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Uhm Phd 9300335 R.Pdf INFORMATION TO USERS This manuscript has been reproduced from the microfilm master. UMI films the text directly from the original or copy submitted. Thus, some thesis and dissertation copies are in typewriter face, while others may be from any type of computer printer. The quality of this reproduction is dependent upon the quality of the copy submitted. Broken or indistinct print, colored or poor quality illustrations and photographs, print bleedthrough, substandard margins, and improper alignment can adversely affect reproduction. In the unlikely event that the author did not send UMI a complete manuscript and there are missing pages, these will be noted. Also, if unauthorized copyright material had to be removed, a note will indicate the deletion. Oversize materials (e.g., maps, drawings, charts) are reproduced by sectioning the original, beginning at the upper left-hand corner and continuingfrom left to right in equal sections with small overlaps. Each original is also photographed in one exposure and is included in reduced form at the back of the book. Photographs included in the original manuscripthave been reproduced xerographically in this copy. Higher quality 6" x 9" black and white photographic prints are available for any photographs or illustrations appearing in this copy for an additional charge. Contact UMI directly to order. U·M·I University Microfilms International A Beil & Howell Information Company 300 North Zeeb Road. Ann Arbor. M148106-1346 USA 313/761-4700 800/521-0600 Order Number 9300335 Behavioral ecology of reproduction in the pomacanthid angelfish, Centropyge potteri Lutnesky, Marvin Michael Francis, Ph.D. University of Hawaii, 1992 Copyright @1992 by Lutnesky, Marvin Michael Francis. All rights reserved. U·M·I 300 N. Zeeb Rd. Ann Arbor. MI 48106 BEHAVIORAL ECOLOGY OF REPRODUCflON IN THE POMACANTHID ANGELFISH, CENTROPYGE POTIERI A DISSERTATION SUBMfITED TO THE GRADUATE DMSION OF THE UNIVERSITY OF HAWAII IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE DEGREE OF DOcrOR OF PHIWSOPHY IN ZOOLOGY AUGUST 1992 BY Marvin M. F. Lutnesky Dissertation Committee: Ernst S. Reese, Chairperson Milton Diamond Leonard A Freed George S. Losey John S. Stimson o Copyright by Marvin M. F. Lutnesky 1992 All rights reserved iii This work is dedicated to all those who have taught me the value of exploration, especially my parents, Patricia R. Lutnesky and the late Marvin L. Lutnesky, and my wife Kimberly D. K Lutnesky. iv ACKNOWLEDGMENTS I thank E. Reese for his guidance and encouragement throughout this project, and also the other members of my dissertation committee: M. Diamond, L Freed, G. Losey, and J. Stimsonfor their advice on this project and their contributions to my development as a scientist. In addition to my dissertation committee, I also thank G. Barlow, T. Hourigan, D. R. Robertson, R. Ross, W. Searcy, D. Shapiro, W. Tyler,and R. Warner for their commentson early versions of various chapters in the dissertation. This work was possible due to the generosity of my dive buddies, D. Anderson, C. Fiedler, R. Fitzhardinge, J. Godwin, R. Kosaki, J. Mahon, O. McMillan, J. Mendez, L. Privitera, D. R. Robertson, S. Romano, M. Sarrazin, U. Schober, RJ.F. Smith, F. Stanton, W. Tyler, and D. Worthington. My dive buddies and D. Chu, B. Danilowicz, D. Gulko, S. Haley, D. IroDS, C. Kelley, B. Kessing, K. Lutnesky, S. Murphy-Walker, R. Pyle, and T. Tricas are also thanked for helpfuldiscussions and assistance with various aspects of this project. I am especially grateful to R. Kosaki for helping me test the temporal-threshold model of polygynous mating. Finally, I am most grateful to my wife, Kimberly, for her patience and encouragement during my pursuit of this study. Financial support was provided by the ARCS foundation, the Department of Zoology at the University of Hawaii, the Sigma Xi Society, and a grant from the Edwin W. Pauley Foundation, "Summer Program in Marine Science," to the Hawaii Institute of Marine Biology. v ABSTRACT The Hawaiian angelfish, Centropyge potteri, was used as a model system to develop and test hypotheses regarding the integration of stimuli from both physical and social environments into reproductive strategies; specifically the social control of sex change, and temporal patterns of mating. Prerequisite to these topics are descriptions of sexual dimorphism and dichromatism, and protogynous sex change. These descriptions include differences between the sexes in meristics, morphometries, and color pattern; a field-test of identification of the sexes; and histological descriptions of the gonads. Encounter-rate models are developed for territorial-haremic fishes. They show how a combination of social-group composition and density may form predictable patterns of contact between social-group members. The patterns of contact are asserted as cues for sex change. The models are tested with S;. potteri by demonstrating that encounters with smaller fish are needed for sex change, encounters with larger fish prevent sex change, and changingthe density of a social group alters encounter rates and can induce sex change. Sex change