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Cereus Pedunculatus Und Actinia Equina (Anthozoa, Actiniaria)

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Cereus Pedunculatus Und Actinia Equina (Anthozoa, Actiniaria) I:tELGOL~,NDER ME ERESUNTERSUCI-iUNGEN Helgol~nder Meeresunters. 34, 451-461 (1981) Fortpflanzung und Sexualit~it von Cereus pedunculatus und Actinia equina (Anthozoa, Actiniaria) W. Sch~fer Institut ffir Zoologie (Lehrstuhl I) der Universita't Erlangen - Nfirnberg; D-8520 Erlangen, Bundesrepubtik Deutschland ABSTRACT: Reproduction and sexuality of Cereus pedunculatus and Actinia equina (Anthozoa, Actiniaria). Sexuality and reproductive behaviour of Cereus pedunculatus (Pennant) and several forms (subspecies) of Actinia equina (L.) from populations collected along the French Atlantic Sea coast and in different habitats along the European Mediterranean coast were studied. At the stage of 96 septae C. pedunculatus and A. e. atlantica II exhibited mature oocytes which developed parthenogenetically into larvae. The latter appeared simultaneously in the gastrocoele. Adolescent A. e. atlantica II developed very few mature oocytes and larvae. Following a sterile period, oocytes and young individuals of different age groups were present almost throughout the whole year in adult anemones. A. e. mediterranea I was dioecious and oviparous in any habitat observed. Samples of the larviparous A. e. mediterranea II (collected near Banyuls, France) exhibited male gonads exclusively and contained larvae. Spontaneous longitudinal fission was occasionally observed in adult A. e. mediterranea I and adolescent A. e. atlantica II. EINLEITUNG Die Befunde /ilterer Autoren tiber die Fortpflanzungsbiologie von Cereus peduncu- tatus und Actinia equina, insbesondere iiber Sexualit~t und Fortpflanzungsrnodi beider Arten, kbnnen oft nicht in Einklang rniteinander gebracht werden (vgl. Lacaze Duthiers, 1872; Graeffe, 1884; Lo Bianco, 1909; Stevenson, 1935; Leloup, 1952). Neuere verglei- chende Untersuchungen lassen fiir beide Species eine Aufspaltung in mehrere Unterar- ten, gegebenenfalls Formen (Actinia), notwendig erscheinen (Schrnidt, 1971; Schmidt, 1972a, b; Rossi & Calenda, 1974; Rossi, 1975; Carter & Funnell, 1980). Hierbei hat sich gezeigt, daf~ die geographisch unterschiedlich verbreiteten Subspecies bzw. Formen unter anderem stark im Fortpflanzungsverhalten variieren, wobei viele Fragen, insbe- sondere fiber die Sexualit/it larviparer Formen, noch nicht zufriedenstellend gekl~irt werden konnten (Chia & Rostron, 1970; Cain, 1974; Black & Johnson, 1979; Larkrnan, 1980), Die unterschiedlichen Beobachtungen und die vielen noch offenen Probleme erfor- dern weitere ausffihrliche Untersuchungen der Fortpflanzungsbiologie von Actinia equina und Cereus pedunculatus, wobei in der vorliegenden Arbeit grundlegende Fragestellungen wie Geschlechtsreife und Fortpflanzungszyklen einerseits sowie Sexualit~it und Fortpflanzungsmodi andererseits untersucht werden. © Biologische Anstalt Helgoland 0174-3597/81/0034/0451/$ 02,00 452 W. Sch6fer MATERIAL UND METHODE Zur Untersuchung gelangten lebende Tiere aus Aquarienpopulationen (franz6si- sche Atlantikkiiste) und von mehreren Fundorten der europ6ischen Mittelmeerk(iste (Costa Brava (iber C6te Vermeille