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Zur Morphologie Und Systematii( Deh Alloeocoela Cumulata Und Rhabdo- Coela Lecithophora (Turbellaria)

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Zur Morphologie Und Systematii( Deh Alloeocoela Cumulata Und Rhabdo- Coela Lecithophora (Turbellaria) ACTA ZOOLOGIeA FENNIC.\. 26 EDIDIT SOCIE1'AS PRO FAUNA ET FLORA FENNTeA ZUR MORPHOLOGIE UND SYSTEMATII( DEH ALLOEOCOELA CUMULATA UND RHABDO­ COELA LECITHOPHORA (TURBELLARIA) VON TOR G. KARLING ~IIT 20 ABBILDUNGEN IM TEXT UND 17 TAFELN 7.U~1 DRUCK I;;INGELIEFFRT A~I 11. APRIL 1939 HELSINGFORSIAE 1940 HELSINGFORS 1940 DnUCK VON A.-G. F. TILGMANN Inhaltsverzeichnis. Seite Vorwort 5 Ei.rlleitung ? Technisches ". .............. .. 12 FtUldorte und Material der tmterSllchten Tiere '1:~ I. Spezieller TeD 15 '1. Fam. Pseudostomidae ....................................... .15 A. Prolecithoplana lutheri Ii. gen., 11. sp. 15 B. Pseudostomum klostermanni (Graff); P. quadrioculatulll (Lcnck.) 26 :l. Falll. Cylindrostomidae 41 A.. Enterostomula dura (Fuhrm.) ......................... .. 41 13. Sc1eraulophorus cephalatus 11. gen., n. sp. ........... .. 55 3. Falll. Plagiostomidae 62 Acmostomum diokum (Meczn.); Plicastoma norvegicum n. sp.; Plagio- st01ll11m torquatu111 n. sp. 62 ,'1. Fam. Provorticidae ........................................ .. 98 Hangethellin calceifera n. ,~en., n. sp.; H. c. norvegica 11. subsp.; Pro- vortex balticus M. ScllUltze ,.......................... 98 ::;. Fanl. Solenopharyngidae 1'l!l Anthopharynx vaginatus 11. gen., 11. sp.; Trisaccopharynx westbladi n. gen., n. sp. t 1ft IJ. Allgemeiner Teil 134 1. Vergleichende Morphologie ................................ .. 13/i A. Körperform und Farbe 134 B. Deckepithel , 135 C. Basalmembran , 139 D. Hauteinschlüsse und HautdrÜsen , H1 E. 11uskulatur 14G F. Nervensystem 148 G. Augen " 159 H. Chemoreceptoren ...................................... .. '1 GO 1. Parenchym ,............................... 1G'l J. Verdauungsapparat 162 K. Protonephridien . 177 T(. Geschlechtsapparat .......................... .. 178 a. Hoden und Vasa deferentia ..................... .. '1 ') 8 b. Begattungsorgan 185 c. Weibliche Gonaden ,....................... 194 d. Weibliche Ausführwege , .. 202 'r Tor G. Karling, Zur Morphologie und Systematik der Turbellarien e. Begattung und weibliche Hilfsapparate 20 /1 f. Atrium und Genitalöffnungen 209 2. Phylogenie und Systematik 212 A. Die Verwandtschaftsbeziehungen in der Unterordnung Cumulata 212 B. Über die Abstammung der Lecithophora 218 C. Ein Beitrag zur Stammesgeschichte und. Klassifikation der Turbellaria .......................................... .. 223 D. Entwurf zu einem neuen System der Turbellaria 229 E. Übersicht Über die diagnostischen Merkmale der neuen Arten .. 235 Nachtrag , , 239 Literaturverzeic1mis 242 Erklärung der Abbildungen. .................. .. 