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Fossil and Extant Spiders (Araneae) Fossile Und Heutige Spinnen BEITR. ARANEOL., 5 (2008) FOSSIL AND EXTANT SPIDERS (ARANEAE) FOSSILE UND HEUTIGE SPINNEN Joerg Wunderlich BEITR. ARANEOL., 5 (2008) BEITRAEGE ZUR ARANEOLOGIE, 5 (2008). – ISBN 978-3-931473-11-2. FOSSIL AND EXTANT SPIDERS (ARANEAE) Photos on the front cover of the book (the bodies of four extraordinary fossil spiders): Photo ABOVE ON THE LEFT: Male spider of the extinct family Spatiatoridae in Eocene Baltic amber, body length 4.3 mm. This spider existed 40 to 50 million years ago. Note the raised “caput”. See photo 139. Photo ABOVE ON THE RIGHT: Male spider of the family Borboropactidae in Baltic amber, body length 3.5 mm. This is a relict family of the tropics. Note the powerful anterior “capture legs”. See photo 380. Photo BELOW ON THE LEFT: Male spider of the family Pumiliopimoidae in Baltic amber, body length 1.9 mm. The family is almost extinct; a single extant species, which I regard as related, has recently been described from North America. See photo 156. Photo BELOW ON THE RIGHT: Juvenile spider of the extinct family Lagonomegopidae in Lower Cretaceous amber from Burma, body length 1.2 mm. The spider existed about 100 million years ago. Note the very large eyes in a lateral position. See photo 90. Fotos auf dem Buchdeckel (Körper vier außergewöhnlicher fossiler Spinnen): Foto OBEN LINKS. Männliche Spinne der ausgestorbenen Familie Spatiatoridae in Bal- tischem Bernstein, Körper-Länge 4.3 mm. Diese Spinne lebte vor 40–50 Millionen Jahren. Ihr “Kopfteil” ist deutlich erhöht. Siehe Foto 139. Foto OBEN RECHTS: Männliche Spinne der nahezu ausgestorbenen tropischen Familie Borboropactidae in Baltischem Bernstein, Körper-Länge 3.5 mm. Ihr vorderes Beinpaar ist als mächtige Fangbeine ausgebildet. Siehe Foto 380. Foto UNTEN LINKS: Männliche Spinne der nahezu ausgestorbenen Familie Pumiliopimoi- dae in Baltischem Bernstein, Körper-Länge 1.9 mm. Eine vermutlich verwandte Art wurde kürzlich aus Nord-Amerika beschrieben. Siehe Foto 156. Foto UNTEN RECHTS: Junge Spinne der aus der Unteren Kreidezeit bekannten ausgestor- benen Familie Lagonomegopidae in Burmesischem Bernstein, Körper-Länge 1.2 mm. Diese Spinne lebte vor etwa 100 Millionen Jahren und besaß ein Paar enorm entwickelte Augen in seitlicher Position. Siehe Foto 90. Photos on the backcover of the book: Photo above: An extant Cretaceous spider from Brazil (Zygiellinae). Photo below: An extant spider from Australia (Theridiidae). 1 2 BEITR. ARANEOL., 5 (2008) FOSSIL AND EXTANT SPIDERS (ARANEAE) Editor and author: Joerg Wunderlich © Publishing House: Joerg Wunderlich, 69493 Hirschberg, Germany Print: GROSCH! Druckzentrum, 69214 Eppelheim Orders for this volume: Publishing House JOERG WUNDERLICH Oberer Haeuselbergweg 24, 69493 Hirschberg, Germany E-Mail: [email protected] ISBN 978-3-931473-11-2 3 BEITR. ARANEOL., 5 (2008) FOSSIL AND EXTANT SPIDERS (ARANEAE) Phylogeny, diversifications, extinctions, biogeography, ecology and ethology; with descriptions of new fossil and extant taxa JOERG WUNDERLICH Remark: Unfortunately just before the printing of this volume a defect in the soft- ware caused some trouble with the manuscript, e. g. some letters and words were lost or changed, and some corrections as well. 4 BEITR. ARANEOL., 5 (2008) CONTENTS Key words of most papers of this volume page Acknowledgements: See the paper on Cretaceous spiders in this volume. ...... 10 Introduction.. 11 (1) CONTRIBUTION ON FOSSIL AND EXTANT SPIDER ................... 14 Abstract . 14 Key words ........................................................ 14 Corrections and supplements regarding vol. 3 Beitr. Araneol. (2004) (“Spiders in Amber and Copal”) ................................... 14 Remarks on J. DUNLOP’s (2005, 2006) review of J. WUNDERLICH’s (2004) “Spiders in Amber and Copal” .................................... 17 Material .......................................................... 19 (1) Origin of the material ........................................... 19 (2) Storing the material/institutions ................................... 19 (3) Preservation of the material . 20 (4) Selection of holotypes and the importance of paratypes . 21 Techniques of study and documentation ................................. 22 Methods of study . 22 Preservation of selected structures of the inclusions ....................... 24 Fakes of inclusions, imitations and changes by mistake ..................... 27 PHYLOGENETICS AND BIOGEOGRAPHY ............................ 29 Evolutionary “trends” . 29 Phylogenetic relationships: Morphological/taxonomical connecting (“missing”) links, the taxonomical importance of fossil/extinct spider taxa, the concepts of genera and remarks on spider’s diversity ............ 31 On “Faunas connecting fossils”. 