Fauna E Flora Del G Lfo Di Napoli
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FAUNA E FLORA DEL G LFO DI NAPOLI Anthomedusae/Athecatae (Hydrozoa, Cnidaria) of the Mediterranean PART I CAPITATA BY ANITA BRINCKMANN - VOSS with 11 colour - plates drawn by ILONARICHTER With contribution from the Office of Naval Research (NONR 2100) and the National Science Foundation, U.S.A. EDIZIONE DELIA STAZIONE ZOOLOGICA DI NAPOLI CONTENTS AND SYNOPSIS OF FAMILIES AND GENERA Preface . pag. 1 Introduction . )) 3 Pictorial Key . È 8 Family Corymorphidae . .a 11 genus Corymorpha . È 11 genus Vannuccia . È 14 genus Euphysa . È 16 genus Eucodoniz~m . È 19 genus Paragotoea . 20 genus Branchiocerianthus . 2 22 Family Tubulariidae . 22 genus Ectopleura . ) 22 genus Tubularia . 28 genus Rhysomedusa . 32 Family Velellidae . 33 genus Velella . 34 genus Porpita . 38 Family Pennariidae . 40 genus Pennaria . È 40 Family Acaulidae . 44 genus Acaulis . 44 Family Corynidae . ) 46 genus Coryne . 47 genus Sarsia . 3 57 Family Cladocorynidae . : 69 genus Cladocoryne . 69 Family Zancleidae . , . ) 71 genus Zanclea . 72 Family Cladonematidae . 75 genus Cladonema . 76 Family Eleutheriidae . 78 genus Eleutheria , . 79 genus Staurocladia , . 81 Family Paracorynidae . 84 genus Paracoryne , . 84 Family Halocorynidae . > 84 genus Halocoryne . 84 Family Ptilocodiidae . > 85 genus Thecocodium . 85 PREFACE This monograph was first planned by the Naples Zoological Station in 1957. My own interest in Hydrozoa was awakened during student excursions under the guidance of Prof. A. PORTMANN(University of Basel, Swit- zerland) in Villefranche and Banyuls from 1953 to 1958. Finding that literature on this region was extre- mely poor and scattered, I started to think of writing a comprehensive report on A-nthomedusae-Athecatae of the Mediterranean, even as a student. I worked on this monograph in Naples from 1958 to 1963 continu- ously (BRINCKMANN,1959-65 ; BRINCKMANN-VOSS,1966-67). The five years in Naples were spent collecting live specimens and observing them in the laboratory. From 1963 on I worked on the morphology of the pre- served specimens and completion of the monograph in Winnipeg. The geographical area of this study comprises the Western Mediterranean, including the Adriatic Sea (NEPPI, 1912-22). Almost nothing is known from the Mediterranean east of the Adriatic Sea. In 1967 I was given the opportunity to look through the athecate hydroid collection of the Agassiz Mu- seum at Harvard University. The specimens there were mostly from collections made by ALEXANDERand LOUISAGASSIZ and A. G. MAYERand comparison of this material with the Mediterranean forms was most valuable. Discussions with and letters from the late Dr. W. J. RJZES,as well as his papers, were tremendously helpful (REES, 1936-66). After the appearance of the Synopsis of Medusae of the World by P. L. KRAMP in 1961, the work on the medusae of this monograph proceeded much faster. KRAMP'Sgeneric and specific de- finitions are often used in this paper (KRAMP,1911-1968). Although it does not deal with Mediterranean species, RUSSELL'SMonograph on the Medusae of the British Isles (1953) strongly influenced this public- ation, and his descriptions - in cases where the Northern and Mediterranean species are the same - are often cited, because they are so accurate. The order Athecatae-Anthomedusae should have been completed in a single volume. The plates do in fact include species of the whole order. But the detailed study of each species was so much more time-consum- ing than anticipated that it was decided by the author and publishers to print the suborder Capitata to- gether with the plates for Capitata and Filifera first and then, in a second publication, the text and text-figs. of the Filifera. (The order Anthomedusae-Athecatae was split into two suborders Capitata and Filifera by KUHN,1913. This division is actually only of value for the hydroid stage and I suppose it will be difficult to uphold the two suborders in the future; see REES, ed. 1966, p. 396). So while the division of Capitata and Filifera is maintained in this monograph, it is done for practical reasons only. The plates and text-figs. 6, 11, 43, 54, 59, 60, 66,79, 82, 83, 87 89 were drawn by Miss ILONARICHTER under the supervision of the author. All the drawings by Miss RICHTERwere done from living specimens. Dr. M. YA- MADA (Hokkaido University, Japan) kindly gave me the permission to publish text-fig. 26 of Ectopleura la- rynx. Dr. K. PETERSEN(Zoologisk Museum, Kopenhagen, Denmark) did the illustration of a living colony of Ptilocodium repens, text-fig. 100. Unless stated otherwise, all other text-figs. are my original drawings. Numerous are the people who helped me in the course of this work. Most of all I am deeply indebted to my former technician, MARIA-SOFIAGIAQUINTO-BOAG (formerly Naples, now Edmonton) for her skill and patience in keeping my hydroids and medusae alive under laboratory conditions. Sincerest thanks are due to Dr. P. DOIIRN,director of the Naples Zoological Station for