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An Analysis of the Distribution of a Commensal

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AN ANALYSIS OF THE DISTRIBUTION OF A COMMENSAL POLYNOID ON ITS HOSTS by JOHN BEACH PALMER , " . :.".~. ~ .. " ': j A DISSERTATION March 1968 VITA NAME OF AUTHOR: John Beach Palmer PLACE OF BIRTH: Grand Rapids, Michigan DATE OF BIRTH: February 22, 1941 UNDERGRADUATE AND GRADUATE SCHOOLS ATTENDED: Oberlin College ~. University of Oregon DEGREES AWARDED: Bachelor of Arts, 1963, Oberlin College AREAS OF SPECIAL INTEREST: Teaching and research in Population and Community Ecology, Evolution, and Invertebrate Zoology PROFESSIONAL EXPERIENCE: Research Assistant to Dr. Peter W. Frank, University of Oregon, 6/63 to 9/63, 9/64 to 6/65. ~eaching Assistant in Invertebrate Zoology under Dr. Peter W. Frank, University of Oregon, 9/63 to 12/63. Teaching Assistant in General Biology, University of Oregon, 1/64 to 6/64. Teaching Assistant in Invertebrate Zoology under Mr. Richard Darby, Oregon Institute of Marine Biology, Summer Session 1964. Teaching Assistant in Marine Ecology under Dr. John P. Kerr, Oregon Institute of Marine Biology, Summer Session 1965. General Biology Instructor for Division of Continuing Education, University of Oregon, 9/64 to 6.65. Teaching Assistant in Invertebrate Zoology under Dr. Stephen Brown, Oregon Institute of Marine Biology, Summer Session 1967. Instructor in Biology, University of Oregon, 9/67 to 6/68. ~ ..; f i 1 VITA - continued AWARDS AND HONORS: Finalist National Merit Program 1959. Scholarship to Oberlin College 1959-1963. Scholarship to Marine Biological Laboratories, Woods Hole, Mass. 1962. Elected to Sigma Xi, 1963. High Honors in Biology, 1963. NSF Graduate Fellowship, 1965 to 1967. PUBLICATIONS: Distribution of a commensal polynoidon a keyhole limpet (Abstract), Bull. Ecol. Soc. Am. 46(4):160-161. !. Settling and growth of a commensal polynoid, Arctonoe vittata (Abstract), Bull. Ecol. Soc. Am. 47(2):88. ii AN ANALYSIS OF THE DISTRIBUTION OF A COMMENSAL POLYNOID ON ITS HOSTS by JOHN BEACH PALMER A DISSERTATION Presented to the Department of Biology and the Graduate School of the Universfty of Oregon in partial fulfillment of the requirements for the degree of Doctor of Philosophy March 1968 .~;~.. NOT APPROVED (Adviser for the thesis, Peter W. Frank) It iv I would like to acknowledge the help and guidance of Dr. Peter W. Frank in developing this problem with Arctonoe. v To Patsy and Robert for braving cold Pacific waters; To Ann for bravely typing; And to all the Cryptochiton that have ever lived for bravely being what they are. vi TABLE OF CONTENTS Page INTRODUCTION •••••• 1 METHODS AND STUDY AREAS 6 RESULTS 13 Habitat and Habits of Hos ts ••••••••••• 13 Distribution of the Commensal on Its Hosts 16 Negative Worm-worm Interaction •• ••••• 23 Recruitment of the Commensal •• ••••• 29 Growth Rate and Structure of Commensal Population 45 Growth Rate and Structure of Host Population 50 Variation in Population Structure with Location, Host and Species of Commensal ••••••• 58 SUMMARY AND DISCUSSION 79 BIBLIOGRAPHY 95 APPENDIXES 1. Success of Tagging Methods for Cryptochiton and Diodora ,. ...... ,. ....... 102 2. Host Factor Responses of ~. vittata •• 106 3. Recapture Record of Marked Cryptochiton 108 4. Recapture Record of Marked Diodora 110 vii LIST OF TABLES Table Page 1. Comparisons of observed distributions of various ectoparasites to the Poisson distribution 3 2. Distribution of the number of ~. vittata per host Diodora from the Cape Arago region of the Oregon coast, 1964 to 1966 .••• 17 3. Distribution of the number of A. vittata per host Cryptochiton from the Cape Arago region of the Oregon coast, 1964 to 1965 ••••• 18 4. Distribution of the size of A. vittata from host Diodora in the Cape Arago region of the Oregon coast, 1964 to 1966 •••.••••••••• 20 5. Distribution of the size of A. vittata from host Cryptochiton in the Cape Arago region of the Oregon coast, 1964 to 1965 ••••••••• • 21 6. A binomial model of occupancy for the commensals on Cryptochiton compared with the Poisson distribution of the same average occupancy .