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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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Alignment. The sequences were curated and aligned using ClustalW implemented in MEGA (v.5.2.1) (Tamura et al. 2012). 28S and 16S were further aligned with MAFFT v7 (Katoh and Standley 2013). Concatenation of the four different markers were done using Mesquite v2.75 (Maddison and Maddison 2011).

Phylogenetic analyses. To verify the edited sequences MEGA (v.5.2.1) (Tamura et al. 2012) was used by running Neighbor-Joining analysis under the Kimura 2Parameter model. Models of nucleotide substitution and sequence evolution were evaluated for each gene under different alignment strategies using jModeltest v0.1.1 (Posada 2008). The estimated models were further used to test phylogenetic relationships among taxa under Bayesian inference (BI) (using MrBayes v3.2.1 after Huelsenbeck and Ronquist 2001; Ronquist and Huelsenbeck 2003). Parallel to BI analyses, the phylogenetic relationships were tested using Maximum likelihood inference (using raxmlGUI v0.95 after Silvestro and Michalak 2011) The resulted trees were viewed and manipulated in Figtree v. 1.4. (available at http://tree.bio.ed.ac.uk/software/figtree/). The divergence time among taxa were estimated by BEAST v1.7.2 (Drummond et al. 2012) by calibrating BI trees using fossil and biogeographic data.

2.3. Distributional data processing

Eusparassus species locality data were obtained from collection labels and direct

sampling in the field using GPS. Internet sources such as Google Earth:

http://www.earth.google.com used to verify the accuracy of the localities. The online global gazetteers version 2.2 (http://www.fallingrain.com/world) was used to find correct name of places and also states and provinces of the country of distribution. It also provided geographical coordination for the species locations when the coordinates were not recorded by the samplers. All formats were converted to decimal degree to be mapped in DIVA-GIS version 7.4.0 (available at http://www.diva-gis.org/, Hijmans et al. 2005).

3. RESULTS The results of this study are presented in three chapters. Chapter 3.1 is about the taxonomic revision of the genus Eusparassus in Eurasia. In chapter 3.2 I present an overview on the systematics of Eusparassus along with the revision of Afro-Arabian species. And finally, in chapter 3.3 the molecular phylogenetic relationships of Eusparassus, Eusparassinae and Sparassidae are investigated.

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Chapter 3.1.

Taxonomic revision of the genus Eusparassus Simon, 1903 (Araneae: Sparassidae) in Eurasia This chapter is based on the following paper in a slightly modified version.

Status: published (28 September 2012) Type of publication: Research Article Journal: Journal of Natural History Citation: Moradmand, M. and Jäger, P., 2012. Taxonomic revision of the huntsman spider genus Eusparassus Simon, 1903 (Araneae: Sparassidae) in Eurasia. Journal of Natural History, 46 (39-40), 2439–2496.

Results: Chapter 3.1: Eusparassus in Eurasia

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The huntsman spider genus Eusparassus Simon, 1903 in Eurasia is revised to include 13 valid species. The type species is redescribed, and additional diagnostic characters are presented for the genus. Neotypes are designated for Eusparassus dufouri Simon, 1932 from Portugal, and Eusparassus walckenaeri (Audouin, 1826) from Egypt to establish their identity.

Consequently, E. kronebergi stat. nov. Denis, 1958 from Afghanistan and E. doriae stat. nov.

(Simon, 1874) from central Iran which were considered junior synonyms of E. walckenaeri are re-established as valid species. Lectotypes and paralectotypes are designated for: E.

kronebergi stat. nov., E. maynardi (Pocock, 1901) comb. nov. and E. pearsoni (Pocock,

1901) comb. nov. Two new synonymies are proposed: E. nanjiangensis (Hu & Fu, 1985) as junior synonym of E. potanini (Simon, 1895) from Xinjiang Uyghur autonomous region in China and E. doriae stat. nov. as senior synonym of E. fontanieri (Simon, 1880). Three new combinations are proposed: Eusparassus xerxes (Pocock, 1901) comb. nov. from Makran coast in Pakistan and Iran, E. maynardi (Pocock, 1901) comb. nov. from Baluchistan in Pakistan and E. pearsoni (Pocock, 1901) comb. nov. from Ghats in India (all transferred from the genus Olios Walckenaer, 1837). The latter two species are proposed as valid species and

are removed from junior synonymy with E. xerxes comb. nov. One new species is described:

