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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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Later on, around 35 MA, the walckenaeri-group split from the doriae-group with their common ancestor probably lived somewhere in the Middle East and/or NE Africa. These two groups recovered to be closely related phylogenetically. The members of the doriae-group (seven species) are distributed in the Middle East toward Central Asia and parts of South Asia. The walckenaeri-group occurs in eastern Mediterranean and in NE Africa and Arabia. It is likely that the doriae lineage diverged from the common ancestor with the walckenaeri-group by losing their intermarginal denticles of the chelicerae and subsequent modifications in the copulatory structures. The closing age of the Turgai Strait is in concordant with the divergent time of the doriae- and walckenaeri-group. This event might play a role in the divergence of these two groups. Later on in Miocene, a land bridge of arid savannah between Arabia and southern Iran (Shmida, 1985) around where the Persian Gulf is currently present might have affected further exchanges.

The phylogenetic position and relationships of the vestigator-group members could not be tested in this study due to missing fresh samples for DNA isolation. The vestigator-group composes of three species, two occur in Central and East Africa and one endemic to western India. The presence of this isolated lineage (E. pearsoni) in India far from Africa can be explained by secondary distribution of the hypothesised ancestor from eastern Africa to India after the reattachment of the two plates via great Somalia around 65–60 MA.

Finally for the same reason as for the previous, the phylogenetic relationships of E. xerxes (from Arabia to Pakistan) and E. pontii (Himalayas) to rest of congeners could not be tested. Both species could not be placed in the proposed species groups. They demonstrate a transition in characters between the dufouri-, doriaeand walckenaeri-group. E. pontii might have diverged from the doriae-group in highlands of the Himalayas and adapted to very high elevations (~4000 m). These entire hypotheses must be tested by a robust phylogeny and molecular dating approach after a rich sampling of the noted taxa.

5. Zusammenfassung

Die Spinnengattung Eusparassus Simon, 1903 (Araneae: Sparassidae:

Eusparassinae; Stein-Riesenkrabbenspinnen) wird weltweit revidiert, sie umfasst 30 valide Arten ausschließlich aus Afrika und Eurasien. Die Typus-Art E. dufouri Simon, 1932 wird wiederbeschrieben und ein Neotypus aus Portugal designiert. Eine erweiterte Diagnose für die Gattung wird vorgeschlagen. Acht neue Arten werden erstmals beschrieben: Eusparassus arabicus Moradmand, 2013 (Männchen, Weibchen) von der Arabischen Halbinsel, E. educatus Moradmand, 2013 (Männchen, Weibchen) aus Namibia, E. reverentia Moradmand, 2013 (Männchen, Weibchen) aus Burkina Faso und Nigeria, E. jaegeri Moradmand, 2013 (Männchen, Weibchen) aus Südafrika und Botswana, E. jocquei Moradmand, 2013 (Männchen, Weibchen) aus Simbabwe, E. borakalalo Moradmand, 2013 (Weibchen) aus Südafrika, E. schoemanae Moradmand, 2013 (Männchen, Weibchen) aus Südafrika und Namibia sowie E. mesopotamicus Moradmand & Jäger, 2012 (Männchen und Weibchen) aus dem Irak, Iran und der Türkei. 22 Arten werden wiederbeschrieben, wovon sechs aus der Gattung Olios Walckenaer, 1837 nach Eusparassus transferiert werden. Sechs Artengruppen werden aufgestellt: die dufouri-Gruppe [8 Arten: 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); Iberische Halbinsel bis Nordwestafrika (teilweise)], walckenaeri-Gruppe [3 Arten: E. walckenaeri (Audouin, 1826), E. laevatus (Simon, 1897), E. arabicus; östlicher Mittelmeerraum bis Arabien und Nordostafrika (teilweise)], doriae-Gruppe [7 Arten: E. doriae (Simon, 1874), E. kronebergi Denis, 1958, E. maynardi (Pocock, 1901), E. potanini (Simon, 1895), E. fuscimanus Denis, 1958, E. oculatus (Kroneberg, 1846) und E. mesopotamicus; Mittlerer Osten bis Zentral- und Südasien], vestigator-Gruppe (3 Arten: E. vestigator (Simon, 1897), E.

