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Kugelspinnen - Eine Einführung (Araneae, Theridiidae)

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Kugelspinnen - Eine Einführung (Araneae, Theridiidae) ZOBODAT - www.zobodat.at Zoologisch-Botanische Datenbank/Zoological-Botanical Database Digitale Literatur/Digital Literature Zeitschrift/Journal: Denisia Jahr/Year: 2004 Band/Volume: 0012 Autor(en)/Author(s): Knoflach Barbara, Pfaller Kristian Artikel/Article: Kugelspinnen - eine Einführung (Araneae, Theridiidae). 111-160 © Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at Kugelspinnen - eine Einführung (Araneae, Theridiidae) B. KNOFLACH & K. PFALLER Abstract: Comb-footed spiders - an introduction (Araneae, Theridiidae). Comb-footed spiders rep- resent a comprehensive, derived and successful orbicularian spider family. They fascinate by a wide spec- trum of morphological and biological traits. The present paper gives an overview on their morphology and behaviour, with emphasis on the European species. Key words: Theridiidae, morphological characterisation, behaviour. Einleitung 111 Einleitung Morphologische Charakterisierung... 115 Der Name Theridiidae geht auf das alt- Körpergröße 115 griechische Theridion zurück und bedeutet Färbung, Zeichnung 115 kleine Bestie. So wurde die Typusgattung Prosoma 117 auch als Wildspinne bezeichnet (MENGE Augen 123 1868: 164). Kugelspinnen (oder Hauben- Cheliceren 123 netzspinnen) faszinieren durch enorme Labium 123 Vielgestaltigkeit hinsichtlich Färbung, Ha- Sternum 123 bitus, Genitalmorphologie und Biologie. Weiblicher Palpus 123 Die deutschen Namen weisen auf das kugel- Beine 127 förmige Abdomen vieler Arten bzw. auf die Borstenkamm 127 Form des Schlupfwinkels hin (WlEHLE Stridulationsorgan 127 1937). Als charakteristisches Familienmerk- Abdomen 129 mal gilt der Borstenkamm an den vierten Epiandrische Drüsen 130 Tarsen, der in der englischen Bezeichnung Spinnwarzen 132 „comb-footed spiders" verankert ist. Mit sei- Genitalorgane 135 ner Hilfe bewerfen sie die Beute mit Leimfä- Diagnose 143 den. Die Leimtropfen werden von besonde- Biologie 145 ren Spinndrüsen, den Glandulae aggregatae, Netze 145 produziert. Auf diese Weise sind Theridiidae Beutefang und Beute 147 in der Lage, verhältnismäßig große und Giftigkeit 148 wehrhafte Beutetiere, u.a. Ameisen, zu über- Kokon und Brutfürsorge 149 wältigen. Kugelspinnen sind wegen ihres Parasiten, Feinde 152 Giftes auch für den Menschen von Bedeu- Abwehrverhalten 153 tung. Die berüchtigten „Schwarzen Wit- Verbreitung 154 wen" (Gattung Latrodectus, Abb. la-f, 28d, Dank 155 29d) waren schon in der Antike gefürchtet Literatur 155 und sind schon bei ARISTOTELES erwähnt (KOBERT 1901; DAHL 1926; HABERMEHL 1994). In Amerika, Afrika und Australien verhalten sie sich zunehmend synanthrop. Manche Arten breiten sich über Verkehr Denisia 12, zugleich Kataloge und Handel aus. So haben Latrodectus der OÖ. Landesmuseen Neue Serie 14 (2004), 111-160 © Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at Abb. 1a-f: Die „Schwarzen Witwen" - prominenteste Vertreter der Theridiidae. a: Latrodectus tredecimguttatus (Rossi), 9 am Kokon (Kroatien, Rovinj, Juli 1993). b: Idem, ventral. Die charakteristische Sanduhrzeichnung ist unterbrochen, c, d: L. geometricus C.L. KOCH, 9, „Braune Witwe", ventral, mit Sanduhrzeichnung (d) (c Jemen, Wadi Warazan, Aug. 1999; d Socotra, Okt. 2000). e: L renivulvatus DAHL, 9 subadult, mit Zeichnung (Jemen, Sana'a, Okt. 1998). f: L. dahli LEVI, 9 adult, Zeichnung weitgehend verschwunden (Jemen, Socotra Okt. 2000). Für die Gattung charakteristisch: Seitenaugen deutlich getrennt (c, f). Exemplare von Jemen leg. A. VAN HARTEN. 