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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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Collections and curators CRB — Collection of Robert Bosmans, Brussells IOZB — Institute of Zoology, Chinese Academy of Sciences, Beijing (Li Shuqiang) MCSN — Museo Civico di Storia Naturale ―Giacomo Doria‖, Genoa (Maria Luisa Tavano) MHNG — the Muséum d‘histoire naturelle, Genève (Peter Schwendinger) MIZ — Zoological Museum, Polish Academy of Science, Warsaw (Dominika Mierzwa) MVHN — Museu Vallencià d‘Historia Natural, Vallencià (Sergio Montagud Alario) MMB — Moravian Museum, Brno (Petr Baňař) MNCN — Museo Nacional de Ciencias Naturales, Madrid (Javier Sánchez Almazán) MNHN — Muséum National d‘Histoire Naturelle, Paris (Elise-Anne Leguin, Christine Rollard) MNM — Museo Civico di Storia Naturale di Milano, Milan (Andrea Sabbadini, Carlo Pesarini) MZH — Finish Museum of Natural History, University of Helsinki (Ritva Talman) NHM — Museum of Natural History, London (Janet Beccaloni) NHMW — Naturhistorisches Museum, Vienna (Christoph Hörweg) NRM — Swedish Museum of Natural history, Stockholm (Gunvi Lindberg, Kjell Arne Johanson) SMF — Senckenberg Research Institute, Frankfurt am Main (Julia Altmann, Peter Jäger) SNSD — Senckenberg Naturhistorische Sammlungen, Dresden (Katrin Schniebs) SZMN — Siberian Zoological Museum, Novosibirsk (Galina Azarkina) ZIP — Zoological Institute, Academy of Science, St. Petersburg (Kirill Mikhailov) ZMB — Museum für Naturkunde, Berlin (Anja Friederichs, Jason Dunlop) ZMMU — Zoological Museum of Moscow state University, Moscow (Kirill Mikhailov) ZMUC — Zoological Museum, University of Copenhagen (Nikolaj Scharff) ZSM — Zoologische Staatssammlung München, Munich (Stefan Friedrich, Roland Melzer)

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Micrommata [part] – Latreille, 1818: 517; Dufour, 1820: 299, pl. 2 (misidentification).

Sparassus [part] – Walckenaer, 1830: 108, pl. 7, fig. 1; Walckenaer, 1837: 584, 585; Simon, 1880: 290; Bonnet, 1958: 4098; Levy, 1989: 138, fig. 20. (misidentification).

Eusparassus Simon, 1903: 1020, 1023, 1025– Strand, 1906: 630; Strand, 1907: 437 ; Strand, 1908: 19; Simon, 1909: 31; Järvi, 1912: 57, 175, fig. 49, pl. 4, figs 9, 10; Järvi, 1914: 173– 175; Reimoser, 1919: 200; Petrunkevich, 1928: 155; Gravely, 1931: 238; Schenkel, 1936: 9, 283; Roewer, 1928: 118, pl. 2, figs 38–39; Roewer, 1955: 775; Roewer, 1962: 4, figs 82–84;

Caporiacco, 1935: 216, pl. 6, f. 4; Caporiacco, 1939: 353; Caporiacco, 1941: 109, f. 40;

Denis, 1945: 54; Denis, 1947: 49, pl. 2, f. 12; Denis, 1958: 102, f. 30; Barrientos & Urones, 1985: 356, figs 4, 5; Jäger, 1999: 1, 4, 6; Song et al. 1999: 467, f. 268H, K; Jäger, 2001: 16, 18, figs a–c, ä, ö; Jäger & Yin, 2001: 132; Jäger and Kunz, 2005: 168, 169, figs 205–213;

Urones, 2006: 100, figs 1–43; Dunlop et al. 2011; Deltshev, 2011: 28; Gabriel, 2011: 9–12, figs 2, 9.

Notes. Simon (1903) created the generic name Eusparasus to substitute it for the name Sparassus; because he suspected that Sparassus was a junior synonym of Micrommata (Simon 1903: 1020). He designated the type species as Eusparassus argelasius denoted by a new replacement name (nomen novum) for misidentified Micrommata argelasia (published in Latreille 1818). Since this species could be mistaken with Olios argelasius, the type species of Olios Walckenaer, 1805 (sub Sparassus argelasius) Simon (1932) proposed Eusparassus dufouri as a new species.

