Biology of Aspidobothrian Trematodes David William Fredericksen Iowa State University

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Biology of Aspidobothrian Trematodes David William Fredericksen Iowa State University Iowa State University Capstones, Theses and Retrospective Theses and Dissertations Dissertations 1973 Biology of aspidobothrian trematodes David William Fredericksen Iowa State University Follow this and additional works at: https://lib.dr.iastate.edu/rtd Part of the Zoology Commons Recommended Citation Fredericksen, David William, "Biology of aspidobothrian trematodes " (1973). Retrospective Theses and Dissertations. 5010. https://lib.dr.iastate.edu/rtd/5010 This Dissertation is brought to you for free and open access by the Iowa State University Capstones, Theses and Dissertations at Iowa State University Digital Repository. It has been accepted for inclusion in Retrospective Theses and Dissertations by an authorized administrator of Iowa State University Digital Repository. For more information, please contact [email protected]. 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Xerox University Microfilms 300 North Zeeb Road Ann Arbor, Michigan 48106 74-538 FREDERICKSEN, David William, 1943- BIOLOGY OF ASPIDOBOTHRIAN TREMATODES. Iowa State University, Ph.D., 1973 Zoology University Microfilms, A XEROX Company, Ann Arbor, Michigan © 1973 DAVID WILLIAM FREDERICKSEN ALL RIGlirS RESERVED THIS DISSERTATION HAS BEEN MICROFILMED EXACTLY AS RECEIVED. Biology of aspidobothrian trematodes by David William Fredericksen A Dissertation Submitted to the Graduate Faculty in Partial Fulfillment of The Requirements for the Degree of DOCTOR OF PHILOSOPHY Department: Zoology and Entomology Major: Zoology (Parasitology) Approved: Signature was redacted for privacy. In Cnaige of Major Work Signature was redacted for privacy. For the Major Department Signature was redacted for privacy. For th^ G^a<fuate College Iowa State University Ames, Iowa 1973 Copyright^David Willicun Fredericksen, 1973. All ri reserved. ii TABLE OF CONTENTS Page INTRODUCTION 1 HISTORICAL REVIEW OF SUBCLASS ASPIDOBOTHRIA 3 MATERIALS AND METHODS 7 ADULT MORPHOLOGY 14 External Morphology 14 Cotylogaster occidentalis 15 Aspidogaster conchicola 16 Cotylaspis insignis 18 Other Aspidobothria 18 NERVOUS SYSTEM MORPHOLOGY 22 Cotylogaster occidentalis 24 Aspidogaster conchicola 29 Cotylaspis insignis 33 SENSORY STRUCTURES 4 7 Cotylogaster occidentalis 49 Aspidogaster conchicola 50 Cotylaspis insignis 51 THE COTYLOCIDIUM LARVA 54 DEVELOPMENT 70 Previous Studies 70 Specimen I 73 Specimen II 75 Specimen III 7 5 iii Page Specimen IV 76 Specimen V 77 Specimen VI 77 Young Adult 78 Host-Parasite Relationships 82 Cotylogaster occidentalis 84 Aspidoqaster conchicola 92 Cotylaspis insignis 93 PHYLOGENY 9 7 SUMMARY AND CONCLUSIONS 100 LITERATURE CITED 105 ACKNOWLEDGMENTS 117 APPENDIX 121 PLATES 122 Abbreviations I 1 INTRODUCTION Among the Platyhelminthes there occurs a small, interest­ ing subclass of trematodes, the Aspidobothria Burmeister 1856. The group is comprised of nine recognized genera and at least 27 species, occurring in fresh and marine waters as parasites of both molluscs and poikilothermous vertebrates. In one case, however, a marine crustacean may serve as a host. In all known instances, aspidobothrians develop directly, without asexual stages, and maturity is usually reached in a molluscan host. They are easily recognized by their large, unarmed, ventral adhesive discs divided into many alveoli or separated into individual suckers. They lack the typical oral sucker of other trematodes and have, instead, a buccal disc at the anterior end of a retractable neck. The dorsal posterior portion of the aspidobothrian body usually protrudes beyond the ventral adhesive disc as a muscular cone containing the excretory pore(s). A simple rhabdocoelous gut is common to all members of the subclass. There are six genera known to occur in the fresh-waters of the world, and of