«Taxonomic Revision, Molecular Phylogeny and Zoogeography of the huntsman spider genus Eusparassus (Araneae: Sparassidae) Dissertation for attaining ...»
Taxonomic Revision, Molecular Phylogeny and
Zoogeography of the huntsman spider genus
Eusparassus (Araneae: Sparassidae)
for attaining the PhD degree of Natural Sciences
Submitted to the Department 15, Fachbereich Biowissenschaften
der Johann Wolfgang Goethe-Universität
in Frankfurt am Main
from Shahin-Shahr, Isfahan, Iran
Frankfurt am Main 2013
vom Fachbereich 15 Biowissenschanften
der Johann Wolfgang Goethe - Universität
als Dissertation angenommen.
Dekanin: Prof. Dr. Anna Starzinski-Powitz
Prof. Dr. Michael Türkay Prof. Dr. Sven Klimpel Date of Disputation:.−..−.
III IV I would like to dedicate my dissertation to my family for all their motivations, loves and supports, in particular to my parents, my wife Maryam and my son Bardia V VI Table of Contents
1.1 Motivation and preface to this study
1.2 General introduction to spider classification
1.3 Family Sparassidae
1.4 Genus Eusparassus and subfamily Eusparassinae, a historical review
1.5 Aims of my dissertation
2. Material and methods
2.1 Morphological studies
2.2 Molecular studies
2.3 Distributional data processing
3.1 Taxonomic revision of Eusparassus in Eurasia
3.2 Systematics and zoogeography of Eusparassus with revision of the genus in Afro-Arabia.........84
3.3 Molecular phylogeny of Sparassidae with focus on Eusparassus and Eusparassinae...............230
4.1 Taxonomic revision and systematics of Eusparassus based on morphology
4.2 Molecular phylogenetics
4.2.1 Eusparassus and its species-groups
4.2.2 subfamily Eusparassinae
4.2.3 Sparassidae and the rest of subfamilies
4.2.4 ‘Laterigradae’, Dionycha and the RTA-clade
4.3 Origination and zoogeography of Eusparassus
5. Zusammenfassung (Summary in German language)
AbstractThe spider genus Eusparassus Simon, 1903 (Araneae: Sparassidae: Eusparassinae;
stone huntsman spider) is revised worldwide to include 30 valid species distributed exclusively in Africa and Eurasia. The type species E. dufouri Simon, 1932 is redescribed and a neotype is designated from Portugal. An extended diagnosis for the genus is presented. Eight new species are described: Eusparassus arabicus Moradmand, 2013 (male, female) from Arabian Peninsula, E. educatus Moradmand, 2013 (male, female) from Namibia, E. reverentia Moradmand, 2013 (male, female) from Burkina Faso and Nigeria, E. jaegeri Moradmand, 2013 (male, female) from South Africa and Botswana, E. jocquei Moradmand, 2013 (male, female) from Zimbabwe, E. borakalalo Moradmand, 2013 (female) from South Africa, E.
schoemanae Moradmand, 2013 (male, female) from South Africa and Namibia and E. mesopotamicus Moradmand and Jäger, 2012 (male and female) from Iraq, Iran and Turkey. 22 species are re-described six of them are transferred from the genus Olios Walckenaer, 1837. Six species-groups are proposed: the dufouri-group [8 species: E. dufouri, E. levantinus Urones, 2006, E. barbarus (Lucas, 1846), E.
atlanticus Simon, 1909, E. syrticus Simon, 1909, E. oraniensis (Lucas, 1846), E.
letourneuxi (Simon, 1874), E. fritschi (Koch, 1873); Iberian Peninsula to parts of north-western Africa], walckenaeri-group [3 species: E. walckenaeri (Audouin, 1826), E. laevatus (Simon, 1897), E. arabicus; eastern Mediterranean to Arabia and parts of north-eastern Africa], doriae-group [7 species: E. doriae (Simon, 1874), E. kronebergi Denis, 1958, E. maynardi (Pocock, 1901), E. potanini (Simon, 1895), E. fuscimanus Denis, 1958, E. oculatus (Kroneberg, 1846) and E. mesopotamicus; Middle East to Central and South Asia], vestigator-group (3 species: E. vestigator (Simon, 1897), E.
reverentia, E. pearsoni (Pocock, 1901); central to eastern Africa and an isolated area in NW India], jaegeri-group [4 species: E. jaegeri, E. jocquei, E. borakalalo, E.