E. mesopotamicus spec. nov. (male and female) from Iraq and Iran. New geographical records are presented: E. pontii Caporiacco, 1935 and E. kronebergi stat. nov. are recorded for the first time from India. E. fuscimanus Denis, 1958, E. oculatus (Kroneberg, 1846) and E.

levantinus Urones, 2006 are redescribed using new material. E. lilus Strand, 1907, described from Java, is proposed as nomen dubium because the type material could not be found and no longer seems to exist. Misplaced Olios flavovittatus comb. nov. (Caporiacco, 1935) from Karakoram is transferred from the genus Eusparassus. Almost all the species are redescribed for the first time and illustrations from male and female copulatory organs including intraspecific variations are provided using a large number of specimens.

Keywords: Eusparassinae, neotype, lectotype, new species, Eurasia

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Eusparassus Simon, 1903 are medium to large sized huntsman spiders which are among the foremost arthropod predators of deserts and semiarid areas (Levy 1989). Silken papery retreats, stuck firmly to underside of large flat stones, are used as a shelter for moulting and hiding during the day (Figure 1A). Females lay their eggs enclosed in a silken sac inside the retreat (Gerhardt 1928; Levy 1989; Gabriel 2011). They are distributed accross part of the Old World from Southern Africa to Mediterranean Europe and through the Middle East into Central and South Asia. The single Neotropical report of the genus, ―Eusparassus shefteli‖ Chamberlin, 1916 is not congeneric with Eusparassus (based on original illustrations and picture of holotype female epigyne) and actually belongs to the genus Polybetes Simon, 1897 (Cristina Anne Rheims, personal communication). Thus, Eusparassus spp. are restricted to Africa and Eurasia.

Currently, Eusparassus comprises 28 nominal species, of which 10 are described from Eurasia and 18 from Africa (Platnick 2012). The majority of the species are known merely by a single gender and by their original description, thus they have never been recorded since.

Some species were placed originally or subsequently in the genus Olios and therefore the necessity to investigate their types was unavoidable. The systematics of the taxon is obscure and no comprehensive taxonomic revision has been carried out so far. It was only Levy (1989) who in a small revisionary work, emphasized on the female‘s copulatory organs and the lateral view of the vulva as a diagnostic character, and redescribed E. walckenaeri (sub Sparassus). Presently, definitions of characters and species boundaries of Eusparassus species are incomplete. As in other groups of spiders, early diagnoses were based mostly on variable somatic and non-copulatory characters, which poorly defined species boundaries.

Eusparassus show a striking uniformity in somatic and copulatory characters. These similarities in traits as well as some intraspecific variations have challenged a discrimination of species. The challenging taxonomy of Eusparassus was admitted by previous workers as well (i.e., Denis 1947; Levy 1989; Jäger and Yin 2001).

Results: Chapter 3.1: Eusparassus in Eurasia

FIGURE 1. (A) Papery retreats of Eusparassus walckenaeri underside of flat stones in Mügla, Turkey, (B) Habitus of E.

walckenaeri at the entrance of its retreat, (C) Habitus of Eusparassus mesopotamicus spec. nov. from Birecik, Turkey. Photos by D. Kunz (A, B) and B. Göcmen (C).

Results: Chapter 3.1: Eusparassus in Eurasia

Eusparassus was erected by Simon (1903) to replace nominal species published under the name Sparassus by Simon in 1880. The type species is E. dufouri Simon, 1932, designated originally sub misidentified ―Eusparassus argelasius‖. Simon (1903: 1020) proposed the synonymy of Sparassus Walckenaer, 1805 with Micrommata Latreille, 1804.

The time lapse between proposing the genus and describing the type species was due to the taxonomic puzzle of the generic name Sparassus. At that time, it had been subjected to complex disputes. It was in use simultaneously for species of Olios Walckenaer, 1805 and Eusparassus (Simon 1874 and 1895; Pocock 1901). Some workers (Bonnet 1958; Levy 1989) originally used Sparassus for describing and recording species of Eusparassus until Sparassus was considered by Jäger (1999) as a junior synonym of Micrommata. Simon (1897) classified the Sparassidae by means of eyes (pattern and size) in seven sub groups and placed Sparassus in Sparassinae (sub ―Sparasseae‖). After creating the genus Eusparassus, Simon (1903) classified his new genus along with several other genera in Delenineae (sub ―Deleneae‖).