these, only three occur commonly in fresh-waters of North America, namely Cotylaspis insignis Leidy 1857 , Cotylogaster occidentalis Nickerson 1902 , and Aspidogaster conchicola v. Baer 1827. Although each of these parasites occurs in the intestines of various poikilothermous 2 vertebrates, all three are primarily parasites of pelecypod molluscs, and each occupies a different niche in the mussel host. Cotylaspis insignis, an ectoparasite, is found on the epidermis covering the foot, kidney, and in the water chambers of the mussel gill. Cotylogaster occidentalis, a lumen dweller, occurs in the intestines of mussels, whereas Aspidogaster conchicola resides in the pericardial cavity and kidney of mussels. Due to their nature, these three members of the family Aspidogastridae Poche 1907 lend themselves to interesting biological comparisons, the subject of this thesis. These three parasites have been investigated from various standpoints of their adult morphology, initial larval stage, development, and host-parasite relationships. Data gained from studies on the aforementioned aspects have many bio­ logical and phylogenetic implications concerning the nature of aspidobothrian trematodes. 3 HISTORICAL REVIEW OF SUBCLASS ASPIDOBOTHRIA Aspidobothrians have long been of uncertain taxonomic status because of their unique characteristics and paucity of information regarding their biology. Monogenean and digenean trematodes fall within well defined boundaries, but such is not true of aspidobothrians. In all previous studies, development among members of the latter group has been shown to be direct as in the monogeneans. However, aspidobothrians lack a major monogenean characteristic, namely, sclerotized elements in their well-developed adhesive discs, Aspido­ bothrians resemble digeneans morphologically, and many authors presently consider these groups closely related. However, evidence needed to assess their precise position has been lacking and consequently the group has remained enigmatic. Their uncertain status is borne out by the fact that since their discovery they have sometimes been placed within the Monogenea, sometimes witnin the Digenea as well as having been given status coordinate with these two trematode groups. Aspidogaster conchicola v. Baer 1827, was the first aspido- bothrian fluke to be described, and in Dujardin's 1845 system of trematode classification, A. conchicola was included with the group now known as the distomate flukes. Diesing (1850) devised a system of classification which placed A. conchicola alone in the subtribe Rhabdocoela, but in the same tribe with 4 certain monogenean and digenean trematodes. Aubert (1855) studied the development of A. conchicola and noted that it undergoes neither metamorphosis nor alternation of genera­ tions. Aspidobothrian trematodes were first recognized as a separate group by Burmeister in 1856 when he referred to this group of worms as the Aspidobothrii, with status equal to his other two groups Malacobothrii and Pectobothrii. The latter two groups have been interpreted by Stunkard (1962, 1963) as containing Digenea and Monogenea, respectively. Van Beneden (185 8) divided the trematodes into the two groups now known as Monogenea and Digenea, and, contrary to the report of Stunkard (1963) , van Beneden placed A. conchicola with the monogenean group Polystomatidae. Taschenberg (1879) reviewed the systematics of the Trematoda and included aspidcbothrians with the monogenean group Microcotylidae. Leuckart (1879) suggested that aspidobothrians are essentially sexually mature rediae and thus linked them to the Digenea. Cunning­ ham described Stichocotyle nephropis in 1887 and indicated it should be included with the monogenean group Polystoma­ tidae. Hoyle (1888), in a review of the trematodes, grouped them in accordance with van Beneden'e (1858) views, and placed the aspidobothrians among the Monogenea. Hoyle (1888) was unable to assess
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