henderson etalk 2011 league food habits of three colubrid tree snakes (genus uromacer) on hispaniola

8/3/2019 Henderson Etalk 2011 League Food Habits of Three Colubrid Tree Snakes (Genus Uromacer) on Hispaniola

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Herpetologists' League

Food Habits of Three Colubrid Tree Snakes (Genus Uromacer) on HispaniolaAuthor(s): Robert W. Henderson, Albert Schwartz, Teresa A. Noeske-HallinReviewed work(s):Source: Herpetologica, Vol. 43, No. 2 (Jun., 1987), pp. 241-248Published by: Herpetologists' LeagueStable URL: http://www.jstor.org/stable/3892057 .

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Herpetologica,3(2), 1987,241-248C 1987by The Herpetologists'League,Inc.

FOOD HABITS OF THREE COLUBRID TREE SNAKES(GENUS UROMACER) ON HISPANIOLAROBERT W. HENDERSONI, ALBERT SCHWARTZ2,AND TERESA A. NOESKE-HALLIN'

'Section of VertebrateZoology,MilwaukeePublic Museum, Milwaukee, WI 53233, USA2BiologyDepartment, Miami-Dade Community College-NorthCampus,Miami, FL 33167, USAABSTRACT: Examinationof 687 specimensof threespeciesof Uromaceryielded 214 prey items.The slender-bodied, ong-snoutedspecies (U. frenatus and U. oxyrhynchus)are sit-and-wait for-agers and exclusivelysaurophagous;U. frenatus is a lizard generalist,and U. oxyrhynchusis anAnolis specialist. Uromacercatesbyi has a heavier body and blunter snout and exhibits sit-and-wait and active foragingmodeswhich are reflected n its diet (diurnallyactive lizardsandquiescenttree frogs).Uromacerfrenatus exhibitsgeographicvariationin diet, with Ameiva chrysolaemabeing ex-ploited more frequently on Ile de la Gonave,Haiti, than elsewhere in its range.Similarly,only U.catesbyi from Ile-a-Vache,Haiti, preyed on Leiocephalus. Although U. frenatus makes an onto-genetic shift from anoles to more robust terrestrial izards (Ameiva), the prey mass/snake massratio remainsunchanged.All three species of Uromacer, regardlessof morphologicalspecialization,eat those vertebrateprey items that are the most geographicallyand ecologically widespreadand are, relatively, themost abundant.Key words: Anolis; Diet; Hispaniola;Trophic ecology; Tree snakes;Uromacer

HISPANIOLA has therichest nake aunaof any West Indian island, and it is theonly island in the Greater Antilles withcolubrid snakesspecialized for an arborealexistence. Uromacer is endemic to His-paniola, where three species are nowrecognized, although there has been con-siderable taxonomic uncertainty until re-cently (Henderson and Schwartz, 1984a;Horn, 1969;Schwartz, 1979). Uromacerisone of three polytypic colubridgenera (theothers are Alsophis and Ialtris) occurringon Hispaniola, but it is the only genus inwhich members are sufficiently abundantthat an analysisof trophic relationscouldbe made.Uromacer catesbyi is the most widelydistributed of the three species on the His-paniolan main island and on at least sevensatellite islands. It is, compared with itscongeners, a heavy bodied snake [xT t. +1 SE for four snakes70-75 cm snout-ventlength (SVL) is 71.6 ? 7.3 g] with a widerhead and blunter snout. It is generallyfound in wooded habitats, including des-ert oases, but also occurs in more openAcacia forests. Uromacer frenatus is lim-

ited to the "south island" of Hispaniola(Williams, 1961), including five satelliteislands,occurring in a variety of woodedhabitats (xeric and mesic). It is a slendersnake (x wt. of 11 snakes 70-75 cm SVLis 24.8 ? 1.2 g) with an attenuated snout.Uromacer oxyrhynchus is widespread onthe historical"north sland,"but it has alsoinvaded the eastern portion of the southisland and at least three satellite islands.It too is elongated (x wt. of four snakes70-75 cm SVL is 33.3 ? 3.8 g) with anattenuated snout. Uromacer catesbyi isusuallyfound syntopicallywith one of thelong-snoutedspecies. At the few localitieswhere frenatus and oxyrhynchus havebeen taken together, neither species iscommon (Henderson and Schwartz, un-published observation). The two long-snouted species do not occur together onany of the satellite islands.Photographsofall three species appear in Henderson andSchwartz (1984b).Enough is now known about other as-pects of the biology of Uromacer [e.g.,morphology (Henderson, 1984; Horn,1969); internal topography (Martinez et

241


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242 HERPETOLOGICA [Vol. 43, No. 2TABLE 1.-Means ? 1 SE (in mm) of several morphological variables for three species of Uromacer, sexescombined. Sample size is given in parentheses. Different letters in a row indicate significant differences inmorphological variables among snake species (Student-Newman-Keuls test, P < 0.05).

