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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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Walckenaer CA. 1830. Aranéides. In Faune française ou histoire naturelle générale et particulière des animaux qui se trouvent en France, constamment ou passagèrement, à la surface du sol, dans les eaux qui le baignent et dans le littoral des mers qui le bornent par Viellot, Desmarrey, Ducrotoy, Audinet, Lepelletier et Walckenaer. Paris, livr. 26: 97–175, livr. 29: 177–240.

Walckenaer CA. 1837. Histoire naturelle des insectes. Aptères. Paris, 1:1–682.

Results: Chapter 3.1: Eusparassus in Eurasia Results: chapter 3.2: Systematics and zoogeography with revision of Afro-Arabian species Chapter 3.2. Systematics and Zoogeography of Eusparassus with revision of the African and Arabian species This chapter is based on the following paper in a slightly modified version.

Status: published (17 June 2013) Type of publication: Monograph Journal: Zootaxa Citation: Moradmand, M., 2013. The stone huntsman spider genus Eusparassus (Araneae: Sparassidae): systematics and zoogeography with revision of the African and Arabian species. Zootaxa, 3675 (1), 1-108.

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

Abstract

An overview on the systematics of the stone huntsman spider genus Eusparassus Simon, 1903 and an identification key to the known species are presented. Six species-groups are proposed.

The walckenaeri group (3 species, Eastern Mediterranean to Arabia and parts of North-Eastern Africa), dufouri group (8 species, Iberian Peninsula to parts of North-western Africa), vestigator group (3 species, Central to Eastern Africa and an isolated area in India), jaegeri group (4 species, Southern and South-Eastern Africa), tuckeri group (2 species, South-Western Africa) and doriae group (7 species, Middle East to Central and South Asia). Two species, E. pontii Caporiacco, 1935 and E. xerxes (Pocock, 1901) could not be placed in any of the above groups.

The species from Africa and Arabia are revised. The following ten species are re-described:

Eusparassus barbarus (Lucas, 1846), E. atlanticus Simon, 1909 stat. nov., E. syrticus Simon, 1909, E. oraniensis (Lucas, 1846), E. letourneuxi (Simon, 1874), E. fritschi (Koch, 1873) stat.

rev., E. walckenaeri (Audouin, 1826), E. vestigator (Simon, 1897) comb. nov., E. laevatus (Simon, 1897) comb. nov. and E. tuckeri (Lawrence, 1927) comb. nov. The latter three species are transferred from Olios Walckenaer, 1837. Seven new species are described: Eusparassus arabicus spec. nov. (male, female) from Arabian Peninsula, E. educatus spec. nov. (male, female) from Namibia, E. reverentia spec. nov. (male, female) from Burkina Faso and Nigeria, E. jaegeri spec. nov. (male, female) from South Africa and Botswana, E. jocquei spec. nov.

(male, female) from Zimbabwe, E. borakalalo spec. nov. (female) from South Africa and E.

schoemanae spec. nov. (male, female) from South Africa and Namibia. Three taxa, E. dufouri maximus Strand, 1906 syn. nov., E. rufobrunneus Caporiacco, 1941 syn. nov. and Olios furcatus Lawrence, 1927 syn. nov. are proposed as junior synonyms of E. oraniensis, E. vestigator comb.

nov. and E. tuckeri comb. nov. respectively. Males of E. atlanticus stat. nov. and E. fritschi stat.

rev. are described for the first time as in the female of E. vestigator comb. nov. Neotypes are designated for E. barbarus, E. oraniensis and E. letourneuxi (all from Algeria). The male and female of Cercetius perezi Simon, 1902, which was known only from the immature holotype, are described here for the first time. This resulted in recognizing the monotypic and little used generic name Cercetius Simon, 1902 as a synonym of the widely used name Eusparassus. Nearly all the species are illustrated for the first time. Eusparassus concolor Caporiacco, 1939 is transferred to Olios Walckenaer, 1837 and the replacement name Olios quesitio is proposed because of secondary homonymy. For the majority of the species, new geographical records are presented. The systematics and zoogeography of the currently known species and species groups are discussed. A brief note on the copulation process of E. walckenaeri is presented.

Key words: taxonomy, Eusparassinae, Cercetius, identification key, copulation process, evolutionary hypothesis.