32 Biogeography: Changes of ranges in two araneoid families . 32 Remarks on the Bitterfeld deposit and on Rovno amber ..................... 33 Diversifications of taxa of the family Theridiidae during the Palaeo- cene and the Eocene ............................................. 34 Diversifications during the Cretaceous and Tertiary; gaps in the spider fauna of the Baltic amber forest and on geological old and young higher (suprageneric) taxa ......................................... 34 5 Proof of several diversifications in the same taxon, faunal gaps etc. ........... 35 Extinctions of higher spider taxa, and the results of the Cretaceous- Tertiary extinction events .......................................... 38 TAXONOMY . 40 How discoveries of fossil taxa may change the range of higher taxa ........... 40 Remarks on resurrected taxa ......................................... 41 How different subjective concepts of the taxonomical range of genera may influence conclusions on fossil and extant spiders. 41 Remarks on the species concept in palaeontology ......................... 42 (2) DESCRIPTIONS OF FOSSIL SPIDER TAXA MAINLY IN BALTIC AMBER AS WELL AS OF CERTAIN RELATED EXTANT TAXA ............ 44 Abstract . 44 Key words ........................................................ 45 Family Segestriidae. 45 On extant and fossil (Eocene and Cretaceous) taxa of the new spider subfamily Orchestininae (Araneae: Oonopidae) ................... 53 Family Spatiatoridae ................................................ 79 Family Hersiliidae in Dominican amber . 81 Family Tetragnathidae, with descriptions of new fossil and extant taxa, and remarks on the Guizygiellinae . 81 Family Aranidae: Subfamily Araneinae? ................................ 103 Family Mimetidae ................................................. 109 Family Pimoidae .................................................. 109 Fossil and extant spiders of the “linyphioid branch” of the superfamily Arane- oidea (Araneae), with the description of the new family Pumiliopimoidae ....117 Descriptions of a cryptic fossil and an enigmatic extant genus of the family Synotaxidae. 130 (3) ON EXTANT EUROPEAN AND FOSSIL (EOCENE) COMB-FOOTED SPIDERS (ARANEAE: THERIDIIDAE), with notes on their sub- families, and with descriptions of new taxa .......................... 140 Abstract . 140 Key words ....................................................... 140 INTRODUCTION including remarks on behaviour, biogeography, diversity, evolution, phylogeny and taxonomy . 141 DIAGNOSIS of the family Theridiidae ................................. 144 Subfamilies and tribus . 146 KEY TO THE FOSSIL AND EXTANT SUBFAMILIES .................... 154 PHYLOGENETICS ............................................... 158 Origin/age of the family Theridiidae and its diversifications; co-evolution . 158 Evolutionary “trends”/convergences, reductions and reversals; geologi- cally oldest proofs of peculiar structures and behaviour. 159 Intrafamiliar relationships of higher theridiid taxa ......................... 165 Surviving genera .................................................. 170 Ancestors of extant species?. 170 Remarks on possible subspecies in the fossils . 171 ECOLOGY, ETHOLOGY and FREQUENCY of the Theridiidae in amber ..... 171 6 Cryptic body shape/mimesis ......................................... 172 Sex ratio ........................................................ 172 Prey, leg amputations and healing events. 173 Questionable cannibalism . 173 Parasitizing Acari. 173 Fauna, biodiversity ................................................ 174 “HISTORICAL TAXONOMY”. 176 SYNONYMY of fossils ....................... 176, 186f, 205, 220f, 283f, 316f LIST of the subfamilies, fossil genera and species ........................ 181 DESCRIPTIONS OF THE TAXA IN BALTIC AMBER AS WELL AS OF FEW EXTANT TAXA (Episinae: Episini; ?Pholcommatinae) ........... 183 Asageninae ...................................................... 184 Enoplognathinae .................................................. 238 Pholcommatinae .................................................. 248 Magnopholcommatini n. trib., an extant taxon from Australia. 269 Phoroncidiinae .................................................... 271 Hadrotarsinae .................................................... 279 Episinae. 312 Episini. 316 two extant genera from Africa and Malaysia. 340 Spinitharini n. trib ............................................... 343 Argyrodinae (regarded as a fake) ..................................... 359 Anelosiminae n. subfam ............................................ 361 Theridiinae (questionable). 367 The EXTANT GENERA of the W-Palaearctic Theridiidae ................ 374 LIST OF THE WEST-PALAEARCTIC GENERA and selected characters . 374f New combinations of extant Theridiidae
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