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The Ant Genus Cardiocondyla (Inserta: Hymenoptera: Formicidae) - a Taxonomic Revision of the G Elegans, G Bulgarica, G Batesii, C

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The Ant Genus Cardiocondyla (Inserta: Hymenoptera: Formicidae) - a Taxonomic Revision of the G Elegans, G Bulgarica, G Batesii, C ©Naturhistorisches Museum Wien, download unter www.biologiezentrum.at Ann. Naturhist. Mus. Wien 104 B 203 - 338 Wien, März 2003 The ant genus Cardiocondyla (Inserta: Hymenoptera: Formicidae) - a taxonomic revision of the G elegans, G bulgarica, G batesii, C. nuda, G shuckardi, G stambuloffii, G wroughtonii, G emeryj, and G minutior species groups B. Seifert* Abstract A taxonomic revision of all holarctic species groups of the ant genus Cardiocondyla is given, including all worldwide species groups, with at least one tramp species. General aspects of Cardiocondyla biology such as habitat selection, nest construction, behaviour, the ergatoid male syndrome, and gyne polymorphism are treated. 48 species are recognized, 20 of which are described as new: Cardiocondyla bicoronata sp.n. (Israel, Jordan, United Arab Emirates, Yemen, Turkestan), C brachyceps sp.n. (E Turkey, Iran, Afghanistan), C. breviscapus sp.n. (India), C. gallilaeica sp.n. (Israel), C. goa sp.n. (India), C. israelica sp.n. (Egypt, Israel), C. littoralis sp.n. (S Kazakhstan), C. longiceps sp.n. (Yemen), C. melano sp.n. (Yemen), C. nana sp.n. (Brunei), C. opaca sp.n. (India), C. opistopsis sp.n. (Kuwait), C. paranuda sp.n. (Tunisia), C. persiana sp.n. (Iran, Israel), C. rugulosa sp.n. (Yemen), C. semirubra sp.n. (Asia Minor), C. shagrinata sp.n. (India), C. tenuifrons sp.n. (Jordan), C. tibetana sp.n. (Tibet, Tarim Basin) and C unicalis sp.n. (Iran). Sixteen taxa are newly synonymized: Cardiocondyla bulgarica FOREL, 1892 (= C. elegans var. eleonorae FOREL, 1911, syn.n.), C. elegans EMERY, 1869 (= C. dalmatica SOUDEK, 1925, syn.n., = C. provincialis BERNARD, 1956, syn.n.), C.fajumensis FOREL, 1913, stat.n. (= C. schatzmayri FINZI, 1936, syn.n., = C. nilotica WEBER, 1952, syn.n.), C. mauritanica FOREL, 1890 (= C. ectopia SNELLING, 1974, syn.n.), C. minutior FOREL 1899, stat.rev. (= C. tsukuyomi TERAYAMA, 1999, syn.n.), C. nigra FOREL, 1905 (= C. elegans var. torretassoi FINZI, 1936, syn.n.), C. obscurior WHEELER, 1929, stat. n. (= C. bicolor DONISTHORPE, 1930, syn.n.), C. shuckardi FOREL, 1891 (= C. shuckardoides FOREL, 1895, syn.), C. stambuloffii FOREL, 1892 (= C. bogdanovi RUZSKY, 1905; syn.n.), C. venustula WHEELER, 1908 (= C globinodis STITZ, 1923, syn.n., = C. badonei ARNOLD, 1926, syn.n.), C. wroughtonii (FOREL, 1890) (= C. wroughtonii ssp. quadraticeps FOREL, 1912, syn., = C. longispina KARAVAJEV, 1935, syn.n., = C. yamauchii TERAYAMA, 1999, syn.n.). Five taxa are elevated to species rank, and the status of two controversial taxa is revised. Nine names are listed under "Incertae Sedis". Each species is described based upon workers and gynes, as far as either caste is known, and a detailed morphometry by high-resolution steromicroscopy is provided. Each species is depicted in three or four standard viewing positions preferentially on the basis of type material. Key words: Cardiocondyla, taxonomic revision, Holarctic, tramp species, ergatoid males, gyne