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University Microfilms International 300 North Zeeb Road Ann Arbor, Michigan 48106 USA St INFORMATION TO USERS This material was produced from a microfilm copy of the original document. While the most advanced technological means to photograph and reproduce this document have been used, the quality is heavily dependent upon the quality of the original submitted. The following explanation of techniques is provided to help you understand markings or patterns which may appear on this reproduction. 1. The sign or "target" for pages apparently lacking from the document photographed is "Missing Page(s)". If it was possible to obtain the missing page(s) or section, they are spliced into the film along witn adjacent pages. This may have necessitated cutting thru an image and duplicating adjacent pages to insure you complete continuity. 2. When an image on the film is obliterated with a large round black mark, it is an indication that the photographer suspected that the copy may have moved during exposure and thus cause a blurred image. You will find a good image of the page in the adjacent frame. 3. When a map, drawing or chart, etc., was part of the material being photographed the photographer followed a definite method in "sectioning" the material. It is customary to begin photoing at the upper left hand corner of a large sheet and to continue photoing from left to right in equal sections with a small overlap. If necessary, sectioning is continued again - beginning below the first row and continuing on until complete. 4. The majority of users indicate that the textual content is of greatest value, however, a somewhat higher quality reproduction could be made from "photographs" if essential to the understanding of the dissertation. Silver prints of "photographs" may be ordered at additional charge by writing the Order Department, giving the catalog number, title, author and specific pages you wish reproduced. 5. PLEASE NOTE: Some pages may have lndistinct received. University Microfilms International 300 North Zeeb Road Ann Arbor, Michigan 48106 USA St. John's Road, Tyler's Green High Wycombe, Bucks, England HP10 8HR 78-10,254 LIGIITER, Frederick John, 1942- 1HE SOCIAL USE OF SPACE: ASPECTS OF ECOL<XN, ETI-lOLCXJY .AND ENDOCRINOL<XN IN 'TIlE GHOST CRABS OCYPODE CERATOPIm-IAIMUS (PALLAS) AND OCYPODE LAEVIS DANA. University of Hawaii, Ph.D., 1977 Zoology University Microfilms International, Ann Arbor, Michigan 48106 THE SOCIAL USE OF SPACE: ASPECTS OF ECOLOGY, ETHOLOGY AND ENDOCRINOLOGY IN THE GHOST CRABS OCYPODE CERATOPHTHALMUS (PALLAS) AND OCYPODE LAEVIS DANA A DISSERTATION SUBMITTED TO THE GRADUATE DIVISION OF THE UNIVERSITY OF HAWAII IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE DEGREE OF DOCTOR OF PHILOSOPHY IN ZOOLOGY DECEMBER 1977 By Frederick John Lighter Dissertation Committee: Ernst S. Reese, Chairman Franklin Chang Fred I. Kamemoto George S. Losey John Stimson ABSTRACT This study concerns the social use of space in two species of ghost crabs, Ocypode ceratophthalmus (Pallas) and Ocypode laevis Dana. The work involves four major aspects: the spatial patterns of the burrows of both species in nature, the space-related behaviors of the two species, a test of a mathematical model of socia. interaction for ~. ceratophthalmus, and the influence of the androgenic gland on burrowing behavior and growth in ~. ceratophthalmus. To determine the natural spatial patterns of the two species, nearest neighbor, I mean crowding' and 'patchiness' measurements are made for thirteen populations of Q. ceratophthalmus and seven popula­ tions of O. laevis. Measurements of density, 'mean crowding I and 'patchiness' fail to show a close association with dispersion pattern as measured by nonparametric regressions. Rank correlations demonstrate no significant association between size of individual and distance to nearest neighbor. All individual O. laevis interact to produce a total population dispersion pattern. In contrast, the reproductively­ active male ~. ceratophthalmus, through behavioral competition for space, exert the major influence in determining population dispersion. Three categories of space-related behaviors are observed in the field and laboratory: burrow defense, defense of an area surrounding the burrow, and defense of individual distance. Nine modal action patterns are utilized by ~. ceratophthalmus in its space-related behavioral repertoire; six are found in O. laevis. ~. ceratophthalmus defends space in order to guarantee ownership of a burrow, to reduce iii competitive social stimulatior. between neighbors, and to provide for an area for sexual display. ~. laevi~ defends space in order to provide ownership of a burrow and to reduce competitive social stimula­ tion between neighbors. Crowding experiments with ~. ceratophthalmus show that, as density