250 Vorwort. Seit 1929 habe ich im Laufe von mehreren Sommern während kÜrzerer oder längerer Perioden die Turbellarienfauna - vornehmlich auf Sand­ boden -- in der Umgebung der Zoologischen Station Tvärminne studiert und dabei ein reichhaltiges Material von Formen aus verschiedenen Grup­ pen zusammengebracht. Nur einen geringen Teil der gefundenen Arten habe ich früher - teilweise auch nur vorläufig - beschrieben (vgl. Literatur­ verz.). In der vorliegenden Arbeit habe ich einige Formen von Tvärminne mit einigen Arten aus dem Material zusammengestellt, das ich anlässlich eines fünfwächigen Aufenthaltes im Sommer 1934 an der Biologischen Sta­ tion (des Museums von Bergen) auf Herdia gesammelt habe. Diesem Matedal sind zwei Arten beigefügt worden, die mir Herr Dr. EINAR WESTBLAD freund­ lichen;\'eise zur Bearbeitung überlassen hat. Das konservierte Material ist teils im Zoologischen Institut der Uni­ versität Helsingfors, teils während eines zweimonatigen Aufenthaltes im Herbst 1936 im Naturhistorischen Reichsmuseum in Stockholm bearbeitet worden. Es ist mir eine angenehme Pflicht, hiermit meinem geschätzten Lehrer, Herrn Prof. Dr. ALEX. LUTHER, meinen besten Dank auszusprechen für das grosse Interesse, das er meiner Arbeit entgegengebracht hat, für seine Bereitwilligkeit als Präfekt der Zoologischen Station Tvärminne mir einen Arbeitsplatz zur VerfÜgung zu stellen, und für die freundliche Erlaubnis, seine reichhaltige Spezialbibliothek benutzen zu dürfen. Einige Schnittprä• parate einer der behandelten Arten hat er mir gleichfalls in liebenswürdiger Weise Überlassen. Aufrichtiger Dank gebührt ferner dem Präfekten der Biologischen Station Herdla, Herrn Prof. Dr. AUGUST BRINKMANN und dem Amanuensis Herrn Dr. D. RusTAD sowie dem Vorstand der Evertebratab­ teilung des Naturhistorischen Reichsmuseums in Stockholm, Herrn Prof. Dr. SIXTEN BOCK, die mit Rat und Tat meine Arbeit in den genannten Instituten erleichtert haben. Ich benutze auch die Gelegenheit, meinen guten Freund und Kollegen Herrn Dr. EINAR WESTBLAD meiner Dank­ barkeit zu versichern nicht nur für das bereits erwähnte wertvolle Material, sondern auch fÜr die vielfach erwiesene Freundlichkeit während meines 6 Tor G. J{arZing, Zur JHorphologie und Systematik der Turbellarien Aufenthaltes in Stockholm und für den lehrreichen Gedankenaustausch Über Fragen aus unserem gemeinsamen Interessengebiet. Die Mikrophotogramme sind mit Sachkenntnis und Interesse von Fräu• lein DAGMAR BERG am Naturhistorischen Reichsmuseum in Stockholm ausgeführt worden, was ich mit Dankbarkeit erwähne. Meinen Dank möchte ich auch Herrn Dr. M. MEHLElVI aussprechen, der die sprachliche Korrektur meiner in deutscher Sprache verfassten Arbeit besorgte. Für die im Ausland betriebenen Studien habe ich Stipendien von der Studentenkorporation >}Nylands Natiolll> und vom Unterrichtsamt erhalten, für welche UnterstÜtzungen ich meinen ergebensten Dank zum Ausdruck bringen möchte. J. Einleitung. In GRAFFS Rhabdocoelidenmonographie (1882, p. 2) finden wir erstmals die Namen Rhabdocoela und Alloiocoela 1) als Bezeichnung fÜr gleichwertige Gruppen in der Unterordnung Rhabdocoelida. Als dritte Gruppe gehörte zu dieser Kategorie die Acoela. GRAFF fand es aber später für nötig (1904, p. 207-208), auf die UUANINsche Einteilung der Turbellarien in zwei Hauptgruppen Acoela und Coelata (1870) zurückzugreifen, wonach die Rhabdocoelida nur die Rhabdocoela und A lloeocoela umfassten. Die unter dem Namen Alloeocoela zusammengeführten Formen wurden schon von UUANIN unter dem Namen Monocelinea zusammengefasst (1870). Das Hauptmerkinal dieser Gruppe stellten die Hoden dar, die aus zahlreichen, kleinen, im ganzen Körper zerstreuten oder im Vorderkörper angehäuften Blasen bestehen