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

INTRODUCTION

Members of the spider genus Eusparassus Simon, 1903 (Sparassidae: Eusparassinae) are among the most conspicuous arachnid predators in arid and semiarid deserts of Africa and most parts of Eurasia. As these spiders inhabit stony habitats and build their retreats underside of large flat stones and also in the crevices of rocks (Levy 1989, Gabriel 2011), the common name ―stone huntsman spiders‖ is proposed here. They are small to very large huntsman spiders distributed in Africa and Eurasia. The fossil stone huntsman spider, E. crassipes (Koch & Berendt, 1854) from Eocene era found in Northern Europe amber fossil, is dated back to approximately 50 Ma (Dunlop et al. 2011). Recently, Moradmand and Jäger (2012a) revised the Eurasian representatives (excluding Arabia) and provided an historical review of the systematics of the genus. They provided diagnostic characters of the genus Eusparassus and recognized 13 valid species in Europe, the Middle East, and Central and South Asia. Before that, the genus had never been revised with the exception of a brief review by Levy (1989) who re-described E.





walckenaeri (sub Sparassus) and mentioned some diagnostic characters (e.g. female vulva and colouration of the ventral opisthosoma) for species identification, along with a revision of some Middle Eastern Sparassidae.

The systematic position of Eusparassus within Sparassidae remains vague, since the majority of Sparassidae genera have not yet been revised. Simon (1897a) placed Eusparassus (sub Sparassus) in his proposed ―Sparasseae‖ group. Later, Simon (1903) moved the genus to another group named ―Deleneae‖ along with several other genera. Simon’s classifications were based on somatic characters, e.g. the arrangement of eyes. Järvi (1912, 1914) was the first who applied characters of the copulatory organs (exclusively female) to classify Sparassidae. He proposed the subfamily Eusparassinae Järvi, 1912 (sub ―Eusparaseae‖) for Eusparassus including the genera: Pseudomicrommata Järvi, 1914 and Rhitymna Simon, 1897. Of these two genera, only the African endemic Pseudomicrommata, known as the grass huntsman spider, has some kind of similarities to Eusparassus. Jäger (2003) proposed that Rhitymna represent a different phylogenetic lineage in Asia. Jäger and Kunz (2003) in a congress abstract proposed some diagnostic characters for Eusparassinae and assumed that a number of African endemic genera could be placed in this subfamily (e.g. Arandisa Lawrence, 1938).

Huntsman spiders in Africa and Arabia have received little taxonomic attention (Jäger & Kunz 2005). Despite having great diversity and living in various habitats, the majority of the African huntsman spiders remained unexplored compared to their relatives in other parts of the Results: chapter 3.2: Systematics and zoogeography with revision of Afro-Arabian species world. Dippenaar-Schoeman and Jocqué (1997) gave an historical review of systematic research into the huntsman spiders of Africa (sub Heteropodidae). Jäger and Kunz (2005) provided an overview of the known genera of the huntsman spiders and presented a generic identification key for Africa and nearby regions. They listed 33 nominal genera, and of these, only two have been revised to date: the Afrotropical genus Palystes L. Koch, 1875 (by Croeser 1996) and the AfroAsian genus Cebrennus Simon, 1880 (by Jäger 2000). The nominal and monotypic genus Cercetius Simon, 1902 from Afro-Arabia is considered to be a senior synonym of Eusparassus (Moradmand & Jäger, 2012b). An official proposal (case number 3596) was submitted to the International Commission on Zoological Nomenclature (ICZN) to conserve the name Eusparassus by giving it precedence over the forgotten name Cercetius (Jäger & Moradmand 2012b). The case is under consideration by ICZN, and until a ruling has been made, prevailing usage of the names is to be maintained (ICZN 1999: Article 82). Palaeogeographically closely related to Africa, the Arabian Peninsula is totally neglected in the case of studying sparassids except for a few sparsely recorded species (e.g. Simon 1902; Levy 1989; Jäger 2000; Jäger 2006).

In this paper, an overview of all the known Eusparassus species is provided and the AfroArabian representatives are revised investigating all the available types and a large number of specimens from the major European and African spider collections. Although 18 nominal species are known from Afro-Arabia (Platnick 2013), the taxonomic status of the majority of these species is not clear. However, following the present study and that of Moradmand and Jäger (2012a), 30 valid species of Eusparassus are identified, of which 25 species are known by both sexes and 5 species by females alone. An identification key to all the known species is presented here.