112 © Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at metricus C.L. KOCH und die australische L. (1962) haben die 158 damals anerkannten hasselti THORELL adventiv als Neuankömm- Gattungen auf 43 reduziert. Seitdem ist ihre linge Japan erreicht (ONO 1995 und pers. Zahl wieder auf 80 gestiegen. Die neuen Au- Mitt.)- Kugelspinnen stellen einen be- toren tendieren dazu, „Großgattungen" wie- trächtlichen Anteil der synanthropen Spin- der aufzuspalten bzw. den alten Artengrup- nen (THALER & KNOFLACH 1995). pen Gattungsrang zu geben (u.a. WUNDER- LICH 1991, 1995c, d; YOSHIDA 2001). Die Brutpflege und Sozialverhalten sind bei „trennende Diskontinuität" ist dabei wohl manchen Arten weit entwickelt. Die mittel- Ermessenssache. europäischen Vertreter Theridion impressum L. KOCH und T. sisyphium (Q.ERCK) gehören Die Erforschung der Spinnenfauna hat zu den periodisch-sozialen Spinnen mit ihren Ursprung in Europa. So sind die Mund-zu-Mund-Fütterung durch Regurgita- mitteleuropäischen Vertreter der Familie tion (KULLMANN 1974). In den Tropen fin- gut bekannt (CHYZER & KULCZYNSKI 1894; den sich auch zeitlebens soziale Arten, deren WIEHLE 1937; TULLGREN 1949; LOCKET & gemeinschaftliches Netz enorme Dimensio- MLLLIDGE 1953; MILLER 1971; PALMGREN nen erreicht und Bäume überziehen kann 1974; ROBERTS 1985, 1995; KNOFLACH &. (Achaearanea wau LEVI, LUBIN & ROBINSON, THALER 1998; NENTWIG et al. 2003). Den- insbesondere Anelosimus eximius (KEYSER- noch gibt es auch in diesem Gebiet überra- LING) u.a.). Mehr als ein Drittel der weltweit schende Neuentdeckungen, besonders etwa 20 permanent sozialen Spinnen sind Kleinformen, wie Robertus kuehnae BAUCH- Kugelspinnen (AviLES 1997; AGNARSSON HENSS & UHLENHAUT, Theonoe sola THALER 2002). Andere Arten leben als Kleptoparasi- & STEINBERGER, Camiella brignolii THALER ten in Netzen anderer Spinnen und stehlen & STEINBERGER, Vertreter einer neuen Gat- deren Beute, wie die im Mittelmeerraum tung aus Kärnten. Weitere Ergänzungen be- weit verbreitete Diebsspinne Argyrodes arg)' treffen die Differenzierung von kryptischen rodes (WALCKENAER) (siehe KULLMANN Arten, so der Sammelarten Theridion denti- 1959,1960). Verwandte Arten erbeuten spe- culatum (siehe WlEHLE 1960) und Enoplo- ziell die Netzinhaber und verhalten sich so- gnatha ovata (CLERCK) (siehe HlPPA & OK- mit araneophag (WHITEHOUSE 1987). SALA 1982), sowie die Unterscheidung von Achaearanea simulans (THORELL) und A. te- Theridiidae besiedeln alle Straten, von pidariorum (C. L. KOCH) (siehe LOCKET & der Bodenoberfläche bis zum Kronenbereich LUCZAK 1974). Sogar unter den Witwen (WIEHLE 1937; BRAUN & RABELER 1969; wurde vor kurzem von MELIC (2000) eine PALMGREN 1974). Besonders stark treten sie neue Art in Spanien entdeckt, Latrodectus in der Kraut- und Strauchschicht und daher lilianae. Die Artenzahl nimmt nach Norden zahlreich in