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Genera Thalassoma, Halichoeres and Hemipteronotus)

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Genera Thalassoma, Halichoeres and Hemipteronotus) STUDIES ON THE FAUNA OF CURAÇAO AND OTHER CARIBBEAN ISLANDS: No. 138. Color as related to Size, Sex, and Behavior in seven Caribbean Labrid Fish Species (genera Thalassoma, Halichoeres and Hemipteronotus) by Machteld J. Roede (Universiteit van Amsterdam) Contents Page Figure/Plate Table Abstract 5 Acknowledgements 6 I. INTRODUCTION The phenomenon of color dimorphism ... 7 reversal 8 Color, sex, and sex Two types of males 9 Senescence 11 Scope of present study 11 Material 12 II. TAXONOMY 1 Thalassoma bifasciatum 16 Halichoeres bivittatus 18 Halichoeres garnoti 19 Halichoeres maculipinna 20 Halichoeres poeyi 21 Hemipteronotus splendens 22 Hemipteronotus martinicensis 23 III. GENERAL INFORMATION 24 Curasao Puerto Rico 25 2 Page Figure/Plate Table IV. ABUNDANCE, ENVIRONMENT, AND DISTRIBUTION Abundance and environment 27 2 J uveniles 31 Bathymetrical distribution 31 Migration and hibernation 33 Geographic distribution 34 V. METHODS Collecting 38 1 Preparation of samples 42 Observations 42 VI. COLOR Introduction 44 Terminology 44 Color production 45 Changes of color 46 in Dichromatism the literature .... 48 Present investigations 50 Observations on color change 50 Description of color phases 51 Thalassoma bifasciatum 53 2 Halichoeres bivittatus 57 3 Halichoeres garnoti 61 4 Halichoeres maculipinna 65 5 Halichoeres poeyi 70 6 Hemipteronotus splendens 73 7 Hemipteronotus martinicensis ... 77 8 Summarizing remarks 78 3-4 VII. MERISTIC DATA AND MORPHOMETRY Fin and 83 5-6 spines rays Morphometry 86 Methods 86 Results 88 7 Allometry of growth and size ... 88 9 8-9 Sex differences 94 Geographic differences 95 Specific differences 95 Shape of fins 96 10, 11 10-16 Conclusions 106 VIII. BEHAVIOR Locomotion and Orientation 108 Wrasses at night 112 Food and Feeding habits 114 Reproductive behavior 119 Territorial defence 125 Comments 126 3 Page Figure/Plate Table IX. REPRODUCTIVE ORGANS Methods 129 Sex 130 Stages of gonadal activity 131 Estimated size of gonads 132 Measured length and width of gonads .135 17-18 Weight of gonads 136 Histology 136 Ovary 137 i_iv 19 Testis 143 V-VI 20 Intersexes 146 VII-IX X. RELATION BETWEEN COLOR, SIZE, AND SEX History 148 21 Present investigations 155 12-20 22 Color and Size 159 23 Color and Sex 163 Sex ratio per species 163 24 Sex and color phase 175 25 Females 176 Males 178 Miscellaneous 179 Conclusions 180 26 XI. GONADAL ACTIVITY Gonadal activity per color phase 183 27 Gonad and total weight length .... 188 21 Summary 189 Gonadal activity and time of the 190 28-29 year ... Gonadal activity and lunar month 197 22-23 30-32 KII. NON-FUNCTIONAL GONADS, IMMATURES AND SENESCENCE; INTERSEXES AND SEX REVERSAL Present investigations 205 Immature gonads 205 Young ovaries 206 Young testes 207 Gonads in reduction 207 Ovaries in reduction 207 No visible gonads 208 Testes - in reduction senescence . 209 . Intersexes 211 Sex reversal 212 Remarks 214 Conclusions 216 History 217 and Sex reversal Intersexuality .... 217 Mechanisms of sex determination and differentiation 220 ........... 4 Page Figure/Plate Table XIII. GROWTH AND HORMONE EXPERIMENTS Growth studies 224 24-25 33-34 Hormone experiments 231 History 233 Present investigations 234 X-XI Conclusions 241 XIV. CONCLUDING REMARKS AND SUMMARY .... 242 XV. REFERENCES 251 EXPLANATIONS OF PLATES 263 Present addres of author: Institute of Human Biology, Utrecht. Abstract Approximately 5500 Caribbean labrid fishes belonging to seven species were The caught, transported, partly onice, to the laboratoryand subsequently studied. examined 20 cent stored majorportion was immediately; some per were briefly (less than three days) at minus 20°C. Color of adult from pattern (and shape fins) may change during life, varying slightly Halichoeres poeyi) to drastically Thalassoma bifasciatum). Per species a classification into successive color phases has been introduced; these color phases to are strongly related body length. In Halichoeres maculipinna, Hemipteronotussplendens and Hemipteronotus martini- to adult color males have censis females are restricted the small, first phase; ex- clusively been found in the larger intermediate and terminal color phases. This that confirms the prevailing opinion labrid dichromatism represents a sexual di- morphism. In Halichoeres poeyi females are also significantly smaller than males. In the most abundant species, however, sex and color/size are not clearly related. In Thalassoma bifasciatum and Halichoeres bivittatus two types of males occur, as functional males are present in both the small, first adult color phase and in the large,terminal color phase. Functional females have been found in all sizes and colors in Halichoeres bivittatus and Halichoeres