The Taxonomy and Functional Anatomy of Southern African Echiurans
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THE TAXONOMY AND FUNCTIONAL ANATOMY OF SOUTHERN AFRICAN ECHIURANS by Ramlall Biseswar Submitted in fulfilment of the requirements r for the degree of Doctor of Philosophy in the Department of Zoology in the University of Cape Town. University of Cape Town Supervisor: Dr Jennifer A. Day Date submitted June, 1986 The University of Cape Town has been given tht right to reprnclt,ce this thesis in whole . f or in pMt. Co::-vn:.;hr is Lc:d by the author. :-... ----.-, ..... ~·-·*'"' ... ,, ..... -~·-· .... The copyright of this thesis vests in the author. No quotation from it or information derived from it is to be published without full acknowledgement of the source. The thesis is to be used for private study or non- commercial research purposes only. Published by the University of Cape Town (UCT) in terms of the non-exclusive license granted to UCT by the author. University of Cape Town PREFACE For the sake of convenience, this thesis is presented in two parts : the first part dealing with taxonomy and distribution and the second with functional anatomy. The thesis consists of published papers, manuscript papers and papers that have already been submitted for publi= cation. In addition, interlinking sections have also been included, especially in the taxonomic part of the thesis,in order to bring together all the information into a single comprehensive account. The following three papers have already been published: 1. Some aspects of the anatomy of Ochetostoma caudex (Echiura) from the east coast of southern Africa with remarks on its taxonomic status. S. Afr. J. Zool. 1983, 18: 47-55. 2 , A key to the species of Anelassorhynchus (Echiura) with a description of a new species from the east coast of southern Africa. S. Afr. J. Zool. 1984, 19: 16-21. 3. The geographic distribution of Echiura from southern Africa. S. Afr. J. mar. Sci. 3: 11-21. The manuscript entitled: "Ochetostoma (Echiura) from southern Africa with the description of a new species" has been submitted for publication in the Annals of the South African Museum. The following three manuscripts are in the process of being submitted for publication: 1. Thalassema (Echiura) from southern Africa with the description of a new species. South African Journal of Zoology. 2. Burrowing, locomotion and other movements of the echiuran Ochetostoma caudex. Journal of Zoology, London o 3, The reproductive system, gametogenesis and annual reproductive cycle of Ochetostoma caudex (Phylum Echiura). Acta &nbryologiae et Morphologiae Experimentalis. A glossary of anatomical terms which are important in the classification and descriptions of echiurans is included at the beginning of the section on taxonomy. Keys for the identification of families and subfamilies of the class Echiurida are provided, followed by a key to the genera of the Echiuridae and Thalassematidae. Although the family Echiuridae is represented in southern Africa by a single species (Echiurus antarcticus), it is also diagnosed and discussed in order to complete the taxonomic account. Species already well known as well as those not available in the present collection have been rediagnosed. The comprehensive reference list at the end of the thesis includes the references given at the end of each manuscript as well as those appearing elsewhere in the thesis. It was felt that a single and complete reference list would be more convenient for future researchers interested in this group of animalso THE TAXONOMY AND FUNCTIONAL ANATOMY OF SOUTHERN AFRICAN ECHIURANS CONTENTS Abstract 1 PART 1 TAXONOMY AND DIS'IRIBUTION Introduction 2 Materials and Methods 3 TAXONOMY Explanation of terms as used for echiurans 6 Diagnosis of phylum, classification and keys 9 Keys to the families and subfamilies of the class Echiurida 13 Key to the genera of Echiuridae and Thalassematidae 14 Family Thalassematidae Genus Ochetostoma Ochetostoma (Echiura) from southern Africa with the description of a new species 17 Key to the species of the genus Ochetostoma 21 Ochetostoma natalense sp. nov. 26 Ochetostoma baronii 41 Ochetostoma fonnosulurn 53 Ochetostoma decameron 64 Ochetostoma kempi 74 Ochetostoma palense 85 Ochetostoma sp. indet. 95 Ochetostorna arkc1ti 106 Ochetostoma ~nse 108 References 111 Publication: Some aspects of the anatomy of Ochetostoma caudex (Echiura) from the east coast of southern Africa with remarks on its taxonomic status. 116 Family Thalassernatidae Genus Thalasserna Thalasserna (Echiura) from southern Africa with the description of a new species. 125 Keys to the species of the genus Thalassema 128 Thalassema jenniferae sp. nov. 133 Thalasserna diaphanes 144 Thalassema philostracum 154 Thalassema sp. A. 162 Thalasserna sp. B. 171 Thalassema thalassemum 176 References 178 Family Thalassematidae Genus Anelassorhynchus Publication: A key to the species of Anelassorhynchus (Echiura) with a description of a new species from the east coast of southern Africa. 