MATERIAL AND METHODS

Material for this study was mostly obtained from the large spider collections in Europe and Africa (listed below). Additional specimens were sampled by colleagues from different areas within the Eusparassus distribution range. The spiders have a cryptic life style in subterranean habitats including crevices in rocks and under large flat stones, where they construct their large papery retreats (Fig. 52c). They are strikingly agile and tricky to be caught. The typical method for sampling Sparassidae (using a headlight to trace reflecting eyes at night) seems to be non effective in these spiders. They are mainly collected inside their retreats by turning over the Results: chapter 3.2: Systematics and zoogeography with revision of Afro-Arabian species inhabited stones by day and removing the spider from their retreats. The use of pitfall traps (R.

Bosmans, in Algeria) and Malaise traps (A. van Harten, in Yemen) are two additional methods that seem to be effective for collecting stone huntsman spiders, especially males (see data on labels).

Examination, illustration and measurements of the specimens follows Moradmand and Jäger (2012a), using a Leica MZ 165C stereomicroscope equipped with a drawing tube. Photos of the copulatory structures and habitus were taken using a Canon EOS 50D camera installed on the microscope. The arrangement of the species in the text is according to species groups and follows the geographical distribution range from North (Sahara) to South (Southern Africa).

Measurements are given in millimetres. Size classes of spiders are according to Jäger (2001) as follows: small (3–10 mm), medium (11–20 mm), large (21–30 mm) and very large (30 mm).

Measurements of palps are listed as: total length [femur, patella, tibia, cymbium]; legs as: total length [femur, patella, tibia, metatarsus, tarsus]. 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. At the beginning of every description part, the given MM number (serial number used by the author to measure and/or illustrate the material) is noted for the single measurement of a particular specimen (i.e., eye sizes), but for the range measurements (i.e., total length) the sum number of studied specimens are noted.

Abbreviations used throughout the text:

AB — anterior bands of epigynal field, ALE — anterior lateral eyes, AME — anterior median eyes, AMLL — anterior margin of lateral lobes, CD — copulatory duct, CO — copulatory opening, DK—field numbers used by Dirk Kunz, dRTA — dorsal RTA, E — embolus, EF — epigynal field, EFB — epigynal field bridge, EM — embolus membrane, Results: chapter 3.2: Systematics and zoogeography with revision of Afro-Arabian species ET — embolus tip, GP — glandular process, Gpo — glandular pores, H — haematodocha, L — lumen, LL — lateral lobes, MM — serial numbers used by the author for measured and/or illustrated material, MS — median septum, Msa — membraneous sac, PLE — posterior lateral eyes, PME — posterior median eyes, PMLL — posterior margin of LL, RTA — retrolateral tibial apophysis, vRTA — ventral RTA, ST — subtegulum, SD — Sparassidae DNA numbers in SMF, SpD — sperm duct, SS — slit sensillum, T — tegulum, TL — turning loop, Vsh — vulva shadow, I–IV — 1st to 4th leg.

Collections and curators AMNH— American Museum of Natural History, New York (Norman Platnick) BMSA (NMBA) — National Museum, Bloemfontein (Leon N. Lotz, Trudie Peyper) CCM — Collection of Christoph Muster, Putbus CRB — Collection of Robert Bosmans, Gent HECO — Hope Entomological Collection, Oxford (Zoë Simmons) ICEAD — Invertebrate Collection of Environment Agency, Abu Dhabi (Anitha K. Saji) MHNG —Muséum d’histoire naturelle, Genève (Peter Schwendinger) MIZ — Zoological Museum, Polish Academy of Science, Warsaw (Dominika Mierzwa) MNHN — Muséum National d’Histoire Naturelle, Paris (Christine Rollard) Results: chapter 3.2: Systematics and zoogeography with revision of Afro-Arabian species MNM — Museo Civico di Storia Naturale di Milano, Milan (Andrea Sabbadini, Carlo Pesarini) MRAC — Musée Royal de l’Afrique Centrale, Tervuren (Rudy Jocqué) MZH — Finish Museum of Natural History, University of Helsinki (Ritva Talman) MZUF —Natural History Museum ―La Specola‖, Florence (Luca Bartolozzi) NHM —Natural History Museum, London (Janet Beccaloni) NHMW — Naturhistorisches Museum, Vienna (Christoph Hörweg)

NMB — Naturhistorisches Museum, Basel (Ambros Hänggi)



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