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Licht and Gorman's (1970) RE PRODUCTIVE AND FAT CYCLES IN CARIBBEAN ANOLIS LIZARDS BY PAUL LICHT and GEORGE C. GORMAN UN I VERSITY OF CALIFORNIA PRESS BERKELEY· LO SANGELES ' LONDON 1970 UNIVERSln ' OF CALIFORNIA PUBLICATIONS IN ZOOLOGY AOVISORY EOITOkS: G, A. BARTHOLOM EW, 1. H. C01<·1NELL. J OHN DAVIS, C . R . GOLDMAN, C,\D ET H A"0, K. S. NORRIS, O. P. 1'f.ARSO:-l, R. 1-1 . ROSENBLATT, GROVER STEPHENS Volume 95 Approved for publication June n . 1970 Issued October 23, 1970 Price $2.00 UNIVERS ITY OF CAUFORNIA P RESS U ERKELEY AND Los A1\;C ELES CALIFORNIA U"IVERSlTY OF CALIFORNIA PRESS, LTD. LoNDON , ENCLAND ISB N: 0-520·09374-7 LUmANY OF CONCRess C.. TAJ.OC CAHD No. : 72-632162 © 1970 Il l' T ilE REGENTS OF Til E UN IV ERSITY 01' CALIFORNIA JlR I :\'TEIl IN TilE U:\, ITED STATES OF AMERICA CONTENTS Introd uction . 1 :Materials and Methods 3 Collection, preservation, and autopsy of spec imens 3 ~'at storage 3 Reproductive status of males 4 Rep r oductive status of females 5 Cl imatological influence on reproduetion 7 Results of Studies on St. Vincent . 8 Geography, climate, and species . 8 Reproductive and fat cycles of Anolis tnnita tis and A. richard; grise .... 9 Resu lts of Studies on Jamaica . 12 Geography, climate, and species . 12 Reproductive and fat cyclcs of A. gmha",i, A. lineatopus, and A. sagre; 13 Results of Studics on Hispaniola . 22 Gcography, climate, and species . 22 Reproductive and fat cycles of A . cybotes 23 Resul ts of Studies on Grand Cayman 25 Geography, climate, and species 25 Reproductive and fat cycles of A. consp ..·slls 27 Results of Studies in Florida. 28 Geography, climate, and species . 28 Reproductivc and rat cyclcs of A. sag,·e; 28 , Results of Studies on Bermuda . 30 Geography, climatc, and specics . 30 Reproductive and fat eyclcs of A. graham; 31 Discussion Existence of reproductive cycles in Anolis . 33 Problems in thc interpretation of the reproductive data 36 Fat cycles in Anolis 37 Environmental correlations with reproductive and fat cycles 40 Conclusions regarding envit'onmental influences on reproduction 46 Sum lnary . 47 Acknowledgments 48 Literature Cited 49 REPRODUCTIVE AND FAT CYCLES IN CARIBBEAN AL~OLIS LIZARDS uy PAUL LTCHT alld GEORGE C. GOR~iAN ( A contribution from the l)cparlmrnt of Zoology nnd the Museum of Vertebrate Zoology of the University of California, Berkeley, California) L'lTRODUC1'ION A WEALTH of data has been accumulated over the past few decades on annual rep roductive activities in temperate-zOllO lizards. Sufficient species have been studied to permit broad gelleralizations regarding variations in reproductive pat­ wrns of temperate species (c.g., St. Girons, 1963; Mayhew, 1968; Licht et aI., 1969) and considerable progress has bcen made toward understanding the mccha­ nisms underlying these cycles. Howeyer, understanding of saurian reproductive strategies is still limited by a paucity of phcnological data for tropical lizards. Although the tropical lizards represent thc richest part of the liza rd fauna, com­ parative data for these arc insufficient to allow any conclusions regarding the natur e of variations in annual reproduetivity or the factors that might be re­ sponsible for such variations where thcy ex ist. The present investigation repre­ sents a preliminary attempt to gain such comparative information on these im­ portant aspects of the ecology of the widely distributed neotropicailizard genus, Anolis_ Anolis is a suitable genus for comparative reproductive studies. There are nu­ mcrous species (approximately 250) and the systema tics of a large proportion of these are well understood. Many are very abundant, hence easily collected. In addition, perhaps more is known about the reproductive cycles of one member of this genus, A. carolill clIsis J both in nature and in the laboratory, than of any other lizard. Anolis is widely distributed in Mexico, all of Central America, the northern I half of Sonth America, on virtually all Caribbean islands, and Oil the continental Un ited States. 1'he genus is di"ided into two major groups, a and fJ (Etheridge, 1960 ), based upon osteological differences, and each of these groups is divided into species series. We have chosen seven species from various islands in the 'Vest Ind ies for intensive study. The species and localities included permit numerous comparisons to be made. (1) Anolis grahami, a nati,·e of Jamaica, has been successfully introduced onto Bcrmuda to the north. Populations from both localities have been sampled, to allow comparison of the same species under rather different environmental con­ dit ions_ (2) A,lOlis conspersu~ is a close relative of grahami on an adjacent island. (3) Anolis lineatopus is symputric with A. grahami in Kingstan, Jamaica; thus sym patric congeners can be compared. (-l) Anolis sagrei is found on the western half of Jamaica, and is compared with thc two preceding species from a different part of that island, as well as with a population of sagrei that lives in Florida. (5) Anolis ,-ichardi griseus and A, t,·initatis are cndemic to the island of St. Vincent. [ 1 I 2 Univ.,'sity of California Publications in Zoology j '0° ' 0° 70° , r o .00, ! MILES U,S.A. - B ERMUDA ATLANTIC OCEAN 20° 20° ~ . , PUERTO .. • RICO -, • .