Uromacer atesbyi Uromacer renatus Uromacer xyrhynchusSnout-vent length (SVL) 617 ? 14A 688 ? JJB 736 ? 15C(51) (103) (37)Midbodycircumference(MBC) 38.8 ? 1.8A 29.4 ? 0.9B 25.6 ? 1.Oc(20) (36) (17)Head width (HW) 9.7 ? 0.4A 7.3 ? 0.2B 7.9 ? 0.3B

(30) (47) (19)Head length (HL) 23.2 ? 0.8A 21.5 ? 0.9A 26.7 ? 0.7B(24) (25) (15)Snout anterior width (SAW) 3.0 ? OlA 2.3 ? O.PB 2.9 ? OJA(24) (47) (15)Snout base width (SBW) 6.2 ? 0.2A 5.1 ? 0.1B 5.9 ? OlA

(24) (47) (15)Snout length (SL) 7.1 ? 0.2A 7.8 ? 0.2B 9.6 ? 0.3c(24) (25) (15)Eye diameter (ED) 3.65 ? 0.1OA 3.2 ? 0.08B 3.70 ? 0.06A(24) (25) (15)

al., 1985); and foraging and prey captur-ing (Hendersonand Horn, 1983; Hender-son et al., 1982; Ulrich and Ford, 1985)],that it is now possibleto determine wheth-er the interspecific differences noted bythese previousstudiesare reflected trophi-cally. Here, based on examination of 687specimens, we describe and compare thediets of all three species of Uromacer,analyze the trophic contribution(in termsof number and volume of animals) of avariety of prey genera and species, andrelate diet to morphology, behavior andecology.

MATERIALS AND METHODSWe examined 258 U. catesbyi, 257 U.frenatus and 172 U. oxyrhynchus. Meth-ods used for dissections, determination ofprey volumes, and snake head and bodymeasurements have been explained else-where (Henderson,1982, 1984). We couldnot identify all prey items to species, norcould we determine volumes for all preyitems; therefore, there are more individ-ual prey items than there are prey vol-umes.The snakes were collected at many lo-calities throughoutHispaniolain both wetand dry seasons over a span of about 80years, and they are housed in collections

at the American Museum of Natural His-tory (AMNH), the Albert Schwartz FieldSeries in Miami (ASFS), the Museum ofComparativeZoologyat HarvardUniver-sity (MCZ), the Milwaukee Public Mu-seum (MPM), the Florida State Museumat the University of Florida (UF-FSM),and the National Museumof Natural His-tory (USNM).Body weight and corresponding SVLare available for 25 U. catesbyi and 76 U.frenatus. Also, live weight and corre-sponding volume are available for a va-riety of prey species. We use these data todeterminerelationshipsof predator o preymass in a portion of our sample, becausewe assume this to be an accurate measureof predator-preymass relationship.Data were analyzed using one-wayanalysisof variance (ANOVA),analysisofcovariance (ANCOVA), regression, andStudent'st-test. Differences among groupmeans were determined with Student-Newman-Keuls test. The alpha level ofsignificance is 0.05. Prey species volumedata were logarithmically transformedprior to analysis to normalize their distri-butions. All analyses were performed us-ing the StatisticalAnalyses Systems (SAS)package.Definitionsof abbreviationsusedfor morphologicalvariablesappear in Ta-ble 1.


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June 1987] HERPETOLOGICA 243TABLE 2.-Summary of prey species eaten by three species of Uromacer.For each prey species, the fre-quency of occurrenceis given (with frequency of calculable volumesin parentheses f different from totalfrequency). "X" indicates that a prey species was not exploited.

Prey species Uromacer atesbyi Uromacerrenatus Uromacer xyrhynchusFrogsHyla pulchrilineata 1 X XOsteopilusdominicensis 30(23) X XLizardsSphaerodactyluselegans X 1 XAnolis caudalis 2Anolis chlorocyanus 2 2 1Anolis coelestinus 4 8 (7) XAnolis cybotes 4 10 (7) 10 (9)Anolis distichus 9 (6) 4 (3) 13 (10)Anolis olssoni X 25 (23) 1Anolis semilineatus 1 2 4

Anolis sp. 9 25 5Leiocephaluslunatus 1 X XLeiocephalusmelanochlorus 3 (2) 5 (4) XLeiocephaluspersonatus X 1 XLeiocephalusschreibersi X X 2 (1)Leiocephalusvinculum X 4 (3) XLeiocephalussp. X 1 1Ameiva chrysolaema X 17 (15) XAmeiva lineolata X X 2Ameiva taeniura X 2 (1) 2Ameiva sp. X 1 1RESULTS