Type species. Eusparassus dufouri Simon, 1932 by original designation in Simon (1903) sub E. argelasius, female from Spain.

Extended diagnosis. Eusparassus spp. can be diagnosed from the other two monotypic genera of Eusparassinae by the number of ventral tibial spines: I–IV 4 (6 in Arandisa and Pseudomicrommata) and by relative diameters of AME which is subequal to or larger than ALE (smaller than ALE in the other two genera); Eusparassus spp. are recognisable by the shape of copulatory structures: parallel embolus and tegulum constructing a U-shaped structure, embolus membrane covering partially embolus tip; dRTA strong and straight, in

Results: Chapter 3.1: Eusparassus in Eurasia

contrast to dRTA, vRTA small and weakly developed (Figures 2A, 11G, 19A); Female epigyne characterized by two large triangular lateral lobes, LL parallel and in contact on the median longitudinal suture,diverging strongly at posterior margins and slightly at anterior margins, and circumscribing MS entirely (Figures 3A, 4C) or partially (Figures 5F, 8A, 16A, 21A); in vulva, dorsal view, two parallel copulatory ducts, straight and fully or partially hyaline, folded and membranous (Figure 3B).

Redescription. Medium to large Sparassidae, body length 10 mm (e.g. E. oculatus) to 30 mm (e.g. E. xerxes comb. nov.); prosoma slightly longer than wide; Leg length formula 2 4 1 3 (most of species) or 2 4=1 3; eyes arranged in two rows, anterior row slightly recurved and posterior row relatively straight, eyes about subequal in size, AME slightly larger than or subequal to ALE and PME smaller than PLE; Basal segment of chelicerae at distal retromarginal end with 1 (Figures 2E, 7B) to 3 or 4 thick bristles (Figures 19E, 22D), in most species onlyone bristle; Chelicerae with 2 anterior and 3 to 6 posterior teeth, Cheliceral furrow with (Figure 4B) or without denticles (Figures 2E, 9B, 21D); ventral tibial spines: I–

IV 4, spination of other parts variable but in most species: Palp 131, 101, 1111, 1013; Legs:

Femur I–III 323, IV 322; Patella I–IV 101; Tibia I–IV 2024/2224; Metatarsus I–III 2024, IV 3034/3036; male palp as in diagnosis with embolus originating at 6:30 – o´clock – position running first distally and bent retrolaterally, tip of embolus pointing in various angles and with diverse shapes, embolus and tegulum form a U-shaped structure in ventral view; small and hyaline conductor situated at distal end of tegulum and covering partially tip of embolus (Figures 11E, 12B) ; Female epigyne consisting of two large triangular lateral lobes, LL parallel and in contact on the median longitudinal suture; MS soft and hyaline (Figure 5F) or hard and sclerotised (Figure 22E), EF fusing anteriorly and constructing EFB (Figures 17F, 20A) or not (Figure 16A); internal duct system with glandular pores situated in a depression (Figures 5G, 6B, D) or on a projection (Figures 2C, 8B, 22F).

Colouration. Pale grey to dark brown spiders, with uniform colouration of body (Figure 1C) or clearly patterend body and banded legs (Figure 1B), ventral opisthosoma with distinct dark marking (Figure 23B–D) or pale (Figure 23A), dorsal opisthosoma with a pattern of small chevrons on posterior half.

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Micrommata argelasia (Walckenaer, 1805) – Latreille, 1818: 517 (misidentification, description of

female, Spain); Dufour, 1820: 299, pl. 2 (misidentification).Sparassus argelasius – Walckenaer, 1830:

108, pl. 7, fig. 1 (misidentification, male); Walckenaer, 1837: 584 (misidentification, female); Simon, 1875: 334 (misidentification); Simon, 1880: 290 (misidentification) Eusparassus argelasius – Simon, 1903: 1020, 1025 (type species designation, new replacement name,

description of the genus) – Järvi, 1912: 57, 175, figs 9, 10, 49, pl. 4; Järvi, 1914: 175; Roewer, 1928:

118, pl. 2, figs 38–39 (misidentification).

