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Stuttgarter Beiträge Zur Naturkunde

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Stuttgarter Beiträge Zur Naturkunde download Biodiversity Heritage Library, http://www.biodiversitylibrary.org/ Stuttgarter Beiträge zur Naturkunde ^ Serie A (Biologie) Herausgeber: Staatliches Museum für Naturkunde, Rosenstein 1, D-70191 Stuttgart Stuttgarter Beitr. Naturk. Ser.A Nr. 537 101 S. Stuttgart, 31.5. 1996 Die Kopulationsorgane des Maikäfers Melolontha melolontha (Insecta: Coleoptera: Scarabaeidae). - Ein Beitrag zur vergleichenden und funktionellen Anatomie der ektodermalen Genitalien der Coleoptera The Copulatory Organs of the Cockchafer Melolontha melolontha (Insecta: Coleoptera: Scarabaeidae). - A Contribution to Comparative and Functional Anatomy of Ectodermal Genitalia of the Coleoptera Von Frank-Thorsten Kr eil, Würzburg o^\tHS0/V% Mit 49 Abbildungen und 1 Tabelle ._„, \ APR 4.1997 Xv S u m m a r y \JJBRARl£5- Sclerites, mcmbranes and muscles of male and female copulatory apparatus of the cockcha- fer Melolontha melolontha (Linnacus, 1758) (Insecta: Coleoptera: Scarabaeidae: Melo- lonthinae) are dcscribcd at rcst and during copulation. For the tust ume, the course ol copu lation of Melolontha is dcscribed complctely. By comparing resting position, copulation and supposed oviposition the functions of Single parts of the copulatory organs are inferred. I>\ means of a functional morphological scenario the functioning of copulation is depicted. \ possiblc strategy of the female to stop copulation before Insemination Starts is described (cryptic female choicc). At rest, the aedeagus lies on its lateral side ("deversement"). The course of cjaculatory duct and trachcae indicate a "retournement", a longitudinal IST rota *) Dissertation zur Erlangung des Grades eines 1 )oktors der Natura issensc haften dei tat für Biologie der Eberhard -Karls-Unn ersität Tübingen. - Herrn Dr. (ii kiiakd Mickoi in (Tübingen) zum 65. ( leburtstag gewidmet. Gedruckt mit finanzieller Unterstützung der Firma Cari 7.\ iss, Jena. download Biodiversity Heritage Library, http://www.biodiversitylibrary.org/ 2 STUTTGARTER BEITRÄGE ZUR NATURKUNDE Ser. A, Nr. 537 tion of the aedeagus during evolution, which is described for the first time in Scarabaeoidea. Male and female ectodermal genitalia possess extensive correspondence in musculature, which is described for the first time in Pterygota. The term homology cannot be used for corre- sponding structures between the sexes. By myological comparison of both sexes the following hypotheses are inferred: 1. Aedeagus and vaginal palps are derivatives of different urites. 2. Spiculum gastrale and vaginal palps are derivatives of urite IX. 3. The caudal part of Spiculum gastrale is of sternal or pleural origin. 4. The aedeagus is derived from urite X (or from inter- segmental area IX/X). 5. The bursa copulatrix is derived from urite x (or from intersegmental area ix/x). 