sollen. Der von EHRENBERG stammende Name Rhabdocoela (1831) und die Be~en­ l1ung Alloeocoela wie auch die Übrigen schon ZU dieser Zeit verwendeten Namen der grösseren Turbellariengruppen liefern einen Ausdruck für die Gestaltung des Darmes, eine Eigenschaft, der schon früh grosser systemati­ scher Wert beigemessen wurde. Früh wurde aber auch schon der Wert der Gruppe Alloeocoela bezwei­ felt und HALLEZ, der mit GRAFF (1882, p. 207-209) die nahe Verwandt­ schaft dieser Gruppe mit den Tricladen betont, sagt aus, dass die Alloeocoela »remarquable comme groupe de passage» seien (1890, p. 44). Auch HOFSTEN konstatiert seinerseits die nahen Beziehungen zwischen den Tricladida und Alloeocoela wie auch die Unmöglichkeit die Gruppe Rhabdocoelida aufrechtzuerhalten, weil die Alloeocoela mehr mit den Tri­ cladida als mit den Rhabdocoela verwandt sind. Er teilt somit die coelaten Turbellarien in vier Ordnungen: Rhabdocoela, Alloeocoela, Tricladida und Polycladida (1907 a, p. 237). GRAFF Übernimmt einige Jahre später diese Einteilungsgrundlage (1912-1917, p. 2701, Fussnote). In der letzten, von BRESSLAU gegebenen Übersicht über das Turbella­ riensystem finden wir endlich die sechs Ordnungen: Acoela, Rhabdocoela, Alloeocoela, Tricladida, Polycladida und Temnocephalida (1928-1933, p. 53). 1) Der Name Alloiocoela wurde später von SPENGEL (1884, p. 183) in Alloeo­ coela abgeändert. 8 Tor G. ]{aI'ling, Zur l\Iorphologie und Systematik ucr Turbellarien GRAPF teilte ursprünglich die Alloeocoela in die Familien Plagiostomida und 111onotida ein. Die erstgenannte Familie wurde in vier Subfamilien, Acmostomina, Plagt'ostornina, Allostomina und Cylindrostomina untergeteilt (1882, p. 2). i. BÖHMIG folgt der Familien- und Subfamilieneinteilung GRAFFS inner­ halb der Alloeocoela; die Diagnosen werden aber für diese, wie für manche Gattungen und Arten, verändert oder neu aufgestellt (1890, p. 464-470). Als dritte Familie wird von VEJDOVSKY Bothrioplanidae den Alloeocoela zugeführt (1895, p. 200). Mit diesem Autor teilt HOFSTEN die letztgenannte Gruppe in drei Familien: Plagiostornidae, IVIonocelididae und Bothrioplanidae (1907 a, p. 237). GRAPF bildet aus diesen Familien die drei Sektionen Holo­ coela, Crossocoela und Cyclocoela. Die ehemaligen vier Subfamilien der Fami­ lie Plagiostomidae werden zu Familien erhöht und erhalten die Namen H alle­ ziidae, Plagiostomidae, Allostomatidae und Pseudostomidae (:1904-1908, p. 2010, Fussnote u. p. 2510-2511). HOFSTEN findet es schwierig, innerhalb der Alloeocoela die Gruppen Crossocoela und Cyclocoela gesondert aufrechtzuerhalten, weshalb er sämtliche zu diesen Gruppen gerechneten Formen zur gemeinsamen Gruppe Crossocoela zusammenführt und den Namen Cyclocoela fallen lässt (1918, p. 58-59). STEINBÖCK begründet die neue Sectio Typhlocoela für die Familie Pro­ rhynchidae (1924 a, p. 241) und später die Sectio H emacoela für die Gattung Hojstenia (1924 b, p. 165). Nachdem die Unhaltbarkeit des frÜheren, auf der Darmkonfiguration aufgebauten Systems ernent hervorgehoben worden war, wurde dem neuen System der Alloeocoela von REISINGER (1924 d, p. 147-148) und STEIN­ nÖCK ('1925, p. 166-169) der Bau der weiblichen Gonaden zugrunde gelegt. Es trat in den drei Sektionen (Unterordnungen nach STEINBÖCK)
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