E. dufouri Simon, 1932: 890 (new replacement name). – Barrientos & Urones, 1985: 356, figs 4, 5;

Urones, 2006: 102, figs 1–24.

Sparassus dufouri (Simon) – Levy, 1989: 138, fig. 20.

Type material: (syntype females unavailable, see notes below), Neotype: male (SD 815), Portugal: Distrito de Portalgere, Montalvão, [39˚36' N, 07˚31' W] 6 May 2011, S. Henriques leg. (SMF) Other material examined. PORTUGAL: 1♂ (SD 834), 1♀ (SD 822), with same data as for neotype (SMF); 1♂, Pulo do Lobo, May 2011, S. Henriques leg. (SMF, SD 838); 1♀,Distrito de Beja: Serpa, Altenju, May 2011, S. Henriques leg. (SMF, SD839); 1♂, 1♀, Pomarao, 120 m, (37°34.5′ N 7°32.100′W) 19–22 May 2006, Cardoso et al. leg. (ZMUC); SPAIN: 1♂, 1♀, Huelva Province: Alajar, Aracena, (37°53′28″N 6°33′40″W) 7 July 1969, A. Senglet leg.

(MHNG); 2♂♂, Jaén Province: Sierra de Cazorla, Guadalquivir, (37°56′12″N 02°57′30″W), 24 July 1971, A. Senglet leg. (MHNG); 1♂, 1♀, Cordoba, 3 June 1909 (MNCN); 1♀, Rabida, June 1959, V. Buddenbrock leg. (SMF).

Diagnosis. Closely related to E. levantinus but differing from it by much more stout embolus tip and more sickle-like dRTA in ventral view (Figure 2A, C); EM sheath-like and covering part of embolus tip in retrolateral view (Figure 2B); vulva differing from that of E. levantinus by glandular process located on a continuous part distinguishable from turning loop (Figure 3C).

Redescription. Male (n=8) Medium-sized Eusparassus species; total length: 9.9–13.8, prosoma length 5.5–6.8, prosoma width 5.3–6.6, anterior width of prosoma 2.6–3.5, opisthosoma length 4.5–7.0, opisthosoma width 3.0–4.5. Eyes subequal, eye diameters (neotype): AME 0.40, ALE 0.32, PME 0.31, PLE 0.34; eye interdistances: AME-AME 0.22, AME-ALE 0.10, PME-PME 0.43, PME-PLE

Results: Chapter 3.1: Eusparassus in Eurasia

0.42, AME-PME 0.32, ALE-PLE 0.22, clypeus height at AME 0.20, clypeus height at ALE 0.27.

Chelicerae with 2 anterior and 3 posterior teeth, cheliceral furrow without denticles; Basal segment of chelicerae at distal end close to base of fangs with 1 bristle (Figure 2E).

Leg formula: 2 4 1 3. Measurements of palp and legs (neotype): Palp 8.3 [2.7, 1.2, 1.0, 3.4], I 25.9 [7.2, 2.3, 6.8, 7.3, 2.3], II 28.7 [8.3, 2.9, 7.6, 7.5, 2.4], III 24.3 [7.2, 2.6, 6.1, 6.3, 2.1], IV 26.7 [7.9, 2.3, 6.8, 7.4, 2.3].

Spination. Palp 131, 000/001, 1111; Legs: Femur I–III 323/424, IV 321/322/422; Patella I–IV 000(1)/101; Tibia I–IV 2024/2224; Metatarsus I–III 1014/2024, IV 3034/3(4)036.

Palp. as in diagnosis with cymbium nearly two times longer than tibia; tegulum shorter than embolus and tip of embolus proximad, embolic projection consists of a large sheath-like part distally and a hyaline part proximally (Figure 2A– C).

Female (n=6) Total length: 16.2–17.5, prosoma length 8.0–8.5, prosoma width 6.7–7.7, anterior width of

prosoma 4.3–4.5, opisthosoma length 8.2–9.0, opisthosoma width 4.5–6.0. Eye diameters:

AME 0.45, ALE 0.41, PME 0.34, PLE 0.40; eye interdistances: AME-AME 0.35, AME-ALE 0.16, PME-PME 0.60, PME-PLE 0.58, AME-PME 0.48, ALE-PLE 0.42, clypeus height at AME 0.27, clypeus height at ALE 0.35.