schoemanae; southern and south-eastern Africa], tuckeri-group [2 species: E. tuckeri (Lawrence, 1927), E. educatus; south-western Africa). Two species, E. pontii Caporiacco, 1935 and E. xerxes (Pocock, 1901) cannot be placed in any of the above groups. Two species are transferred from Eusparassus to Olios: O.
flavovittatus (Caporiacco, 1935) and O. quesitio Moradmand, 2013. 14 species are recognized as misplaced in Eusparassus, thus nearly half of the described species prior to this revision were placed mistakenly in this genus. Neotypes are designated for E. walckenaeri from Egypt, E. barbarus, E. oraniensis and E. letourneuxi (all three from Algeria) to establish their identity. The male and female of Cercetius perezi Simon, 1902, which was known only from the immature holotype, are described for the first time. It is recognized that the monotypic and little used generic name Cercetius Simon, 1902 — a species, which had been known only from the immature holotype — as a synonym of the widely used name Eusparassus. The case proposal 3596 (conservation of name Eusparassus) is under consideration by ICZN.
The first comprehensive molecular phylogeny of the family Sparassidae with focus on the genus Eusparassus is investigated using four molecular markers (mitochondrial COI and 16S; nuclear H3 and 28S). The monophyly of Eusparassus and the dufouri, walckenaeri and doriae species-groups are recovered with the latter two groups more closely related. The monophyly of the tuckeri-group is not supported and the position of E. jaegeri as the only available member of the jaegeri-group is not resolved within the Eusparassus clade. DNA samples of the vestigator-group were not accessible for this study. The origination of the genus Eusparassus around 70 million years ago (MA) is estimated according to molecular clock analyses. Using this recent result in combination with some biogeographic and geological data, the Namib Desert is proposed as the place of ancestral origin for Eusparassus and putative Eusparassinae genera.
Further analyses are done on the phylogenetic relationships of Sparassidae and its subfamilies. The Eusparassinae are not confirmed as monophyletic, with the two original genera Eusparassus and Pseudomicrommata in separate clades and only the latter clusters with most other assumed Eusparassinae, here termed the ―African clade‖. Monophyly of the subfamilies Sparianthinae, Heteropodinae sensu stricto, Palystinae and Deleninae is recovered. The Sparianthinae are supported as the most basal clade, diverging considerably early (143 MA) from all other Sparassidae. The Sparassinae and genus Olios are found to be polyphyletic. The Sparassidae are confirmed as monophyletic and as most basal group within the RTAclade. The divergence time of Sparassidae from the RTA-clade is estimated with 186 MA in the Jurassic. No affiliation of Sparassidae to other members of the ‗Laterigradae‘ (Philodromidae, Selenopidae and Thomisidae) is observed, thus the crab-like posture of this group was proposed a result of convergent evolution. Only the families Philodromidae and Selenopidae are found members of a supported clade. Including a considerable amount of RTA-clade representatives, the higherlevel clade Dionycha is not but monophyly of the RTA-clade itself is supported.
1. INTRODUCTIONIt is estimated that nearly 8.7 million eukaryotic species are living on earth, of those 86% still await to be identified by taxonomists (Mora et al. 2011) and for spiders, the order Araneae, over 2/3 of their diversity is claimed to await discovery (Agnarsson et al. 2013). The accelerating biodiversity loss on earth provides an urgent need to increase the knowledge of the survived species diversity.
1.1. Motivation and preface to this study
It was in 2006 when I visited the Senckenberg Research Institute and Natural History Museum (SMF) during a short time scientific visit targeting taxonomy of the hermit crabs from the Persian Gulf (my MSc thesis at the University of Tehran), via ―the Middle Eastern biodiversity network project‖ funded by the DAAD. Three years later (Sep 2009), I have awarded a PhD scholarship from the Ministry of Science, Research and Technology of Iran to study the taxonomy and phylogeny of spiders in SMF as a student of the Goethe University, Frankfurt am Main. There were three main reasons for this shift (from crabs to spiders). First was my great interest in Arachnida. Secondly I found studying taxonomy of spiders a research gap area not even in Iran but also in the entire Middle Eastern countries where the spider fauna is so diverse and unique but at the same time not well investigated. Third reason points toward SMF itself that houses one of the richest spider collections in the world and employed a renowned spider taxonomist such as Dr Peter Jäger. I have had a great opportunity to learn about taxonomy, nomenclature and systematics as well as phylogeny, biogeography and evolutionary history of spiders in general and Sparassidae in detail.