reverentia, E. pearsoni (Pocock, 1901); Zentral- bis Ostafrika und eine isolierte Region in NW Indien], jaegeri-Gruppe [4 Arten: E. jaegeri, E. jocquei, E. borakalalo, E. schoemanae; Süd- und Südostafrika], tuckeri-Gruppe [2 Arten: E. tuckeri (Lawrence, 1927), E. educatus; Südwestafrika). Zwei Arten, E. pontii Caporiacco, 1935 und E. xerxes (Pocock, 1901) können keiner der obengenannten Gruppen zugeordnet werden. Zwei Arten werden von Eusparassus nach Olios transferiert: O.

flavovittatus (Caporiacco, 1935) und O. quesitio Moradmand, 2013. 14 Arten wurden von früheren Autoren fälschlicherweise in die Gattung Eusparassus gestellt, damit wurden vor dieser Revision fast die Hälfte der beschriebenen Arten irrtümlicherweise in die falsche Gattung gestellt. Neotypen werden designiert für E. walckenaeri aus Ägypten, E. barbarus, E. oraniensis und E. letourneuxi (alle drei letztgenannten aus Algerien), um die Identität dieser Arten eindeutig festzulegen. Männchen und Weibchen von Cercetius perezi Simon, 1902 —eine Art, die nur vom juvenilen Holotypus bekannt war— werden erstmals beschrieben. Hierbei wurde festgestellt, dass der kaum benutzte, monotypische Gattungsname Cercetius Simon, 1902 ein älteres Synonym des weitverbreiteten und sehr häufig benutzten Gattungsnamens Eusparassus ist. Ein Antrag (Nummer 3596), den Namen Eusparassus zu schützen, ist bei der Internationalen Kommission für Zoologische Nomenklatur (ICZN) anhängig.





Die Phylogenie der Sparassidae mit dem Schwerpunkt Eusparassus wird zum ersten Mal anhand von vier molekularen Markern (mitochondriale Gene COI und 16S; nukleare Gene H3 und 28S) untersucht. Die Monophylie von Eusparassus und der dufouri-, walckenaeri- und doriae- Arten-Gruppen wird nachgewiesen, wobei die beiden letztgenannten Gruppen enger miteinander verwandt sind. Die tuckeri-Gruppe ist kein Monophylum, die Position von E. jaegeri (als der einzig verfügbaren Art aus der jaegeri-Gruppe) innerhalb des Eusparassus-Stammbaumes kann nicht aufgelöst werden. DNA-Proben aus der vestigator-Gruppe standen für diese Studie nicht zur Verfügung. Anhand der „Molekularen Uhr―-Analyse wird der Ursprung der Gattung Eusparassus vor ca. 70 Millionen Jahren vermutet. Anhand der vorliegenden Ergebnisse in Kombination mit biogeographischen und geologischen Daten sind Eusparassus und andere mutmaßliche Gattungen der Eusparassinae wahrscheinlich in der Namib-Wüste entstanden.