garnoti. in all - Sex reversal from female towards male - sex a common process species to occurs more or less coincidental color change. As distinct from the Sparidae, Scaridae or Serranidae the reversal proceeds via a total decline of the transforming of functional do not occur ovary. Consequently, temporary stages hermaphroditism in Labridae. ACKNOWLEDGEMENTS With sincere I wish to thanks to Prof. Dr. H. ENGEL and gratitude express my Mrs. M. found Dr. S. J. ENGEL-LEDEBOER who always time to discuss my text. Dr. E. interest in has JACK RANDALL'S my doings been encouraging; only a man of his make it torealize talents could possible an experimental study with unmanageable underwater Dr. INGVAR director of CARMABI his huge cages. KRISTENSEN, the and PAPY staff made the period on Curasao unforgettable. Mr. RAMRATTAN MARTINEZ has been an excellent assistant in collecting labrids. I am greatly indebted to Dr. J. HUIZINGA, who permitted me to continue this "reversal" from frank study notwithstandingmy a marine to a human biologist. The and stimulating criticism of my work offered by Dr. G. W. F. EDGAR is greatly appreciated.Many members of the staff of the Institute of Human Biology, Utrecht, valuable effort write Mrs. have given their help to my to this thesis, particularly ANNE TOL. Jos RUTING, VINUS VAN BAALEN and the technical staff of the Zoological made the thanks H. Laboratory, Utrecht, drawings; special are due to Mr. VAN KOOTEN for his expert contribution to the plates. I am gratefulto Dr. P. WAGENAAR HUMMELINCK and Drs. LOUISE J. VAN DER STEEN for their patience and their consci- entious treatment of Mrs. L. W. GEBHARDT-THURBER kind my manuscript. was so as to clear the final text of a number of language inaccuracies. Part of the statistical work has been performed by the Afdeling Waarnemings- uitkomsten, T.N.O. The present study was supportedby a grant of the Netherlands Foundation for the Advancement of Research in Suriname and the Netherlands Antillen (WOSUNA). I. INTRODUCTION Labrid dichromatism has been subjected to numerous investi- gations leading to controversial and sometimes confusing results. In an attempt to clarify the inconsistencies outlined below, the between and interrelationship color, size, sex, gonadal activity was studied in approximately 5500 specimens belonging to seven labrid from the The distribution in different species Caribbean waters. sex color phases and the possibility of sex reversal have been the main points of investigation. THE PHENOMENON OF COLOR DIMORPHISM The of the - the Labridae - is of fishes family wrasses a group and colors. showing a most diversified variation of beautiful bright Color within be than color differences a species may more striking differences between from the of species. Juveniles may differ adults which in show distinct color the same species, turn may two patterns (dichromatism). the In some species dichromatism is so pronounced that color variants be the were considered to two different species. Thus, species pairs Thalassoma nitidum and Th. bifasciatum, Labrus ossifa- Coris gus and L. mixtus, giofredi and C. julis, Stethojulis renardi and S. well varius and G. tricolor strigiventer as as Gomphosus were only 8 each recently recognized as belonging to one single species, viz. Thalassoma bifasciatum, Labrus bimaculatus, Coris julis, Stethojulis strigiventer and Gomphosus varius respectively (LONGLEY, 1914; LÖNNBERG & GUSTAFSON, 1937; BACCI & RAZZAUTI, 1957; RANDALL, 1955a; STRASSBURG & HIATT, 1957). this list far dichromatism Probably, is from complete because may have been in other For this not recognized as such species. instance, may be pertinent to wrasses enlisted for the Indo-Australian archi- pelago (DE BEAUFORT, 1 940) in which no dichromatic species have been described. Recent studies in the area of the Indo-Pacific by RANDALL (MS) could pinpoint a dozen incorrectly classified di- unite chromatic pairs of wrasses. He, e.g. proposes to Anampses into the rubrocaudatus and A. chrysocephalus one species under last name. Color dimorphism is not present in every labrid species. For in- stance, it does not occur in Ctenolabrus rupestris (QUIGNARD, 1966). In dichromatic labrid species dull colors are found often in small whereas individuals, large specimens display a gaudy pigmentation. In a few cases the exact relation between color and size has been analyzed (LÖNNBERG & GUSTAFSON, 1937; REINBOTH, 1957). Also the of the fins alter when sizes In shape may larger are attained. some species the caudal fin changes from a truncatetowards a lunate form. In various lost with cases either elongate dorsal spines get growth (young razor fishes) or develop ( Coris julis). Some razor fish are characterized increased first ventral species by a strongly ray when they are bigger. How far changes in color and shape run parallel, has hardly
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