182 Anelassorhynchus indivisus 188 Anelassorhynchus moebii 190 Family Echiuridae 192 Genus Echiurus 192 Key to the species of Echiurus 193 Echiurus antarcticus 194 DISTRIBUTION Publication: The geographic distribution of Echiura from southern Africa 197 Discussion 208 CONTENTS PART II FUNCTIONAL ANATOMY Introduction 218 Burrowing, locomotion and other movements of the echiuran Ochetostoma caudex 220 The reproductive system, gametogenesis and annual reproductive cycle of Ochetostoma caudex (Phylum Echiura) 262 Discussion 301 Summary 305 Acknowledgements 307 References 308 ABSTRACT The southern African echiuran fauna currently comprises two families, four genera and eighteen species. The taxonomic positions of three other species, one of Ochetostoma and two of Thalassema, remain to be resolved. Of the recorded species, three are new to science and a further five are new records for the southern African region. This survey re= veals that there are ten species of Ochetostoma, four of Thalassema and three of Anelassorhynchus. The family Echiuridae contains a single genus, Echiurus and in southern Africa,~- antarcticus is the only species re= corded to date. Full descriptions of the new species and those formerly inadequately described are given. The diagnoses of some others are modi= fied and, where necessary, additional notes added. Keys for the identifi= cation of families, genera and species are provided and the taxonomic characters are evaluated and discussed. The zoogeographic distribution of the southern African species is mapped and an attempt has been made to analyse the recorded species on the basis of faunistic provinces. The phylogenetic position of the Echiura in the system of the animal kingdom is discussed. The lack of any form of segmentation of the meso dermal bands during embryonic developnent seems to justify their status as a separate phylum. The biomechanics of burrowing, locomotion and trunk irrigatory movements of Ochetostoma caudex are described and compared with the activities of some other animals with unsegmented coelomic cavities. Pressure changes within the coelom have been measured with the use of electronic recording techniques and the results interpreted in relation to direct visual observation. The anatomy of the reproductive system and the process of gametogenesis in O. caudex have been investigated with light and electron microscopes. Some observations on aspects of breeding behaviour and spawning are also included. PART 1 TAXONOMY and DISTRIBUTION 2 INTRODUCTION Echiurans are soft-bodied, burrowing animals with a spatulate proboscis at the anterior end as a result of which they are coumonly referred to as spoon or tongue worms. They are conmon inhabitants of the rocky s hores of the intertidal zone, although some deep water forms have been reported. A few species h~ve also extended into estuaries. They are widely distributed and have been reported from ~11 seas and all latitudes. At present, very little is J<nown about the echiuran fauna of southern Africa. No scientist from southern Africa has taken any interest in this group of animals. There is no single comprehensive report on the recorded species or on their range of distribution. The whole of the existing information on the phylum is confined to a very small number of publications. The only notable works on this group in southern Africa are those of Wesenberg-Lund (1959, 1963) and Stephen and Cutler (1969), A few publications, however, have referred to new species and new records (Jones & Stephen 1955, Biseswar 1983, 1984, 1985), The descriptions of a number of species are extremely brief and lack critical information which can be used to distinguish them. It is thus apparent that many species need re-examination and redescription. In addition, ·some of the species are based on ~ingle individuals which means that nothing is known about the variations that occur within the species. 3 Only ten species have previously been recorded from the southern African region. A further three new species are described here and several new records added bringing the total number of named species for southern Africa to eighteen. The discovery of additional localities of some of the species previously recorded in the region sheds further light on their range of distribution. There are three further specimens in the present collection, one belonging to the genus Ochetostoma and two to Thalassema, that are probably new to science. Unfortunately two of them, belonging to the genus Thalassema, are too badly damaged to allow adequate descriptions. Any further decisions regarding their taxonomic positions will