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Systematics of the Euretidae (Porifera: Hexactnellida: Hexactinosa)

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Systematics of the Euretidae (Porifera: Hexactnellida: Hexactinosa) SYSTEMATICS OF THE EURETIDAE (PORIFERA: HEXACTNELLIDA: HEXACTINOSA) Benjamin Wheeler Redpath Museum and the Department of Biology McGi 11 University, Montreal February , 1 999 "A rhesis szrbmilted to the Facuhy of Graduute S~udiesand Reseurch in parliai fui$llmen( cf rhe reqtrirernents of the degree of Master of Science" 0 Wheeler, 1999 National Library Bibliothèque nationale du Canada Acquisitions and Acquisitions et Bibliographie Services services bibliographiques 395 Wellington Street 395. nn, Wdlingtoci CMawaON K1AW ChtawaON K1AW Canada CaMde The author has granted a non- L'auteur a accordé une Licence non exclusive Licence aiiowing the exclusive permettant à la National Library of Canada to Bibliothèque nationale du Canada de reproduce, loan, disûibute or seil reproduire, prêter, disîribuer ou copies of this thesis in microfom, vendre des copies de cette thèse sous paper or electronic formats. la forme de microfiche/fh, de reproduction sur papier ou sur format électronique. The author retins ownership of the L'auteur conserve la propriété du copyright in this thesis. Neither the droit d'auteur qui protège cette thèse. thesis nor substantiat extracts fkom it Ni la thèse ni des extraits substantiels may be printed or otheMiise de celle-ci ne doivent être imprimés reproduced without the author's ou autrement reproduits sans son permission. autorisation. Table of contents TABLEOF CONTENTS ......................................................................................................... ,7 ABSTRACT........................................................................................................................ 3 RESUME............................................................................................................................ 3 ACKNOWLEDGMENTS........................................................................................................ 4 INTRODUCTION ........................................ .. ................................................................... 5 LITERATUREREVIEW ............................... ,... ........ .... 7 METHODS........................................................................................................................ 25 SYSTEMATICS............................................................................................. 28 KEYTO GENERA............................................................................................................. 34 Subfamily Euretinae ZIITEL ................................................................................... 35 EURETE SEMPER, 1868 ........................................... ... .............................. 35 CALY PTORETE OKADA, 1 925 .................................................................. 42 CONOFtETE IJIMA, 1927 ............................................................................... 47 BATHYXIPHUS SCHULZE, 1899 ................................................................. 54 PARARETE IJIMA, 1 927 ............................................................................... -58 PLEUROCHORIUM SCHRAMMEN, 191 2 ................................................ 64 VERRUCOCOELOIDEA REID, 1969 ........................................................... 68 -. Subfamily Iphitinae (n. sf.)................................ .. ............................................... 12 IPHITEON BO WERBANK, 1869A ................................................................ 72 CHONELASMA SCHULZE. 1887~............................................................... 79 LEFROYELLA THOMSON. 1 877 ................................................................. 85 MYLIUSIA GRAY . 1 859 .............................................................................. -92 PERIPHRAGELLA MARSHALL. 1875 ...................................................... 100 PTYCHODESIA SCHRAMMEN, 19 12 ....................................................... 105 .i Heterorete-group "..... ...................................................................................... 111 HETERORETE DENDY . 19 1 6 ..................................................................... 111 GYMNORETE IJIMA, 1927 ........................................................................ iI5 ENDORETE TOPSENT, 1928....................................................................... 120 Drscljssro~.................................................................................................................. 124 GEOGRAPHICDISTRIBUTION .......................................................................................... 134 CONCLUSIONS............................................................................................................... 