••••••••• 22 7. Goodness-of-fit analysis of all worms less than 20mm in length from a sample of 51 Diodora collected in July 1966 from Simpson Reef at Cape Arago 24 8. Variance:mean analysis of all worms less than 20mm long from 9 samples of hosts •••••••••• 24 9. Frequencies of worms from marked Diodora compared to those from previously undisturbed hosts ••• 26 10. Comparison of the frequencies of the larger small worms (10-19mm size class) on Diodora and Cryptochiton from Sunset Bay, Oregon in 1965 •••• 27 11. Comparison of abundance of A. vittata on Diodora and Solaster from San Juan Island, Washington in 1966 28 12. Reinfestation of marked Cryptochiton 31 13. Reinfestation of marked Diodora 31 viii LIST OF TABLES - continued Table Page 14. Sizes of WOTITIS collected from marked hosts after 1 week of exposure to recruitment ..•. 32 15. WOTITI vagility experiments 34 16. Seasonal settlement as indicated from field census data . 0 0 •••••• It • " 0 '" ~ ••••••• 37 17. Seasonal settlement as indicated by data from the recapture of marked Cryptochiton •.•• 38 18. Seasonal settlement indicated by data from the recapture of marked Diodora •.•• 39 19. Abundance of small WOTITIS (2Ornm or less in length) in f three samples of Diodora collected during one week in July 1966 •. 44 20. Abundance of newly recruited WOTITIS on marked Diodora collected within three days in October 1965 44 21. Data for calculations of turnover time • 48 22.--Correlation Coefficients of the size relation of host and wonn ........ " ... 58 23. Distribution of the number of A. vittata per host Cryptochiton collected in Washington and California 60 24. Distribution of the size of A. vittata from host Cryptochiton collected in California 60 25. Distribution of the number of A. vittata per host Diodora collected off San Juan Is., Washington in March 1966 •••••••••.•.••• 62 26. Distribution of the size of A. vittata from Diodora collected off San.Juan Is., Washington in March 1966 • 62 27. Analysis of the distribution of the number of A. vittata per host Solaster collected off San Juan Is., Washington .••••••••• 64 28. Distribution of the size of A. vittata from Solaster collected off San Juan Is., Washington •..••• 64 ix LIST OF TABLES - continued Table Page 29. Distribution of the size of Solaster collected off San Juan Is., Washington. 65 30. Distribution of the number of ~. pulchra and A. vittata on Megathura collected at Pacific Grove, California in 1966 •..•.•.•• •• .•• •. •. 67 31. Distribution of the size of ~. pulchra and A. vittata on Megathura collected at Pacific Grove, California in 1966 ...•...••. ••. •• 67 32. Analysis of the number of ~. pulchra per host Parastichopus collected off San Juan Is., Washington. 67 33. Distribution of the size of A. pulchra from Parastichopus collected off San Juan Is., Washington. •• •• 68 34. Analysis of the distribution of the number of ~. pulchra per host on 3 of its hosts collected at Punta Banda, Mexico in August 1966 ••••••••••• •• 68 35. Distribution of the size of ~. pulchra from 3 hosts collected at Punta Banda, Mexico in August 1966 68 36. Distribution of the number of Podarke per host Patiria collected at Pacific Grove, California 75 37. Distribution of the size of Podarke pugettensis from host Patiria collected at Pacific Grove, California in March 1967 .••••••••• 76 38. Cases of the regular distribution of commensals 93 x LIST OF FIGURES Figure Page 1. Collection and return stations in the Cape Arago region of the Oregon coast ••••• 7 2. Habitat of Diodora and Cryptochiton in the Cape Arago region of the Oregon coast • ·. 15 3. Histories of 16 ~. vittata in marked Cryptochiton at the Light House Channel between July 1965 and February 1966 •••••••••••••••••.• , 36 4. Frequency of sexually mature ~. vittata from Crypto­ chiton and Diodora in the Cape Arago region of the Oregon coast, 1964 to 1966 ••.••••••••• .' 41 5. Size frequency distribution of very recently settled worms on Diodora and Cryptochiton collected in August 1965 ••• ., 43 6. Growth in A. vittata • 46 7. Growth in Cryptochiton 51 8. Size-frequency distributions of Cryptochiton • 52 9. Growth in Diodora 54 10. Size-frequency distributions of Diodora ·. 