6. The female genital tract has no structures corresponding to the aedeagus. Keywords: Coleoptera, Scarabaeidae, functional morphology, homology, copulatory apparatus, cryptic female choice. Zusammenfassung Sklerite, Membranen und Muskeln des männlichen und weiblichen Kopulationsapparates des Maikäfers Melolontha melolontha (Linnaeus, 1758) (Insecta: Coleoptera: Scarabaeidae: Melolonthinae) werden in Ruhe und in Copula beschrieben. Erstmalig wird der Ablauf der Kopulation bei Melolontha vollständig dargestellt. Aus dem Vergleich zwischen Ruhelage, Kopulationspositionen und mutmaßlicher Ovipositionsstellung werden die Funktionen der einzelnen Elemente der Kopulationsorgane erschlossen. In einem funktionsmorphologischen Szenario wird das Funktionieren der Kopula geschildert. Eine mögliche Strategie des Weib- chens, die Kopulation vor der Insemination zu beenden, wird beschrieben (cryptic female choice). Der Aedoeagus liegt in Ruhe auf der Seite („deversement"). Der Verlauf von Ductus ejaculatorius und Tracheen spricht für ein „retournement", eine Längsrotation des Aedoeagus um 180° während der Evolution, was erstmalig für die Scarabaeoidea festgestellt wird. Die männlichen und weiblichen ektodermalen Genitalien weisen eine entsprechende Muskelaus- stattung auf, was bei Pterygota erstmalig festgestellt wird. Der Terminus Homologie kann für Entsprechungen zwischen den beiden Geschlechtern einer Art nicht angewandt werden. Aus dem myologischen Vergleich beider Geschlechter werden folgende Hypothesen abgeleitet: 1. Aedoeagus und Vaginalpalpen sind Derivate unterschiedlicher Urite. 2. Spiculum gastrale und Vaginalpalpen sind Derivate des Urits IX. 3. Der caudale Teil des Spiculum gastrale ist sterna- 1 len oder pleuralen Ursprungs. 4. Der Aedoeagus ist ein Derivat des Urits X ) (oder des Inter- segmentalbereichs IX/X). 5. Die Bursa copulatrix ist ein Derivat des Urits x (oder des Inter- segmentalbereichs ix/x). 6. Der weibliche Genitaltrakt besitzt keine dem Aedoeagus entspre- chenden Sklerite. Inhalt 1 Einführung 3 2. Material, Methoden und Abkürzungen 5 2.1. Material 5 2.2. Methoden 6 2.3. Abkürzungen 6 3. Terminologie 7 3.1. Myologie 7 3.2. Sklerite und Membranen 8 4. Ablauf der Kopulation 9 4.1. Ablauf und Dauer der Kopulation bei Melolontha melolontha 9 4.2. Kopulationsvorgang bei nahe verwandten Arten 11 ') Als Urite werden im von KfiLERSchen (1963) Sinne die vollständigen Abdominalsegmente bezeichnet und nicht nur deren Ventralbereich. Die Urite werden mit römischen Ordinal- zahlen bezeichnet, und zwar die der männlichen Urite mit großen (VIII), die der weibli- chen Urite mit kleinen Lettern (viii). Sind die Urite beider Geschlechter gemeint, werden große Lettern verwendet. ... download Biodiversity Heritage Library, http://www.biodiversitylibrary.org/ KRELL, KOPULATIONSORGANE VON MELOLONTHA MELOLONTHA 3 5. Oviposition 12 5.1. Oviposition bei Melolontha melolontha 12 5.2. Oviposition bei nahe verwandten Arten 13 6. Morphologie des männlichen Kopulationsapparates von Melolontha melolontha (Sklerite, Membranen und Drüsen) 14 6.1. Urite VII und VIII 14 6.1.1. Third connecting mambrane 14 6.2. Urit IX 15 6.2.1. Spiculum gastrale mit Treffpunkt 1 16 6.2.2. Second connecting membrane mit Treffpunkt 2 16 6.3. Aedoeagus - „retournement" und „deversement" 16 6.3.1. Phallobasis und Phallobasalapodem 21 6.3.2. Parameren 22 6.3.3. First connecting membrane 22 6.3.4. Der Endophallus und seine Sklerite 22 7. Morphologie des weiblichen Kopulationsapparates von Melolontha melolontha (Sklerite, Membranen und Drüsen) 23 7. 