polymor- phism, new species, new synonymy, new status, behaviour, nest, habitat. Zusammenfassung Eine taxonomische Revision der holarktischen Vertreter der Ameisengattung Cardiocondyla sowie aller kosmopolitischen Artengruppen, in denen mindestens eine Tramp-Spezies vorkommt, wird präsentiert. Allgemeine Aspekte der Cardiocondyla-B\o\ogie wie Habitatwahl, Nistweise, Verhalten, das Syndrom der ergatoiden Männchen und Gynenpolymorphismus werden behandelt. Es werden 48 Arten unterschieden, von denen 20 als neu beschrieben werden: Cardiocondyla bicoronata sp.n. (Israel, Jordan, United Arab Emirates, Yemen, Turkestan), C. brachyceps sp.n. (E Turkey, Iran, Afghanistan), C. breviscapus sp.n. (India), C. gallilaeica sp.n. (Israel), C. goa sp.n. (India), C. israelica sp.n. (Egypt, Israel), C. littoralis sp.n. * Dr. Bernhard Seifert, Staatliches Museum für Naturkunde Görlitz, PSF 300154, D-02806 Görlitz, Germany ©Naturhistorisches Museum Wien, download unter www.biologiezentrum.at 204 Annalen des Naturhistorischen Museums in Wien 104 B (S Kazakhstan), C. longiceps sp.n. (Yemen), C. melana sp.n. (Yemen), C. nana sp.n. (Brunei), C. opaca sp.n. (India), C. opistopsis sp.n. (Kuwait), C. paranuda sp.n. (Tunisia), C. persiana sp.n. (Iran, Israel), C. rugulosa sp.n. (Yemen), C. semirubra sp.n. (Asia Minor), C. shagrinata sp.n. (India), C. tenuifrons sp.n. (Jordan), C tibetana sp.n. (Tibet, Tarim Basin) und Cardiocondyla unicalis sp.n. (Iran). Sechzehn Taxa erscheinen als neue Synonyme: Cardiocondyla bulgarica FOREL, 1892 (= C. elegans var. eleonorae FOREL, 1911, syn.n.), C. elegans EMERY, 1869 (= C. dalmatica SOUDEK, 1925, syn.n., = C. provincialis BERNARD, 1956, syn.n.), C.fajumensis FOREL, 1913, stat.n. (= C. schatzmayri FINZI, 1936, syn.n., = C. nilotica WEBER, 1952, syn.n.), C. mauritanica FOREL, 1890 (= C. ectopia SNELLING, 1974, syn.n.), C. minutior FOREL 1899, stat.rev. (= C. tsukuyomi TERAYAMA, 1999, syn.n.), C. nigra FOREL, 1905 (= C. elegans var. torretassoi FINZI, 1936, syn.n.), C. obscurior WHEELER, 1929, stat. n. (= C. bicolor DONISTHORPE, 1930, syn.n.), C. shuckardi FOREL, 1891 (= C. shuckardoides FOREL, 1895, syn.n.), C. stambuloffii FOREL, 1892 (= C. bogdanovi RUZSKY, 1905; syn.n.), C. venustula WHEELER, 1908 (= C. globinodis STITZ, 1923, syn.n., = C. badonei ARNOLD, 1926, syn.n.), C. wroughtonii (FOREL, 1890) (= C. wroughtonii ssp. quadraticeps FOREL, 1912, syn.n., = C. longispina KARAVAJEV, 1935, syn.n., = C. yamauchii TERAYAMA, 1999, syn.n.). Fünf Taxa erfuhren eine Rangerhöhung und der Status zweier umstrittener Taxa wird revidiert. Neun Namen werden unter 'Incertae Sedis' aufgelistet. Jede Art wird anhand von Arbeitern und Gynen, soweit diese bekannt sind, beschrieben, durch eine detailierte Morphometrie mittels Hochleistungsstereomikroskopie vorgestellt und in drei oder vier Standardansichten abgebildet, wobei Typenmaterial bevorzugt dargestellt ist. Content 1. Introduction 206 2. Material studied 207 3. Methods and terminology 208 3.1 Definition of descriptive terms 208 3.2 Definition of numeric characters 209 3.3 The drawings 211 3.4 Discriminant functions 212 3.5 Predicting characters of unknown workers or gynes 212 4. The allometric variance of characters 212 5. Taxonomic significance of characters 213 6. General aspects of Cardiocondyla biology 213 6.1 Habitats, nesting, and