is associated with conditions that yield high smaller-fish encounter, and low larger-fish encounter, for the sex-change candidate. A temporal-threshold model of polygynous mating (TMPM) is developed. This model shows how females that mate with polygynous males may offset polygyny costs by mating during less advantageous times for which fewer females compete. The model's potential for broad applicability is shown to be worthy of study with examples from African elephants and C. potteri. The TMPM is empirically tested using C. potteri. Females compete for mating order. Competition results in a larger variance in daily time of mating, vi independent of the time required for mating, when females are in a larger group. Females probably disperse mating in time to avoid polygyny costs, including interference by competing females. These results were .iJ. priori predictions of the TMPM. The conclusion is that information from both physical and social environments is integrated into J;. potten reproductive strategies. In this way females may minimize potential costs associated with changing sex, and reproducing in a polygynous mating system. vii TABLE OF CONTENTS ACKNOWlEDGMENTS v ABSlRACf vi LIST OF TABLES x liST OF FIGURES xi I. GENERAL INTRODUCTION 1 Organization of the dissertation 3 II. SEXUAL DIMORPHISM, PROTOGYNOUS HERM­ APHRODITISM, AND SOCIAL BEHAVIOR OF THE POMACANTHID ANGELFISH, CENTROPYGE POTTERI.... 6 Introduction 6 Methods 8 Collections 8 Identification of sex 9 Sexual dimorphism and dichromatism 10 Histology 13 Sex change demonstration 14 Behavior 16 Reproductive behavior 16 Results and Discussion 17 Identification of sex 17 Sexual dimorphism and dichromatism 18 Histology 26 Sex change demonstration 31 Behavior 34 Reproductive behavior 37 Conclusions 44 III. DENSITY-DEPENDENT PROTOGYNOUS SEX CHANGE IN TERRITORIAL-HAREMIC FISHES: MODELS AND EVIDENCE 45 Introduction 45 Proximate control theory 46 Encounter-rate models 50 Assumptions 50 Absolute encounter-rate threshold hypothesis 53 Differential encounter-rate threshold hypothesis 54 Inhibition-only hypothesis 57 Testing the models 57 Methods 58 viii Field 58 Enclosures 59 Results and Discussion 64 Field 64 Enclosures 67 General Discussion 76 Conclusions 81 N. A TEMPORAL-THRESHOLD MODEL OF POLYGYNOUS MATING IN CYCliCAL ENVIRONMENTS 82 Introduction 82 A temporal-threshold model ofpolygynous mating 86 Assumptions 86 Predictions 88 Discussion 93 Elephan.ts 94 Angelfish 99 limitations and tests of the temporal-threshold model ofpolygynous mating 104 Conclusions 105 v. A TEST OF THE TEMPORAL-THRESHOLD MODEL OF POLYGYNOUS MATING 107 Introduction 107 Methods 109 System 109 Enclosures 112 Field 115 Results and Discussion 116 Conclusions 135 VI. CONCLUDING REMARKS AND FUTURE RESEARCH 136 UTERATURE crrsn 143 ix LISTOFTABLES Table page 2.1 ~e&~::g::s:upa~~:1 ~~~.~~~~:.~~.~~~~~.~~.~~~~~~ 24 2.2 Spawning ofCentrQP.Yge potteri during all weeks ofthe lunar month 43 3.1 Sex-change experiments using the pomacanthid angelfish, Centropyge potteri 70 x LIST OF FIGURES FIGURE page 2.1 Schematic drawing ofCentroVSge potteri showing line segments used in calculation of the color-pattern index 12 2.2 Color pattern of typical male and female CentrQPY~ ;potten 20 2.3 Plots of the relationships between sexually dimorphic characters and standard length for Centropyge potteri 22 2.4 Histology ofthe gonads of Centrovsge potteri 28 2.5 Plots ofthe relationships between proportion male score and standard length, and proportion male score and female size differences, for Centrovsge potteri 33 2.6 Reproductive behavior in Centropyge potteri 40 3.1 Encounter-rate relationships expected between the alpha female and other members ofher social group, as functions offish density; the encounter-rate threshold models 56 3.2 Plots of the relationships ofharem size, and territory size, over fish density for Centrovsge potteri 66 3.3 Simple linear regressions of alpha-female encounter rates over fish density for Centropyge potteri 69 3.4 Encounter rates in the enclosure experiments using Centropyge potteri 74 4.1 The polygyny-threshold model of Orians (1969) 85 4.2 The temporal-threshold model ofpolygynous mating 91 4.3 Temporal distribution ofbirths by African elephants, Loxodonta africanSJ., as related to rainfall........................................... 97 4.4 Spawning by haremic females ofCentrovsge potteri during April, 1989 102 5.1 Mean adjusted time of mating for Centropyge potteri on a daily time scale 118 5.2 Frequency of daily time of mating in ten-minute time intervals for Centropyge potteri 120 xi 5.3 Interference behavior by haremic CentrQP.Yge potteri females: enclosure 123 5.4 Interference behavior by haremic Centropy~ potteri females: field 125 5.5 Spawningorder as related to size rank in haremic Centropy~e potteri females 128 5.6 Frequency ofmating for CentrQP.Yge pQtteri during April, 1990 131 5.7 Avera~e number of spawns for individualCentropyge potten females during April, 1990 134 xii CHAPTER I GENERAL INTRODUCTION This dissertation is about problems in the behavioral ecology of reproduction in the pomacanthid angelfish, Centropyge potteri.
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