Weitere Analysen zu den phylogenetischen Beziehungen der Sparassidae und deren Unterfamilien weisen die Eusparassinae als polyphyletisch aus, mit den beiden Gattungen Eusparassus und Pseudomicrommata in getrennten Linien. Nur die letztere Gattung gruppiert mit den meisten anderen vermuteten Eusparassinae, hier ―Afrikanische Gruppe‖ genannt. Die Unterfamilien Sparianthinae, Heteropodinae sensu stricto, Palystinae und Deleninae sind monophyletisch. Die Sparianthinae sind ein basaler Abzweig, welcher sich relativ früh (vor ca. 143 Millionen Jahren) von allen anderen Sparassidae abgetrennt hat. Die Sparassinae und die Gattung Olios sind polyphyletisch. Die Sparassidae ist eine monophyletisch Gruppe und innerhalb des sogenannten „RTA-clades― eine basale Abzweigung. Die Abspaltung der Sparassidae vom „RTA-clade― wird vor ca. 186 Millionen Jahre angenommen, also innerhalb des Jura. Es werden keine näheren Beziehungen zu anderen Mitgliedern der ‚Laterigradae‗ (Philodromidae, Selenopidae und Thomisidae) beobachtet, die krabbenartige Stellung der Beine dieser Gruppe wird als ein Ergebnis konvergenter Evolution angesehen. Nur die Familien Philodromidae und Selenopidae werden als Mitglieder ein- und derselben, unterstützten Untergruppe erkannt. Unter Einbeziehung eines beträchtlichen Anteils von Vertretern des „RTA-clades― werden die übergeordneten „Dionycha― nicht als monophyletisch, der „RTA-clade― selbst aber als monophyletisch unterstützt.

6. ACKNOLEDGMENTS I am very grateful to Dr Peter Jäger (SMF) for all his supports and encouragements as my arachnological advisor. I am thankful to Prof Dr Michael Türkay (SMF) for acting as my main supervisor and for all his supports during my doctoral research. I would like to thank Prof Dr Sven Klimpel (Frankfurt University; Biodiversity and Climate Research Centre (BIK-F)) for acting as the second referee of my dissertation.

Prof Dr Jochen Martens (University of Mainz) and Prof Dr Georg Zizka (Frankfurt University; SMF) are gratefully acknowledged as referees for my disputation.

I am very grateful to Dr Axel Schönhofer (University of Mainz) for all his invaluable recommendations, assistances and cooperation on the phylogenetic part of my study. The staff of the Arachnology section (SMF): Mrs Julia Altmann, Dr Steffen Bayer and Mrs Elena Grall are gratefully acknowledged with my especial thanks to J. Altmann for her kind assistances during my lab works. I am very thankful to Dr Mark Harvey (Western Australian Museum, Perth), Dr Charles Griswold (California Academy of Sciences, San Francisco), Dr Nikolaj Scharff (Zoological Museum, University of Copenhagen (ZMUC)) and Dr Jason Dunlop (Museum für NaturKunde, Berlin) for writing recommendation letters for me to apply for SYNTHESYS projects. Dr Marshal Hedin (San Diego State University) and Dr Dimitri Dimitrov (ZMUC) are acknowledged for their comments on the molecular phylogenetic part of my study. Dr Miquel Arnedo (University of Barcelona), Dr Mike Rix (Western Australian Museum, Perth) and Dr Ingi Agnarsson (University of Vermont, Burlington) provided constructive comments on the phylogenetic part of my study, for that I am very thankful. I thank my friend and colleague, Dr Henrik Krehenwinkel (Max Planck Institute for Evolutionary Biology, Plön) for providing facilities and work space at the molecular lab and teaching me novel methods of DNA isolation. I am thankful to Prof Dr Mohammad R. Rahiminijad (Isfahan University) for his comments during my doctoral research as the scientific mentor.