increases, defensive behavior shifts from burrow and area defense to the maintenance of individual distance. Work with models demonstrates that the presence of an adult male within a defended area elicits Threat by a resident reproductively-active male ~. ceratoph­ thalmus. An intruder which exhibits Threat elicits a higher intensity response from the resident than a nonthreat intruder. Calhoun's (1957, 1963) mathematical model of social interaction is tested on field and laboratory data for ~. ceratophtha1mus. The data do not follow the prediction of the model that the frequency of social contacts per unit time increases as group size increases. Rather, the data indicate that the relationship is bimodal, with peak frequencies 2. at 2.5 and 4.5 individua1s/m As an alternative to Calhoun's model, I conclude that increased density produces a period of intensified social competition for space. After this period, Q. ceratophthalmus enters an lI ext ended refractory period,1I which is reflected in periods of low frequency of social activity. The influence of the androgenic gland on burrowing behavior and growth of ~. ceratophthalmus is investigated by implantation and removal experiments. No obvious differences in the growth of carapace width, pollex length, abdomen length and abdomen width are found between implanted individuals, individuals with the androgenic gland iv gland removed, and sham-operated control animals. Changes in burrowing behavior are found in implanted juvenile females and an implanted juvenile male. In these individuals, burrowing behavior is changed toward that of an adult, reproductively-active male. Evidence suggests that androgenic gland secretory activity, and thus male reproductive activity, is synchronized within populations to coincide with the period of the full moon, when time of burrow exposure is greatest. Implanted juvenile females show a masculinization in the value of the abdomen width/ carapace width ratio, while implanted females do not. For females, a critical period exists beyond which androgenic gland implantation no longer produces secondary male sexual characteristIcs. v TABLE OF CONTENTS Page ABSTRACT iii LIST OF TABLES viii LIST OF ILLUSTRATIONS x CHAPTER I. GENERAL INTRODUCTION 1 CHAPTER II. THE SPATIAL PATTERNS OF OCYPODE CERATOPHTHALMUS (PALLAS) AND OCYPODE LAEVIS DANA IN NATURE 3 Introducti on 3 Materials and Methods 4 Results 9 Nearest Neighbor Measurements 9 'Mean Crowding' and 'Patchiness' Measurements 9 Correlation Tests 10 Summary of Statistical Measurements for Each Field Site 10 Di scussi on 13 Conclusions 22 CHAPTER III. THE SPACE-RELATED BEHAVIORS OF THE GHOST CRABS OCYPODE CERATHOPHTHALMUS (PALLAS) AND OCYPODE LAEVIS DANA 38 Introduction 38 Materials and Methods 41 Results: Ocypode ceratophthalmus 47 Burrow Defense 47 Defense of Area Surrounding the Burrow 62 Burrow-Filling by Reproductively-Active Male Ocypode ceratophthalmus 65 vi TABLE OF CONTENTS, (Continued) Page Social Interactions of Ocypode ceratophthalmus Occuring during Foraging Away from Burrow 66 Crowding Experiments with Ocypode ceratophthalmus 68 Use of Models with Reproductively-Active Male Ocypode ceratophthalmus 73 Results: Ocypode laevis 75 Burrow Defense 75 Defense of Area Surrounding the Burrow 79 Social Interactions of Ocypode laevis Occuring during Foraging Away from Burrow 81 Discussion 83 Conclusions 95 CHAPTER IV. A TEST OF A MODEL OF SOCIAL INTERACTION FOR OCYPODE CERATOPHTHALMUS (PALLAS) 97 Introduction 97 Materials and Methods 98 Results 101 Discussion 107 Conclusions 115 CHAPTER V. AN INVESTIGATION OF THE ROLE OF THE ANDROGENIC GLAND IN THE BURROWING BEHAVIOR AND GROWTH OF OCYPODE CERATOPHTHALMUS (PALLAS) 117 Introduction 117 Materials and Methods 118 Results 121 Discussion 136 Conclusions 140 BIBLIOGRAPHY 142 vii LIST OF TABLES Table Page 1. Summary of Field Sites 24 2. Some Characteristics of Each Field Site 26 3. Summary of Spatial Measurements for each Field Site for Ocypode ceratophthalmus 28 4. Summary of Spatial Measurements for each Field Site for Ocypode laevis 30 5. Summary of Spatial Measurements for each Field Site where Ocypode ceratophthalmus and Ocypode laevis occur together in Overlapping Ranges 31 6. Summary of Size Range and Dispersion Pattern for Ocypode laevis for each Field Site 32 7. Summary of Size Range and Dispersion Pattern for Ocypode ceratophthalmus for each Field Site 33 8. Summary of Data for Reproductively-Active Male Ocypode ceratophthalmus for each Field Site 35 9. Comparison of Average Nearest Neighbor Distances (in meters) for Total Populations of Q. ceratophthalmus and for the Reproductively- Active Males within the Population 37 10. Burrow Defense Behavior of Ocypode ceratophthalmus 56 11. Area Defensive Behavior of Ocypode ceratophthalmus 64 12. Values (Number of interactions/individual/hour)
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