vestigator group, E. pearsoni in India, far from its closest relatives in Eastern Africa (Fig. 71a) can be explained by the following hypothesis: the occurrence of this species in Indian plate is a secondary distribution of its ancestral stock from Eastern Africa. The Indian subcontinent was not totally isolated from Africa after its separation from Gondwanaland. India was reattached to northeast Africa via Greater Somalia around 65–60 MYA (million years ago), on its northward drift toward Eurasia (Briggs 2003). The hypothesized Results: chapter 3.2: Systematics and zoogeography with revision of Afro-Arabian species ancestor of E. pearsoni may have dispersed from Eastern Africa to Western India at that time. A similar scenariao was proposed for the distribution of the genus Mallinella Strand, 1906 (Zodariidae) by Dankittipakul et al. (2012). The doriae group - distributed in the Middle East to parts of Central and South Asia - might have evolved from the walckenaeri group by losing their intermarginal denticles. The tuckeri group represents an endemic lineage in Southwest Africa (Fig.71b). Since most parts of the distribution range of Eusparassus species are not explored yet, more species are to be expected, especially in the transition zones between different species groups.

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