his continuous support and interest in this monograph. I am very grateful to Dr. E. W, STRINGAM,Head of the Department of Animal Science, Uni- versity of Manitoba and Prof. G. HODGSONfor the generous space and facilities I was given at their institute for the final work on this monograph. A monographic paper involves a tremendous work on literature, especially on old and remote papers. My thanks go to the efficient staff of the interlibrary loan department of the University of Manitoba for their help. Sincere thanks are due to specialists who sent me comparative material: Dr. C. EDWARDS(Millport), Dr. 3. GOY (Villefranche), Dr. U. MONCHARMONT(Naples), the late Dr. W. J. REES(London), Dr. B. WERNER(Harn- burg). Many thanks go to Dr. G. MACKIE(University of Alberta, Canada), Prof. Dr. A. PORTMANN(University of Basel, Switzerland), Sir F. S. RUSSELL(Plymouth, England), and Dr. MARTHAVANNUCCI (Institute of Oceanography, University of Sao Paulo, Brazil) for reading the manuscript. Editing a monographic paper is a troublesome task. It was done by Dr. R. MARTIN (Head of the Zoology Dept., Naples Zoological Station) to whom I owe many thanks for this. e Writing this paper would not have been possible without a generous grant from the Office of Naval Res- earch and the U.S.A. National Science Foundation. The Office of Naval Research grant was given to the Naples Zoological Station to cover the various expenses connected with this monograph as there are pay- ments for the author, technicians, an artist, divers a.s.0. The National Science Foundation grant was given to me for morphological and histological research on the species, drawings and finishing of the monograph after I left Naples. The National Science Foundation grant was sponsored by the Museum of Comparative Zoology at Harvard University. I thank both Foundations for their trust and generous support. I would like to thank also Dr. B. NEWMAN(formerly Museum of Comparative Zoology, Harvard) and Dr. H. LEVI (Museum of Comparative Zoology, Harvard) for their help and advice. Last not least I would like to thank my husband, Dr. G. Voss, for criticizing this paper sentence by sentence, and who helped to clarify many unintelligible passages. After I had finished this part of the monograph I found that there were so many unsolved problems in numerous species that I would have liked to start investigations again without publishing anything at the moment. Different reasons compel me to publish now, but I am aware of the incompleteness of some chap- ters. Therefore my hope is, that through this monograph there will be more interest in Mediterranean Hyd- rozoa, especially in the life-cycles of Antho- and Leptomedusae. Winnipeg, Institute of Animal Science, University of Manitoba, Canada April 1968 INTRODUCTION Morphology. The morphology of medusae and hyd- such as orders and suborders. To illustrate this point, roids is described in numerous text books. See RUSSELL the classifications from some of the modern textbooks (1953) for all details on medusan morphology, HYMAN are listed here: (1948), KAESTNER(1967), NAUMOV(1960-64) and BOUILLON (1959-67). Unfortunately, the terminology of the ana- tomy of hydroids and medusae varies from one text- Order Hydroida book to another. Therefore text-fig. 1 shows an antho- Suborder Gymnoblastea medusa and three types of hydroids with an explanation Suborder Calyptoblastea of the terminology of structural details as used in this Order Milleporina paper. Order Stylasterina The different types of tentacles require mention be- Order Trachylina cause they are used as taxonomic criteria. Moniliform Suborder Trachymedusae tentacles: cnidocysts arranged in several clusters (text- Suborder Narcomedusae fig. I, h); capitate tentacles: cnidocysts arranged in a Order Siphonophora cluster at the tip of the tentacle (text-fig. 1, p); fili- Suborder Calycophora form tentacles: cnidocysts scattered along the whole Suborder Physophorida length of the tentacle (text-fig. 1, 0); There are transi- tional stages between these three types as will be noted in the detailed descriptions. Class Hydrozoa The taxonomic value of cnidocysts has often been Order Calyptoblastea discussed in recent years (WEILL,1931-1935; WERNER, Order Gymnoblastea 1965; WESTFALL,1965, 1966 a, h). An excellent review Order Hydroida and critical judgment of the different opinions is given Order Trachylina by VERVOORT(1966 b, p. 389-394). Types and sizes of Order Hydrocorallina cnidocysts vary very little from species to species and Order Siphonophora are therefore of little help if any in identifying the species (VANXUCCI,1959); but cnidocysts may be quite important in establishing relationships between fami- lies (REES, 1957 a, d, 1958). Yet, contrary to PICARD Class Hydrozoa (1955 c, 1957, 1958 a), I maintain that cnidocysts must Order Hydroida not be used to the exclusion of every other character in Suborder Anthomedusae the establishment of families or their relation to one Suborder Leptomedusae another. Suborder Limnomedusae Order Milleporina Evolution.