behaviour 213 6.2 The ergatoid male syndrome: all ergatoids show a fighter-phenotype 215 6.3 Gyne polymorphism - an adaptive trait emerging in desert environments 216 7. Diagnosis of Cardiocondyla 218 7.1 Diagnosis of the worker 218 7.2 Diagnosis of the species groups 219 8. A synonymic list of species and their main distribution 221 9. Cardiocondyla incertae sedis 224 10. Treatment by species 225 10.1 Cardiocondyla elegans EMERY, 1869 225 10.2 Cardiocondyla brachyceps sp.n. 228 10.3 Cardiocondyla ulianini EMERY, 1889, stat.rev. 229 10.4 Cardiocondyla gallilaeica sp.n. 230 10.5 Cardiocondyla israelica sp.n. 231 10.6 Cardiocondyla littoralis sp.n. 232 10.7 Cardiocondyla bulgarica FOREL, 1892 233 10.8 Cardiocondyla sahlbergi FOREL, 1913 235 10.9 Cardiocondyla persiana sp.n. 236 ©Naturhistorisches Museum Wien, download unter www.biologiezentrum.at SEIFERT: The ant genus Cardiocondyla - a taxonomic revision of species groups 205 10.10 Cardiocondyla batesii FOREL, 1894 237 10.11 Cardiocondyla semirubra sp.n. 238 10.12 Cardiocondyla kushanica PISARSKJ, 1967 239 10.13 Cardiocondyla nigra F'OREL, 1905 240 10.14 Cardiocondyla bicoronata sp.n. 242 10.15 Cardiocondyla tenuifrons sp.n. 243 10.16 Cardiocondyla rugulosa sp.n. 244 10.17 Cardiocondyla opistopsis sp.n. 244 10.18 Cardiocondyla nuda (MAYR, 1866) 245 10.19 Cardiocondyla paranuda sp.n. 246 10.20 Cardiocondyla atalanta FOREL, 1915, stat.nov. 247 10.21 Cardiocondyla mauritanica FOREL, 1890 248 10.22 Cardiocondyla mauritanica FOREL, morph B 250 10.23 Cardiocondyla kagutsuchi TERAYAMA, 1999 252 10.24 Cardiocondyla strigifrons VIEHMEYER, 1922, stat.nov. 255 10.25 Cardiocondyla shuckardi FOREL, 1891 257 10.26 Cardiocondyla venustula WHEELER, 1908 257 10.27 Cardiocondyla melana sp.n. 259 10.28 Cardiocondyla longiceps sp.n. 259 10.29 Cardiocondyla fajumensis FOREL, 1913, stat.nov. 260 10.30 Cardiocondyla unicalis sp.n. 262 10.31 Cardiocondyla stambuloffii FOREL, 1892 262 10.32 Cardiocondyla koshewnikovi RUZSKY, 1902 266 10.33 Cardiocondyla gibbosa KUZNETZOV-UGAMSKY, 1927, stat.nov. 268 10.34 Cardiocondyla tibetana sp.n. 269 10.35 Cardiocondyla wroughtonii (FOREL, 1890) 269 10.36 Cardiocondyla obscurior WHEELER, 1929, stat.nov. 271 10.37 Cardiocondyla shagrinata sp.n. 275 10.38 Cardiocondyla nana sp.n. 276 10.39 Cardiocondyla emeryi FOREL, 1881 276 10.40 Cardiocondyla weserka BOLTÖH, 1982 280 10.41 Cardiocondyla neferka BOLTON, 1982 280 10.42 Cardiocondyla yemeni COLLINGWOOD & AGOSTI, 1996 281 10.43 Cardiocondyla minutior FOREL, 1899, stat.rev. 283 10.44 Cardiocondyla goa sp.n. 285 10.45 Cardiocondyla tjibodana KARAVAJEV, 1935 287 10.46 Cardiocondyla breviscapus sp.n. 288 10.47 Cardiocondyla carbonaria F OREL, 1907 289 10.48 Cardiocondyla opaca sp.n. 290 10.49 Cardiocondyla britteni CRAWLEY, 1920 291 11. Key and comparative tables to workers 292 12. Comparative tables to gynes 308 13. Figures 313 14. Acknowledgements 334 15. References 334 ©Naturhistorisches Museum Wien, download unter www.biologiezentrum.at 206 Annalen des Naturhistorischen Museums in Wien 104 B 1. Introduction This taxonomic revision considers the complete Holarctic fauna of the ant genus Cardiocondyla EMERY, 1869 and extends its view to all members of any tropical species group in which at least one tramp species is
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