~ ~ 'I'. ~ ... ST. VINCENT I..., , " VENEZUELA Fig. 1. Map of the Caribbean region showing collecting sites of the areas involved in this study. Licht and Gorman: Anolis Reproduct;ve and Fat Cycles 3 .t\gai~ 've can compare sympatl'ic congeners, and can compare these lizards with relatIves on more northcrly islands that experience greater variation in photope­ riod and differcnt rainfall patterns. (6) Anolis cybotes, an cndemic of Hispaniola, is at the same latitudc as Jamaica but experiences different rainfall patterns and thus ?rovides an additional source of comparison. Figure 1 shows the geographical relatIons among the Yarious habitats included in this study. The taxonomic brcakdown of thc species studied is as follows: Gnon' RERIES SPECIES .\lphll fati{ro1!S A. trinilutis, A. ridardi Alpha cl'istatellus A. cybotcs B{'tn flrallami A. gralwmi, A. lineatopw, iJ.. conspcrtrus HeIR sagrri . 1. sagre .. MATERIALS AND METHODS COLLJ.:CTIOX, PHESEUr.\'l'IOX, AND AUTOPSY OF SPEcrMENS Samples of most species werc collected at approximately monthly intervals oyer a one- to two-year period with the assistance of indh'idnals stationed in the va rious localities. Jamaican collections of A. grahami, A.. lineatopus, and A. sagrei we re supervised hy lIfr. Ethon Lowe; St. Vincent collections of A . tr;n;tatis and A. richardi by Mr. Michael DcFrcitas; Bcrmuda collections of A. graham; hy Mr. Dav id Wingatc; Florida collections of A . sagre; by Mr. Phil Baker; and Grand Cayman collections of A. conspersus by Mr. John Gray, Jr. Freshly col­ lected specimens of A. cybotcs from Port-au-Prince, Haiti in 1968-1969 were obtained through Georges Hene; Mr. J. Borno supplied a fcw collections in 1966. Data for other Hispaniola localitics wcre obtained from collections in the Museum of Comparative Zoology at IIanard University through the courtesy of Dr. E. E. Williams. Dr. J. Wril(ht of the I,os Angelcs County Museum and Dr. H. Campbell of the Florida State Museum, University of Florida, provided numerous specimens of A. sagrei from southern Florida and adjacent islands. :\1ost specimens were fixed in 10 percent buffered formalin shortly after collec­ tio n; a few of the A. cybotcs collected in 1966 were obtained alive ahout a week after collection, and the A. sayrci from Florida were preserved in Bouin's fixative. Museum spccimcns of cybotcs and sagre; were stored in alcohol and the method of preservation was unknown. Upon receipt in the laboratory, formalin fixed speci­ me ns were transferred from 10 percent to 4 percent formalin for at least a day rnd t h e n rinsed in running tap watcr fo r a fcw hours before autopsy. Each indi­ 'idua l was mcasured (snollt-to-yent length) and examined for nutritional and rep roductive condition as indicated below. Testis weight was found to vary with the lnethod of fixation: FOl'malin caused an increase of about 16 percent and Bouin's solution causcd a rcduetion of about 15 percent of fresh weight. Thus, testis weights cannot bc compared among all species but wcights should be con­ sistent for species obtained from each collcctor. FAT STORAGE The weights of the paired abdominal fat bodies were taken as an index of encrgy balance; i.e., whether or not fat storage was occurring. Studies on Anolis caro- 4 University of California Publications in Zoology linensis by Dessauer (1955a) have shown that the scasonal variations in fat body size generall y reflect changes in total hody lipids, the fat bodies acconnting for about half of the increase in body lipid with the remainder bcing divided between the liver aJld carcass (Licht and J ones, 1967 ). We initially measured liver weight, but variability due to fixation made these weights too unreliable and this mcasure· ment was discontinued. In gcneral, monthly mean hepatic weights tended to vary in parallel with the wcights of the abdominal fat bodies. In most samples, we found that the weight of the fat bodies was independent of body size (length) within the range of sizes used (sec below). Therefore fat body weight is expressed as an absolute value. REPRODUCTrvE STATUS OF MALES The rcproductive condition of the males was jndged by the weight of the testis an d by the histological appearance of the germinal epithelium and accessory scx· ual structures (epididymis and sexual segmcnt of the kidney). Autopsy pro· cedures were the same as those described in previous studies on Anolis in this laboratory (sec Licht, 1967a) .
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