Chelicerae with 2 anterior and 3 or 4 posterior teeth, Cheliceral furrow without denticles.

Basal segment of chelicerae at distal end close to base of fangs mostly with 1 bristle or 2 bristles.

Leg formula: 2 4 1 3. Measurements of palp and legs: Palp 8.7 [2.3, 1.5, 1.7, 3.2], I 26.1 [7.5, 3.4, 6.2, 6.8, 2.2], II 28.5 [8.5, 3.5, 7.0, 7.3, 2.2], III 24.2 [7.4, 3.2, 5.7, 5.8, 2.1], IV 26.7 [8.0, 3.1, 6.3, 7.1, 2.2].

Spination. Palp 131, 000 (001), 1111, 1013; Legs: Femur I–III 323/(3)424, IV 322(1)/422;

Patella I–IV 000(1)/101; Tibia I–IV 1014/2024; Metatarsus I–III 2024, IV 3034/3036.

Epigyne/vulva. As in diagnosis, epigyne is longer than wide, AMLL are fused together and circumscribe MS entirely, EFB present and combined with AMLL (Figure 3A).

Colouration. Olive-brown with clearly banded legs; ventral opisthosoma with a V-shaped dark marking (Figure 23C).

Results: Chapter 3.1: Eusparassus in Eurasia

FIGURE 2. Eusparassus dufouri Simon, 1932, neotype male from Chanca, Portugal (SMF).

(A) left palp, ventral, (B) left palp, retrolateral, (C) tip of embolus and conductor, ventral, (D) eye arrangement, (E) left chelicera, ventral. Abbreviations: C — Conductor, dRTA — dorsal retrolateral tibial apophysis, E—Embolus, EM—Embolus membrane, ET— Embolus tip, H—Haematodocha, SpD— Sperm duct, ST— Subtegulum, T—tegulum, vRTA — ventral retrolateral tibial apophysis.

Scale bars: (A, B, D, E) 1 mm, (C) 0.5 mm.

Results: Chapter 3.1: Eusparassus in Eurasia

Taxonomic notes. In the description of the genus Sparassus, Walckenaer (1805: 40) just listed Sparassus argelasius without a description (nomen nudum) along with the following nominal species: S. samaragdulus (Fabricius, 1793), S. pallens (Fabricius, 1794), S. roseus (Clerck, 1757) and S. ornatus (Walckenaer, 1802) [for more details see Jäger (1999: 3)]. One year later (1806: 146, table 2) he published a description and illustration of a male under the name Sparassus argelasius, a misidentification that was later transferred to the genus Olios.

Walckenaer‘s original description of Sparassus was actually based on species of the previously established genus Micrommata Latreille, 1804 and a single male of Olios argelasius. Latreille (1818) examining two female specimens from Spain tried to describe the female of Walckenaer‘s species, ―Sparassus argelasius”, and transferred it to Micrommata (sub Micromata argelasia), but he failed to identify it correctly. This misidentification was pointed out later by Simon (1903: 1025), who described his new genus Eusparassus, cited Latreille‘s description and indicated the type species as E. argelasius Latreille. Nevertheless, Latreille‘s misidentification was based on Walckenaer (1905) and the species name was preoccupied by Olios argelasius (sub Sparassus). Simon (1932: 890) realized this confusion when he described and illustrated O. argelasius (Walckenaer) and proposed E. dufouri as a new replacement name (nomen novum) to substitute the previously established name E.

argelasius. Simon noted that ―the species described under name Sparassus argelasius Latreille (in Simon 1875: 334) must take the new name (nom. nov.) as Eusparassus dufouri‖.

Prior to proposing the genus Eusparassus, Simon (1875: 334, 1880: 290) used the nominal species ―Sparassus argelasius‖ to describe E. dufouri. In the literature, Sparassus itself was used to record not only different species but also different generic taxa including Micrommata, Olios and Eusparassus.

Neotype designation. According to all the facts noted, Simon (1932) did not designate any name-bearing type specimen subsequently while referring to Latreille (1818). According to Article 72.4.2 of ICZN when a new nominal species-group taxon (E. dufouri) is based on a published misidentification by an earlier author (M. argelasia Lat.), the type series consists of

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