56 11. Size of Fabia faba and size of host Schizothaerus 72 12. Size of the male and female Fabia collected from Schizothaerus ••••••••••.••• 73 13. Analysis of a sample of Podarke pugettensis and host Patiria miniata collected from Monterey Bay, California in March 1967 ••••••••••••• ·. 77 14. Death schedule (qx) for A. vittata on Diodora and Cryptochiton .......•....•.•... 87 xi INTRODUCTION " .•• In displacing other shells, I found in nearly everyone a similar tenant: the secret was discovered, the worm was a parasite, that lived in peace and good fellowship with the Keyhole •.••At least four out of every six contained a para­ site, and, what is rather strange, the worms were nearly all of one size." From the first written account of contact between Western Science and Arctonoe vittata.--J. K. Lord, 1866. The observed distribution of an animal population in its habitat is the resultant of a large number of factors such as reproductive habits, gregariousness or its lack, catastrophes, resource distribution, habitat selection, etc. It is also influenced by the manner and time of sampling. Distribution is an aggregate phenomenon. Ideally, one might determine the true distribution of the population, then resolve the various forces responsible for it, and finally determine the selec- tive value of this pattern of distribution to the individuals which make up the population. Such an account is probably not complete for any population. Another, less heroic approach is to study population distributions as phenomena with a certain demographic logic of their own. This has been the rationale of the present study. The distribution of many organisms is difficult to analyze because accurate and natural definitions of the unit regions in which the population is distributed are hard to make. Depending on the size of the sample unit, the individuals of a population will, with suffi- ciently large sample size, show aggregation and with smaller sample 1 2 size demonstrate random or regular distribution (Greig-Smith, 1964; Slobodkin, 1961).
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  • Polychaete Worms Definitions and Keys to the Orders, Families and Genera

    Polychaete Worms Definitions and Keys to the Orders, Families and Genera

    THE POLYCHAETE WORMS DEFINITIONS AND KEYS TO THE ORDERS, FAMILIES AND GENERA THE POLYCHAETE WORMS Definitions and Keys to the Orders, Families and Genera By Kristian Fauchald NATURAL HISTORY MUSEUM OF LOS ANGELES COUNTY In Conjunction With THE ALLAN HANCOCK FOUNDATION UNIVERSITY OF SOUTHERN CALIFORNIA Science Series 28 February 3, 1977 TABLE OF CONTENTS PREFACE vii ACKNOWLEDGMENTS ix INTRODUCTION 1 CHARACTERS USED TO DEFINE HIGHER TAXA 2 CLASSIFICATION OF POLYCHAETES 7 ORDERS OF POLYCHAETES 9 KEY TO FAMILIES 9 ORDER ORBINIIDA 14 ORDER CTENODRILIDA 19 ORDER PSAMMODRILIDA 20 ORDER COSSURIDA 21 ORDER SPIONIDA 21 ORDER CAPITELLIDA 31 ORDER OPHELIIDA 41 ORDER PHYLLODOCIDA 45 ORDER AMPHINOMIDA 100 ORDER SPINTHERIDA 103 ORDER EUNICIDA 104 ORDER STERNASPIDA 114 ORDER OWENIIDA 114 ORDER FLABELLIGERIDA 115 ORDER FAUVELIOPSIDA 117 ORDER TEREBELLIDA 118 ORDER SABELLIDA 135 FIVE "ARCHIANNELIDAN" FAMILIES 152 GLOSSARY 156 LITERATURE CITED 161 INDEX 180 Preface THE STUDY of polychaetes used to be a leisurely I apologize to my fellow polychaete workers for occupation, practised calmly and slowly, and introducing a complex superstructure in a group which the presence of these worms hardly ever pene- so far has been remarkably innocent of such frills. A trated the consciousness of any but the small group great number of very sound partial schemes have been of invertebrate zoologists and phylogenetlcists inter- suggested from time to time. These have been only ested in annulated creatures. This is hardly the case partially considered. The discussion is complex enough any longer. without the inclusion of speculations as to how each Studies of marine benthos have demonstrated that author would have completed his or her scheme, pro- these animals may be wholly dominant both in num- vided that he or she had had the evidence and inclina- bers of species and in numbers of specimens.