134 REFERENCES................................................................................................................ 136 APPENDIXA .................................................................................................................. 144 Abstract A long overdue taxonomic revision is performed on the family Euretidae (Hexactineltida). World Iiterature is compiled, world type material is gathered, prepared, re-examined, measured. described and illustrated, tauonomic histories and world species distributions are provided. A generic key to the Euretidae is also provided. Of the 19 generic names associated with the Euretidae at one time or another, 16 are recognised here (Buthpiphus. Calyprorere. Chonciusma. Cortorere. Endarete, Eurece, Gymnorere. Heterorete, iphiteon, Myliwia. Pararele. Periphrugelfa, P~ezrrochorirrm,P~chodesia. Lefioyella and Verrucocoeloidea). Syringidium is put in to sy nonymy with Lefioyellq Joanella and MarguritelZa are synonym ized w itli /phireon: two subspecies and one form are elevated to species sbtus; three new species combinations are presented and two new species are named. Phylogenetic analysis suggests the majority of morphological euretid characters are pi-one to homoplasy and are not useful for phylogenetic classitication. Subfamilies based on channelization and the presence or absence of key spicules are designated. Résumé Une révision taxonomique longuement attendue a été esécutée sur la famille Euretidae (Hesactinellida). Mondialement. la littérature est compilée. le matériel de type du monde est recueil1 i. préparé, esam iné de nouveau. mesuré, décrit, et illustrées; les histoires ta,,onomiques et les distributions des espèces du monde sont fournies. Une clée générique de la famille I'Euretidae est également fournie. Des 19 noms génériques associés à I'Euretidae en même temps ou encore. 16 sont reconnus ici (Bathyxiphlrs, Culyprorere, Chonelasma. Conorete. Endorele. Ezrt-ere, Gynrnorefe.Heterorete, @hiteon, Mvliusia. Pararete. Periphragella, Plcurochorirrrn. Piychodesia. Lefioyella et Vernrcocoeloidea).Syrirlgidizm est m is dans la synon y m ie avec L~'fioyclZu;Joanella et Margaritella sont mis dans synonymie avec /phireon; deux sous-espèces et une forme sont élevées au status d'espèce; trois nouvelles combinaisons d'espèce sont présentées et deus nouvelles espèces sont nommées. L'analyse phylogénétique suggère que la majorité des caractères morphologiques des euretids soient sujets a I'homoplasy et ne sont donc pas utile pour la c!assification phylogénétique. Sous-familles basées sur le channelization et la présence ou absence des spicules principaux sont indiqués. Ackno wledgments Henry Reiswig provided me not only with access to his extensive specimen and iiterature collections, but also to his valuable time and patience (especially with the synonomies!). Without his positive attitude, encouragement and support this study would not have been possible. I'd like to acknowledge Dr. G. Bell and Dr. D. Green for their participation in this project and the "Biodiversity lab" for the use of their advanced technology and positive atmosphere. Ttianks also goes to Jason for his helpful cladistic input. 1 would also Iike to acknowledge the BMNH. the Leiden museum. Musée de Monaco and MC2 for their valuable specimen loans. For financial support 1 would like to recognize NSERC and McGill University. Ingrid, if I could only express how much your help, happiness and reassurances have contnbuted to this project! Thanks for making my Master's a wonderful experience. I'd also like to extend my gratitude to the Anand family for their reassuring support and nurnerous nourishing "take-home" meals! ! Harry. your bright positivity canied me throughout this project. What more cm 1 say but THANKS!! Finally, I'd Iike to thank Chris and Susan, for without whom. I'd no longer have my best friend (Hi Bau-xi!) and would still be somewhere in India!! I'd like to dedicate this thesis to my parents whose example and support defies imagination and great distances. ln troduction Hexactinellids, or glass-sponges, are perhaps one of the more poorly understood and unrecognised marine invertebrate groups. Due to thsir remote abyssal or bathyal habitat, little is known about modes of reproduction. development, diversity, geographic distri bution and evolutionary relationships. Perhaps more practically, hexactinellids have remained obscure due to their unstable taxonomy. which, in turn. has complicated the identification of newly collected material. The majority (approx. 50%) of hexactinellid genera are based upon single. rarely complete. specimens crudely collected and described ofien over one hundred years ago. In fact. nearly 70% of hexactinellid specimens recorded in the literature were descnbed before 1930. To fùrther aggravate matters. historicall y, hexactinellids
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