1 Vagina und Cloaca mit Treffpunkten 1 und 2 24 7.2. Glandulae accessoriae 24 7.3. Palpi vaginales 26 7.4. Bursa copulatrix 26 7.5. Receptaculum seminis 27 7.6. Glandula receptaculi 27 8. Genitalmuskulatur von Melolontha melolontha 28 8. 1 Sollen entsprechende Muskeln des männlichen und weiblichen Genitalapparates als homolog bezeichnet werden? 28 8.2. Muskulatur des Urits VIII 30 8.3. Muskulatur der männlichen ektodermalen Genitalien 31 8.3.1. Extrinsische Genitalmuskulatur 31 8.3.2. Intrinsische Genitalmuskulatur 35 8.4. Muskulatur der weiblichen ektodermalen Genitalien 41 9. Funktionsmorphologisches Szenario von Kopulation und Oviposition 48 9. 1 Kopulation 49 9.2. Oviposition 76 10. Wie verhindert das Weibchen die Insemination? Ein Beitrag zur „Female-choice- Hypothese" 79 11. Entsprechungen im männlichen und weiblichen Kopulationsapparat und deren phylogenetische Interpretation S 1 12. Danksagung 91 l 13. Literatur »l 1. Einführung Im Jahre 1848 entdeckte der französische Apotheker Pi pro Orman< EY, daß der männliche Kopulationsapparat der Käfer im allgemeinen artspezifisch ausgeprägt ist. Ein Jahr darauf publizierte er eine ausführliche Studie /ti diesem Thema (Or- manci.y 1849). Nachdem die Genitalmorphologie in Jen darauffolgenden Jahrzehn- ten nur sehr vereinzelt in der coleopterologischen Forschung berücksichtigt wurde, ist es vor allem l\c\' [nsistenz des Berliner Professors Gustav K.raatz zu verdanken, daß die taxonomische Bedeutung der Genitalmorphologie publik gemacht winde (zum Beispiel Kraatz 1881; Smith 1889: 483f). Doch erst nach dem Erscheinen der umfangreichen Monographie von Sharp cv Mi ir (1912) winde die Genitalunter- suchung außerhalb des deutschen Sprachraumes /ur gängigen taxonomischen Me thode (Nichols 1986). download Biodiversity Heritage Library, http://www.biodiversitylibrary.org/ 4 STUTTGARTER BEITRÄGE ZUR NATURKUNDE Ser. A, Nr. 537 Bis heute werden die äußeren Genitalien zur Diagnostizierung von Biospecies be- vorzugt verwandt, da in den Kopulationsorganen anscheinend unmittelbar ein in- terspezifischer Isolationsmechanismus manifestiert ist: das mechanische Schlüssel- Schloß-Prinzip (lock-key), das Leon Dufour im Jahre 1844 (p. 253) in seinen Unter- suchungen über die Kopulationsorgane der Diptera formuliert hatte. Wie jedoch be- reits Jeannel (1941: 107) erkannte, ist gerade bei Käfern ein mechanisches Schlüssel- Schloß-Prinzip aus konstruktiven Gründen ausgeschlossen. Hier ist zwar der Schlüssel, der männliche Aedoeagus, ausgesprochen vielgestaltig; das weibliche Ge- genstück, die Vagina, weist hingegen keine Sklerite auf, die einem falschen Schlüssel in irgendeiner Weise passiven Widerstand leisten könnten [cf. Eberhard 1992: 1780 („because there is simply no female 'lock,'")]. Aktiver Widerstand ist hingegen mög- lich. Mehrfach konnte ich beobachten, daß ein Weibchen den bereits penetrierten Aedoeagus eines artgleichen Männchens nicht akzeptierte und die Kopulation been- dete, was die „female-choice-hypothesis" (Eberhard 1985) unterstützt. Diese Hypo- these, nach der das Weibchen durch Wahlverhalten sexuelle Selektion bewirkt, er- klärt nach Eberhard (1985) die hohe interspezifische Variabilität der männlichen Ge- nitalien im Tierreich.
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