2007 (2008). The Journal of Arachnology 35:432–462 THE SPIDER GENUS DYSDERA (ARANEAE, DYSDERIDAE) IN CENTRAL EUROPE: REVISION AND NATURAL HISTORY Milan Rˇ eza´cˇ: Department of Entomology, Research Institute of Crop Production, Drnovska´ 507, CZ-161 06 Prague 6-Ruzyneˇ, Czech Republic. E-mail: [email protected] Jirˇ´ı Kra´l: Laboratory of Arachnid Cytogenetics, Department of Genetics and Microbiology, Faculty of Science, Charles University in Prague, Vinicˇna´ 5, CZ-128 44 Prague 2, Czech Republic Stano Peka´r: Institute of Botany and Zoology, Faculty of Science, Masaryk University, Kotla´rˇska´ 2, CZ-611 37 Brno, Czech Republic ABSTRACT. Nine species of the genus Dysdera were found to occur in central Europe: D. adriatica Kulczyn´ski 1897, D. crocata Koch 1838, D. dubrovninnii Deeleman-Reinhold 1988, D. erythrina (Walckenaer 1802), D. ninnii Canestrini 1868, D. hungarica Kulczyn´ski 1897, D. lantosquensis Simon 1882, D. longirostris Doblika 1853, and D. taurica Charitonov 1956. Two species, D. dubrovninnii and D. lantosquensis, are newly recorded from central Europe. The original description of D. hombergi (Scopoli 1763), the name used for a common species of the genus Harpactea, probably refers to D. ninnii. We retain the name D. ninnii as a nomen protectum. Dysdera hamulata Kulczyn´ski 1897 appears to be a junior synonym of D. maurusia Thorell 1873. This North African species probably does not occur in central Europe, and a previous record from Slovakia is probably based on mislabeled material. A review of all species of Dysdera named from outside the Palearctic region demonstrated that D. australiensis Rainbow 1900 and D. magna Keyserling 1877 are junior synonyms of D. crocata, and that D. bicolor Tatzanovski 1874 and D. solers Walckenaer 1837 are erroneously placed in the genus Dysdera; the former is likely to be an oonopid and the latter a caponiid. In central Europe, Dysdera spiders prefer xerothermic forests, particularly sites enriched by calcium. All species probably have biennal life-cycles. The karyotype of males of seven species were examined, and diploid chromosome numbers were found to be extraordinarily variable, ranging from 9 (D. crocata)to40(D. longirostris). Karyotypes consist of holocentric chromosomes. Keywords: Sibling species, karyotype, geographic parthenogenesis, taxonomy, thelytoky Spiders of the genus Dysdera (Dysderidae) and the presence of sibling species (e.g., Deele- are ground dwellers characteristic of xerother- man-Reinhold & Deeleman 1988). mic forests of the Mediterranean and adjacent A modern revision of the genus was initiated areas. During the day, they shelter in gravel by Deeleman-Reinhold & Deeleman (1988), covered by organic material or under stones, focusing on species from the eastern part of the and at night they search for woodlice, their Mediterranean. The genus was redefined and principal prey (Cooke 1965). divided into different species-groups. This Comprising more than 240 species (Platnick paper revises central European species of the 2007), Dysdera is currently the largest genus in genus Dysdera, based mainly on analysis of the family Dysderidae and one of the richest material from the Czech Republic and Slova- Palearctic spider genera. Interestingly, the vast kia. We solve some nomenclatural problems majority of species appear to be endemic to and summarize data on the distribution, only small areas of the Mediterranean region, habitat preferences, phenology and karyotypes and only nine representatives appear to have of the species. We recognize eight species colonized central Europe after the last glacial representing five groups within central Europe: period. Although the species diversity in this D. crocata C.L. Koch 1838 (crocata group); D. region is low, there has been much confusion ninnii Canestrini 1868, D. dubrovninnii Deele- concerning their identification because of the man-Reinhold 1988 (ninnii group); D. hungar- uniformity in both the shape and body color, ica Kulczyn´ski 1897, D. adriatica Kulczyn´ski similarity in external female genitalic features 1897, D. longirostris Doblika 1853 (longirostris 432 Rˇ EZA´ Cˇ ET AL.