und C6te d'Azur bis italienische Riviera, Korsika, jugoslawische Adria, Golf von Korinth). Im Rahmen dieser Arbeit konnten nicht alle bisher bekannten Subspecies untersucht werden, weshalb sich die vorliegende Studie zun~chst auf Actinia equina mediterranea I und II, Actinia equina atlantica II (zur Taxonomie des Genus Actinia vgl. Schmidt, 1971) sowie auf eine Aquarienpopulation yon Cereus pedunculatus beschr6nkt. Fiir histologische Untersuchungen wurden ein- zetne Individuen in einem Medium aus Seewasser und MgC12-L6sung (7,5 %) bet6ubt und mit Formol/Seewasser fixiert. Nach dem Entw~ssern (Alkoholreihe) wurden die Proben fiber Methylbenzoat in Paraplast eingebettet. Die 5-10 [tm dicken Schnitte wurden nacheinander mit Alcianblau/-griin (3:2) und darauf nach Schmidt (1970) mit Mayer's Haemalaun, Eosin/Orange G und Anilinblau gef6rbt Zur Kontrolle des Fortplanzungsmodus wurden Larven yon Actinia equina atIantica II und Cereus pedunculatus aus den Gastrocoelien adulter Tiere entnommen und einzeln in einer Batterie yon Zuchtbeh61tem aufgezogen. Das stets frisch angesetzte Seewasser wurde mehrmals w6chentlich gewechselt Die Fiitterung der Tiere erfolgte mit Artemia-Nauplien und Muschelfleisch. ERGEBNISSE UND DISKUSSION Geschlechtsreife An Hand histologischer Pr6parate wurden heranwachsende Individuen auf das Vorhandensein von Geschlechtsprodukten untersucht Bei Cereus pedunculatus waren bis zum 48-Septenstadium keine Gameten nachzuweisen, jedoch ist unmittelbar vor dem AbschluB des 3. Septenzyklus zwischen Retraktor und Filament eine breitere Zone mit undifferenzierten Entodermzellen zu beobachten. In Tieren, die das 48-Septensta- dium gerade ~iberschritten haben, werden erstmats Keimzellen angelegt. Es handelte sich ausschlieBlich um Oocyten (Abb. 3a). Vollreife Eizellen und erste Larven treten erst nach AbschluB des 4. Septenzyklus auf. Bei Actinia equina atlantica II ist die entscheidende Phase offensichtlich unmittel- bar vor der Vollendung des 4. Septenzyklus erreicht, das jiingste Tier mit Oocyten hatte 86, das jiingste mit Larven 96 Septen. Demgemafl waren in den folgenden Entwick- lungsstadien Jungtiere im Gastrocoel, jedoch keine Oocyten mehr nachzuweisen, da nicht in allen potentiell fertilen Septen Keimzellen angelegt wurden. Erst nach Ablauf einer sterilen Periode fanden sich in adulten bzw. fast adulten Tieren regelm6Big mehr E P[onuto. 4B 96 130 Ad ultus O L Abb, 1. Entwicklungsstadium von Actinia equina atlantica II (E; Zahlen = Septenstadium) in Relation zum Auftreten von Oocyten (O) und Larven (L) im Gastrocoel Cereus pedunculatus und Actinia equina 453 oder weniger zahlreiche Oocyten und Larven. (Einzelheiten fiber die Oogenese siehe Sch~fer & Schmidt, 1980, sowie Schmidt & Sch~fer, 1980). Die vorliegenden Befunde k6nnen zusammenfassend nur so gedeutet werden, dab Actinia equina atlantica II folgende Fortpflanzungsphasen durchmacht (Abb. 1): (a) Das heranwachsende Tier (ca. 96-Septenstadium) entwickelt nur wenig reife Keimzellen und entsprechend wenig Larven 14-8). (b) Es fotgt eine mehrmonatige sterile Periode. (c) Nach deren Ablauf treten schlieBlich tiber das ganze Jahr hinweg Oocyten und Jung- tiere auf. Beziiglich ihrer Geschlechtsreife weisen Cereus pedunculatus und Actinia equina charakteristische Unterschiede