I would like to thank all the curators listed in the ―Material and Methods paragraphs‖ (chapters 3.1 & 3.2) who provided me with important material for my research. I am grateful to Mrs Janet Beccaloni (NHM, London), Dr Jason Dunlop (ZMB, Berlin), Dr Christina Rollard (MNHN, Paris), Dr Nikolaj Scharff (ZMUC, Copenhagen) and Dr Rudy Jocqué (MRAC, Tervuren) for their hospitality during my visits and study to their scientific collections. I am thankful to Dr Tharina Bird (Windhoek), Mr Martin Forman (Prague), Dr Charles R. Haddad (Bloemfontein), Mr Sérgio Henriques (Lisbon), Dr Vladimír Hula (Brno), Mr František Kovařík (Prague), Mr Kadir B. Kunt (Ankara), Mr Dirk Kunz (Frankfurt am Main), Mr Vláďa Trailin (Hradec Králové), Mr Siegfried Huber (Oberuhldingen), Mr Arnaud Henrard (Tervuren), and Dr Axel Schönhofer (Mainz) for collecting several fresh specimens especially for molecular part of my study. I appreciate the assistances and suggestion from Dr Svetlana Nikolaeva (ICZN, London), Dr Ambros Hänggi (Basel), Dr Cristina A. Rheims (São Paulo), Dr Norman Larsen (Cape Town), Dr Astri Leroy (Roodepoort), Dr Christoph Muster (Leipzig), Dr Christian Hof (Frankfurt am Main), Mr Rowley Snazell (Swanage), Mr Imran Khaliq (Frankfurt am Main) and Mr Youcef Alioua (Batna) that improved my research. My excursion to Ethiopia (2011) was arranged by my friends Dr Vladimír Hula and Dr Jana Nivobová (Mendel University, Brno), because of that I am very grateful.

For the financial supports of this dissertation, the Ministry of Science, Research and Technology of Iran (MSRT) is acknowledged for providing a PhD scholarship for me and my family to study and live in Germany. I am indebted to the Department of Biology, Isfahan University (Iran) for introducing me to the MSRT. The Senckenberg Research Institute and Natural History Museum (SMF) provided research space and partially financial supports to visit scientific collections, participate in congresses as well as travel to Ethiopia. This research also received supports from the SYNTHESYS Project (http://www.synthesys.info/) which is financed by the European Community Research Infrastructure Action under the FP7 "Capacities" Program to visit NHM (2011) and RMCA (2012) collections. The EDIT (European Distributed Institute of Taxonomy) provided a grant for me to participate in the ―Phylogenetic systematic and molecular dating course‖ (2011, ZMUC). The GRADE (Goethe Graduate Academy) is acknowledged for providing very useful workshops and courses which I attended and also its financial support under the ―completion scholarship programme for international doctoral candidates‖ which was funded by the STIBET-programme of the DAAD.

7. REFERENCES

Agnarsson, I., Coddington, J. A., Kuntner, M. (2013) Systematics, progress in the study of spider diversity and evolution. In: D. Penny (Ed), Spider Research in the 21st Century. Siri Scientific Press, Manchester, pp. 58-111.

Agnarsson, I., Rayor, L.S. (2013) A molecular phylogeny of the Australian huntsman spiders (Sparassidae, Deleninae): Implications for taxonomy and social behaviour. Molecular phylogenetics and Evolution, (in press).

Arnedo, M. A., Coddington, J., Agnarsson, I. & Gillespie, R. G. (2004) From a comb to a tree:

phylogenetic relationships of the comb-footed spiders (Araneae, Theridiidae) inferred from nuclear and mitochondrial genes. Molecular phylogenetics and Evolution, 31, 225-245.

Barnard, P., Brown, C. J., Jarvis, A. M., Robertson, A. (1998) Extending the Namibian protected areas network to safeguard hotspots of endemism and diversity. Biodiversity and Conservation, 7, 531-547.

Barrett, R. D. H. & Hebert, P. D. N. (2005) Identifying spiders through DNA barcodes. Canadian Journal of Zoology-Revue Canadienne De Zoologie, 83, 481-491.

Bayer, S. & Schönhofer, A. L. (2013) Phylogenetic relationships of the spider family Psechridae inferred from molecular data, with comments on the Lycosoidea (Arachnida : Araneae).

Invertebrate Systematics, 27, 53-80.

Coddington, J. (2005) Phylogeny and classification of spiders. In: D. Ubick, Paquin, P., Cushing, P. E., RothV., (Ed), Spiders of North America: an identification manual. American Arachnological society, pp. 18-24.

Coddington, J. A. & Levi, H. W. (1991) Systematics and Evolution of Spiders (Araneae). Annual Review of Ecology and Systematics, 22, 565-592.



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