—THE GENUS DYSDERA IN CENTRAL EUROPE 433 group); D. taurica Charitonov 1956 (lata Hrusˇov, Slovakia, 2 L; D. adriatica – Korana group); D. erythrina (Walckenaer 1802) and near Plitvicˇka jezera, Croatia, 1 L; D. long- D. lantosquensis Simon 1882 (erythrina group). irostris – Kranevo near Varna, Bulgaria, 4 L; and D. taurica – Kranevo near Varna, Bulgaria, MATERIAL AND METHODS 2 L. Chromosome preparations were examined Distributional data and habitat preferences under immersion lens using an Olympus BX 50 were obtained by analysis of extensive material light microscope. from collections and during our field work. Specimens are lodged in the following Selected localities were visited mainly in the institutions: private collection of Alesˇ Jelı´nek, summers of 1999–2005. Vegetation of inspected Telcˇ, Czech Republic (AJ); Australian Muse- localities was characterized following Chytry´et um, Sydney, Australia (AMS); Museum of al. (2001) and Moravec (1995). Natural History, London, UK (BMNH); pri- Specimens were examined using a Nikon vate collection of F. Gasparo, Trieste, Italy SMZ 645 stereomicroscope and an Olympus (FG); Magyar Terme´szettudoma´nyi Mu´zeum, BX51 light microscope. Before examination, Budapest, Hungary (HNHM); private collec- female vulvae were dissected and cleared in tion of J. Dolansky´, Pardubice, Czech Republic glycerol. The prosoma, chelicerae and bulbi of (JD); private collection of J. Svatonˇ, Martin, selected males were removed, placed on a stub, Slovakia (JS); private collection of L. Kubcova´, coated with gold and examined using a scanning Prague, Czech Republic (LK); private collec- electron microscope JEOL JSM 6400. To tion of M. Antusˇ, Prague, Czech Republic describe structures of the male pedipalp and (MA); Muse´um d’Histoire Naturelle, Gene`ve, the female vulva, we used the terminology of Switzerland (MHNG); Muse´um National Arnedo et al. (2000). d’Histoire Naturelle, Paris, France (MNHN); Phenology was studied both on selected private collection of M. Rˇ eza´cˇ, Prague, Czech localities and by processing data on labels of Republic (MR); Naturhistoriska Riksmuseet, the revised material. Phenological observations Stockholm, Sweden (NHRS); Na´rodnı´ Mu- were performed on data from the following zeum, Prague, Czech Republic (NMPC); Nat- localities: Roksˇtejn [49u199N, 15u439E], Czech urhistorisches Museum, Vienna, Austria Republic (D. ninnii); Vinne´ near Michalovce (NMW); private collection of P. Gajdosˇ, Nitra, [48u489N, 21u589E], Slovakia (D. dubrovninnii); Slovakia (PG); South Australian Museum, Hrusˇov [48u369N, 20u409E] and Vinne´ near Adelaide, Australia (SAM); Naturmuseum Michalovce, Slovakia (D. hungarica); Plitvicˇka Senckenberg, Frankfurt am Main, Germany jezera [44u549N, 15u369E], Croatia (D. adria- (SMF); Univerza v Ljubljani, Slovenia (UL); tica); Rilski monastir [42u079N, 23u209E] and Universidad de La Laguna, Spain (ULCI); Kranevo [43u209N, 28u029E], Bulgaria (D. long- private collection of V. Bryja, Brno, Czech irostris); and Kranevo, Bulgaria (D. taurica). Republic (VB); private collection of V. Hula, For the karyological analyses, the most Brno, Czech Republic (VH); Vihorlatske´mu´- appropriate ontogenetic stage was found to be zeum, Humenne´, Slovakia (VMH); private the adult male shortly after molting, which collection of V. Ru˚zˇicˇka, Cˇ eske´ Budeˇjovice, occurs at the end of the summer in all species Czech Republic (VR); Western Australian studied. Testes at this stage contained numer- Museum, Perth, Australia (WAM); private ous dividing cells suitable for karyotypic collection of Z. Majkus, Ostrava, Czech Re- analysis, namely spermatogonial mitoses as public (ZM); Museum fu¨r Naturkunde, Hum- well as various meiotic stages. The chromo- boldt Universita¨t, Berlin, Germany (ZMHB). some preparations were obtained by the meth- od described in Peka´r & Kra´l (2001). Localities TAXONOMY of karyotyped species were as follows: D. Family Dysderidae C.L. Koch 1837 crocata – Kranevo near Varna, Bulgaria, 1 L; Genus Dysdera Latreille 1804 C¸ aytepe near Ordu, Turkey, 1 L; Mitra near E´ vora, Portugal, 4 L; Bloemfontein, South Type species.—Dysdera erythrina (Walcke- Africa, 1 L; Taborno, Tenerife, Spain, 2 L; D. naer 1802). ninnii – Roksˇtejn near Brtnice, Czech Republic, Remarks.—Comprising more than 240 2 L; D. dubrovninnii – Vinne´ near Michalovce, named species (Platnick 2007), Dysdera is Slovakia, 2 L; D. hungarica – Hradisko near currently the largest genus in the family 434 THE JOURNAL OF ARACHNOLOGY Dysderidae. The vast majority of species region; only nine representatives appear to have appear to be endemic to the Mediterranean colonized central Europe. KEY TO THE SPECIES OF CENTRAL EUROPEAN DYSDERA 1. Carapace smooth with rounded pits . 2(ninnii group) Carapace wrinkled, without rounded pits . ......... 3 2. Cheliceral fang not flattened . ... Dysdera ninnii Cheliceral fang dorsoventrally flattened . Dysdera dubrovninnii 3. Tibiae III and IV with one or more dorsal spines . Dysdera taurica (lata group) Tibiae III and IV without dorsal spines . ........ 4 4. Femur IV with one or more dorsal spines . Dysdera crocata (crocata group) Femur IV without dorsal spines . ....... 5 5. Lateral anterior margins of carapace parallel (dorsal view), inner margin of basal cheliceral segment concave . 6 (erythrina group) Lateral anterior margins of carapace convergent (dorsal view), inner margin of basal cheliceral