Järvi (1912, 1914) proposed a new subfamily Eusparassinae (sub ―Eusparasseae‖) for three genera Eusparassus, Pseudomicrommata Järvi, 1914 and Rhitymna Simon, 1897, according to similarities in female copulatory structures. Rhitymna was later revised and proved to belong to a different phylogenetic lineage (Jäger 2003). Järvi‘s classification appeared in Petrunkevitch (1928) who emphasized some somatic characters and combined Järvi‘s and Simon‘s classifications. Recently, Eusparassinae was re-established by Jäger and Kunz (2003) who found some synapomorphies in both somatic and genital characters, transferred an endemic African genus Arandisa Lawrence, 1938 and proposed some other endemic South African genera to be potentially included in this subfamily.

The recently investigated Eusparassus fossil in Baltic amber, E. crassipes (Koch and Berendt, 1854), uncovered the long existence of the genus which can be dated back to at least 44–49 Ma, during Eocene (Dunlop et al. 2011). Modern widespread distribution of the genus across the Old World and its long existence at least from early Tertiary till present (~50 million years) demonstrate its evolutionary success. The extant individuals occur in a diverse range of elevations from the semi-arid areas at sea level to the mountainous highlands, c. 4000 m, the highest altitude recorded to date for members of Sparassidae. In the present paper, we deal with the Eurasian species excluding Arabia. The Eusparassus fauna of the Arabian Peninsula is mostly related to Northern African elements which will be considered in a revision of African representatives. In this context, we provide descriptions of 13 Eusparassus species and designate neotypes, lectotypes and paralectotypes to fix species identities.

Results: Chapter 3.1: Eusparassus in Eurasia


The specimens were examined, measured and illustrated using a Leica MZ 165C stereomicroscope equipped with a drawing tube. Male palps were observed in 70% ethanol.

Hairs covering the bulb as well as the base of RTA were removed with forceps and fine needles. Hairs surrounding distalo-ventral margin of cymbium and around RTA are partially illustrated in male palps. Female epigynes were dissected and soft tissue surrounding vulva was removed using minute entomological pins (model Sphinx V2A 0.1 x 12 mm).

Subsequently, epigynes were submerged in 96% lactic acid to observe the internal duct system. For a better understanding of the internal duct system, we provide a drawing from anterio-dorso-lateral view of left vulvas. In this view, there is no need to cut the lateral lobes of epigyne and is suitable for type material which should be treated with care. The dorsal view of vulva is not illustrated (except Figure 3B). The order of species is arranged from West (Portugal) to East (India) in the geographical distribution range. Geographic coordinates extracted subsequently from the web site http://www.fallingrain.com/world/ are given in parentheses.

All measurements are in millimetres. Size classes of spiders are according to Jäger (2001: 14). Measurements of palps are listed as: total length [femur, patella, tibia, cymbium];

legs as: total length [femur, patella, tibia, metatarsus, tarsus]. Abbreviations used throughout the text: AB — anterior bands of epigynal field, ALE — anterior lateral eyes, AME — anterior median eyes, AMLL — anterior margin of LL, dRTA — dorsal RTA, EF — epigynal field, EFB — epigynal field bridge, LL — lateral lobes, MS — median septum, PLE — posterior lateral eyes, PME — posterior median eyes, PMLL — posterior margin of LL, RTA — retrolateral tibial apophysis, vRTA — ventral RTA, SD — Sparassidae DNA numbers in SMF, SS — slit sensillum, T — tegulum, TL — turning loop, I–IV — 1st to 4th leg. Palp and leg spination are presented in the following format: prolateral, dorsal, retrolateral and ventral (the latter only if present). Parentheses and slashes are used to state spination variation within a single specimen and among different specimens, respectively. Since most of the specimens are old and long preserved, we provide a general pattern of colouration in the genus description paragraph. Species specific colouration is given briefly in species description.

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