Klopffängen auf (KNOFLACH & hin ab sowie mit steigender Höhenlage. BERTRANDI 1993). In Barberfallen sind sie Dementsprechend leben im Mittelmeer- wegen ihrer sessilen Lebensweise unterre- raum mehr Arten. Doch befinden sich dar- präsentiert. Ihre Netze sind meist dreidi- unter auch zahlreiche „species inquirendae", mensionale Gerüstnetze mit Fangfäden nicht zu deutende oder unzureichend be- (englisch „cob web weavers") und einem schriebene Arten. Zudem sind vielfach gut haubenförmigen Schlupfwinkel (Retraite), charakterisierte Arten nur nach einem Ge- der zum deutschen Namen Haubennetzspin- schlecht bekannt. Zahlreiche rezente Neu- nen geführt hat. Jedoch gibt es zahlreiche beschreibungen (LEVY 1998; WUNDERLICH Abwandlungen und Reduktionserscheinun- 1987, 1991, 1995a, d-f) stehen den „alten" gen (siehe BENJAMIN & ZSCHOKKE 2003). Arten gegenüber. Diese unbefriedigende ta- Theridiidae zählen mit ca. 2.200 Arten xonomische Situation ist nur durch Revi- zu den diversen Spinnenfamilien und stellen sionsarbeiten zu bewältigen (siehe KNOF- 6% aller bekannten Spinnen (PLATNICK LACH 1994, 1996c, 1999; BOSMANS & VAN 2003). In den Tropen erreichen sie die größ- KEER 1999). Die Differenzierung der Arten te Vielfalt. LEVI hat von 1953 bis 1970 (mit beruht überwiegend auf morphologischen Nachträgen) eine weltweite taxonomische Kriterien, insbesondere der Genitalmorpho- Basis geschaffen, besonders durch Revisio- logie. Die Morphospezies decken sich in der nen über die Arten Amerikas. LEVI &. LEVI Regel mit der biologischen Realität. Um 113 © Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at Abb. 2a-h: Vielfalt der Kugelspinnen, Habitus und Färbung, a: Euryopis episinoides (WALCKENAER), er, vagant, Ameisenfresser (Kefallonia, Mai 2002). b, d: Enoplognatha afrodite HIPPA & OKSALA, 9, Farbpolymorphismus (Kefallonia, Mai 2002). c: f. latimana HIPPA & OKSALA, 9 mit Eikokon (Polen, Bialowiezy, Juli 1999). e, f: Macaridion barreti KULCZYNSKI, 9, in Makaronesien endemisch (Teneriffa, Anaga, Feb. 2000). g: Thehdion melanostictum O. P.-CAMBRIDGE, er, pantropisch (Jemen, Sana'a, Juni 1998). h: Theridula gonygaster (SIMON), juv. (Cabo Verde, Santiago, Jan. 2000, leg. A. VAN HARTEN). 114 © Biologiezentrum Linz/Austria; download unter www.biologiezentrum.at dem Konzept der Biospezies (MAYR 1975) planaren Radnetzen der Araneidae als gerecht zu werden, müssen biologische Kri- hochmodifiziert gegenübergestellt und sol- terien berücksichtigt werden, Fortpflan- len zum evolutiven Erfolg der Theridiidae zungsbiologie und ökologische Einnischung. geführt haben. Auf diese Weise konnten Kugelspinnen dem Prädationsdruck durch Die morphologische Mannigfaltigkeit Sphecidae entkommen, im Gegensatz zu spiegelt sich in der Biologie der Theridiidae den Radnetzspinnen (BLACKLEDGE et al. wider. Trotz der umfassenden Bearbeitung 2003). Ihre Stellung war lange Zeit unsi- von NIELSEN (1931, 1932) ist das Wissen zur cher: „Despite much work, the placement of Biologie vieler Arten noch immer lücken- Theridiidae and its relatives within Araneo- haft. Nur wenige sind ausführlich unter- idea remains ambiguous" (CODDINGTON & sucht: z.B. LEVI 1991). Nach HORMIGA et al. (1995) Achaearanea riparia (BLACKWALL): siehe
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