auf. Als MaB dafiir, wie friihzeitig eine Art geschlechts- reif wird, kann der Quotient aus SeptenzahI im adulten Zustand und Septenzahl beim Auftreten der ersten reifen Geschlechtsprodukte dienen; je h6her dieser ausf~llt, um so zeitiger wird eine Art geschlechtsreif. Er betr~gt bei Cereus 768 : 96 ~ 8, bei Actinia auf Grund der Unvollz~hligkeit des letzten Zyklus ca. 180 : 96 -- ca. 1,88. Cereus wird also relativ friih, Actinia retativ sp~it geschlechtsreif. Bei Actinia equina atlantica II ist bemerkenswert, dab im ersten Jahr der Gesehlechtsreife nach meinen Beobachtungen lediglich in einem einzigen der potentiell fertilen Septen (nur die Richtungssepten sind steril) Oocyten angelegt werden. Dies hat zur Folge, dab in fast adulten Individuen Jungtiere, aber keine Keimzellen nachweisbar sind. So fanden Gashout & Ormond (1979) und Carter & Funnell (1980) einige Exemplare von Actinia equina (bei der untersuchten Unterart handelt es sich mit grol3er Wahrscheinlichkeit um Actinia equina equina I [Schmidt, 1971]), die zwar Larven im Gastrocoel entwickelten, jedoch als "sterile" bzw. "non sexuell" beschrieben werden. Es kann nicht ausgeschlossen wer- den, dal~ bei jenen Tieren gerade das Stadium (siehe oben) vorlag, in dem zwar Larven aber keine Keimzellen beobachtet werden kbnnen. Dies wiirde alterdings voraussetzen, dab A. e. atlantica II und A. e. equina I beziiglich ihrer Geschlechtsreife eine ~hnliche Entwicklung durchlaufen (vgl. Abb. 1). Fortpflanzungszyklen Bei der Aquarienpopulation von Cereus pedunculatus unterliegt die Oogenese, das Auftreten der Larven und die Geburt der Jungtiere einem jahreszeitlichen Zyklus. Die vorliegenden Daten wurden in den Jahren 1976 und 1977 gewonnen. Nach Ablauf einer Fortpflanzungsperiode werden in den Herbstmonaten vereinzelt Keimzellen gebildet, die jedoch nicht zur Reife gelangen und zum Teil wieder aufgel6st, m6glicherweise resorbiert werden. Nach einer Erholungsphase werden bereits im Winter wieder neue Oocyten angelegt, die binnen zwei his drei Monaten reifen, so dab schon Ende M~rz bis Anfang April Larven nachgewiesen werden k6nnen. Letztere verbleiben vorerst in den Gastrocoelien der Muttertiere. In den Monaten April bis Juni reifen weitere Oocyten nach (Abb. 2). Im Friihsommer werden Jungtiere verschiedenen Alters gefunden. Letz- tere verlassen die Muttertiere vorwiegend im Juli und August, meist zur Zeit der Temperaturmaxima. Bereits im Friihherbst werden in der Regel keine reifen Gonaden mehr angetroffen, einzelne gr6Bere Oocyten zersetzen sich (Abb. 3b). Die Temperatur mit ihren jahreszeitlichen Schwankungen ist offensichtlich der wichtigste Faktor, der den Fortpflanzungszyklus yon Cereus steuert. Fiir das Wachstum und die Entwicklung der Oocyten werden nicht unbedingt konstant hohe, sondern 454 W. Sch~fer L | 0 o c 25- \ 20 /s/~ ~'x x \\ I -j x-.N "l¢:m Feb Mrz Apt Mai 3un 3ul Aug Sep Okt Nov Dez Abb. 2. Fortpflanzungsperiode von Cereus pedunculatus (Aquarienpopulation) in Relation zur Wassertemperatur (L: Auftreten yon Larven im Gastrocoel, O: Auftreten reifer Oocyten) vielmehr kontinuierlich steigende Temperaturen ben6tigt (Abb. 2). Dagegen scheinen stetig absinkende Temperaturen, auch wenn im Herbst noch relativ hohe Durchschnitts-
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