The huntsman spiders or giant crab spiders of the family Sparassidae Bertkau, 1872 are among the most diverse families of spiders. Despite they comprises the largest spiders in the world, most of the genera are not investigated. For instance, it is surprising that of the 33 genera known from Africa just two have been revised prior to this study. The genus Eusparassus Simon, 1903 is one of these unrevised genera.
Eusparassus was previously included 29 nominal species, a heterogeneous assemblage of taxa. They are distributed from Africa to parts of Eurasia. Moreover, Eusparassus is unique among the sparassid genera by having a well preserved fossil dated back to the Eocene era (Dunlop et al. 2011). All these facts highlight the genus as a fruitful subject for a taxonomic, phylogenetic and biogeographic study.
I have started my research on spiders by revising the cave dwelling genus Spariolenus Simon, 1880 in Iran leading to the description of four new species (Moradmand and Jäger 2011). Then I focused on the taxonomic revision of Eusparassus which resulted in two revisionary publications (Moradmand and Jäger 2012a; in Eurasia) and (Moradmand 2013; in Africa and Arabia) and one case proposal to the International Commission on Zoological Nomenclature (ICZN) (Moradmand and Jäger 2012b). I presented three lectures and one poster of my
results in International (2010, Poland: Siedlce) and European (2012, Slovenia:
Ljubljana) congresses and also two talks as scientific visitor in 2011 at the Natural History Museum (NHM London) and in 2012 at the Royal Museum for Central Africa (MRAC Tervuren). These two latter scientific visits were sponsored by SYNTHESYS for studying the type series at noted spider collections. The most time consuming task during the revision was to find and borrow the (sometimes hidden) type material from various collections in Europe and Africa. Additionally to NHM and MRAC, I had an opportunity to visit the rich spider collections of the Muséum National d‘Histoire Naturelle (MNHN Paris), Zoological Museum, University of Copenhagen (ZMUC) and Museum für Naturkunde (ZMB Berlin). I have learnt new scientific methods and techniques by participating in the ―Phylogenetic systematics and molecular dating course‖ in ZMUC (2011), the ―Geographic Information System‖ (GIS) workshop and the ―Species Distribution Modelling‖ (SDM) course which two latter were held in GRADE (Goethe Graduate Academy, Frankfurt).
After performing the taxonomic review, I classified Eusparassus species into species-groups and tried to raise evolutionary hypotheses about speciation, relationships of species-groups, historical biogeography and the systematic position of the genus Eusparassus within the Sparassidae (see chapter 3.2). To verify these hypotheses I focused on molecular phylogeny and molecular clock analyses (see chapter 3.3). Additionally to Eusparassus, the range of the sampled taxa expanded gradually while my study was the first comprehensive phylogenetic analysis on Sparassidae as well. Thus, representatives of other Eusparassinae genera in particular and Sparassidae in general were included in my research as well as members of the RTA-clade (see chapter 3.3).
1.2. General introduction to spider classification
Spiders, Order Araneae, are one of the most diverse groups of animals belonging to the class Arachnida. They are chelicerate arthropods and share the trait chelicerae with other well-known orders, i.e. Scorpiones, Opiliones and Acari (Foelix 2010).
Currently, taxonomists classified more than 44,000 known spider species into 112 families (Platnick 2013). The Araneae are divided into three suborders: Mesothelae (the oldest group with autplesiomorphic character, e.g. segmented opisthosoma), Mygalomorphae (Tarantulas and their relatives) and Araneomorphae. The Araneomorphae exhibit a vertical form of fangs (labidognath) in comparison to Mygalomorphae which have parallel fangs (orthognath).
The Araneomorphae include the majority (90%) of spider species (Foelix 2010). Some of them have a special spinning plate close to spinnerets named cribellum. Araneomorphae with and without cribellum are called cribellate and ecribellate, respectively. According to the complexity of the copulatory structures, Eugene Simon in his ―Historie Naturelle des Araignées‖ (1892–1903), separated the ecribellate spiders into haplogynes (simple copulatory organs) and entelegynes (complex copulatory organs). Entelegynes are usually divided into Trionycha (legs with three tarsal claws) and Dionycha (legs with two tarsal claws). Another higher level synapomorphy among a relatively large group of entelegynes is the retrolateral tibial apophysis on the male palpal tibia (RTA). The group with representatives sharing this character is called RTA-clade (Coddington and Levi 1999).