MorphometricsThere were significantdifferences in allmorphometric charactersamong Uroma-cer species (ANOVA,P < 0.001; Table 1).Uromacer oxyrhynchus was longest inbody, head, and snout; U. catesbyi wasthickest in body, head, and snout;and U.frenatus was intermediate between U. ca-tesbyi and U. oxyrhynchusin body thick-ness, SVL, and snout length (Student-

Newman-Keulstest, P < 0.05). Uromacercatesbyi had a significantly longer headrelative to total body length (SVL as co-variate) than U. frenatus (ANCOVA,F =6.46, P < 0.003). Head width relative tototal body thickness (MBC as covariate)was greater for U. oxyrhynchus and U.catesbyi than for U. frenatus (ANCOVA,F = 8.56, P < 0.0005).There were significant sexual differ-ences only for U. frenatus: females werelarger than males in all morphometricvariables(Student'st-test, P < 0.05). Thissex difference, however, did not affect theabove comparisons of morphologicalvari-ables among snake species.

DietPreserved specimens (687) yielded 214prey items (62 for U. catesbyi, 108 for U.frenatus, and 44 for U. oxyrhynchus). Ta-ble 2 andFig. 1 summarizeUromacerdiet.The slender-bodied, long-snouted speciesU. frenatus and U. oxyrhynchuswere ex-clusively saurophagous, whereas theheavier-bodied, blunt-snouted U. catesbyitook frogs and lizardswith equal frequen-cy (although frogs contributed a greatervolume of food than did lizards) (Fig. 1).Frog predation by U. catesbyi was vir-tually restrictedto the large hylid Osteo-pilus dominicensis, a geographically andecologically widespread species. Osteopi-lus contributed over 70%of total volumeof the diet of U. catesbyi, a larger per-centage than for any other single preyspecies eaten by any other Hispaniolancolubrid (Henderson, unpublished obser-vation). Several species of Anolis wereeaten by U. catesbyi, with A. distichusbeing the most frequently consumed andA. cybotes providing the greatest total(Fig. 1) and mean (3.6 ? 0.6 cm3) vol-umes. Leiocephalus melanochlorus was


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244 HERPETOLOGICA [Vol. 43, No. 2

50t U.oxyrhyrchus 1 Percent Frequency30 2 { Percent Volume10I30 frena usr

a10 < 77,ii .S10

0) 70 U.catestyi

3010 KtF1r$

Osteopilus Ano/is Anol/s Anonis Anonis Anon/i Leioceph. Leioceph. Ameivadomin. coenlest/is cybotes distichus onssoni semi/in. me/anoch/or vinculum chrysoniemaFIG. 1.-Prey species in the diets of three species of Uromacerby frequency and volume. Not all prey

species that were eaten are represented in the graph; domini. = dominicensis; semilin. = semilineatus;Leioceph. = Leiocephalus;melanochlor.= melanochlorus.eaten by U. catesbyi only on Ile-a-Vache,Haiti, and representedthe only non-scan-sorial species of prey taken by U. catesbyi.The Leiocephalus-eatingU. catesbyi werelonger than those U. catesbyi preying onother genera (ANOVA, F = 7.47, P


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June 1987] HERPETOLOGICA 245*Sphaerodaclylus c/egans* A.o scybotesIl e\A0 ^ A. distichLls ieaV c;0 A. olssoniVA. spp

Leowepf1aw1s spp. >*A.neiva ch,ysolaem-8EJ A. t- Uaeir X8 X GopOop

Ea) Grosse CayeSo*0

04 *

2 2 00 2> onUO

0 * o O 0? O 070 0OCA OV A0 0

40 50 60 70 80 90Snake Snout Vent Length (Cm)FIG. 2.-Relationship between body size (SVL)of

Uromacer frenatus and prey size (volume). Pointson the graphindicatingmost of the larger prey rec-ords were from satellite islandsoutlined with theirlocality of origin noted.

U. frenatus that ate Ameiva (x= 790 mmSVL) were significantly longer than thosethat ate Anolis (x? 667 mm) or Leioceph-alus (x = 699 mm) (ANOVA, F = 10.82,P < 0.0001). The prey volume consumedby U. frenatus depended on snake SVL,MBC, HW, SBW, and SAW (regression,P < 0.0001). Mean mass ratio for all preyitems was 12.5% range1.3-52.2%,n = 68).Leiocephalus spp. had the greatestweight(6.0 ? 1.4 g, n = 8) and the highest massratio (18.6%).Massratio for anoles rangedfrom 8.5%for A. distichus to 20.9%for A.coelestinus and A. chlorocyanus com-bined; all Anolis combined had a meanmassratio of 11.5%.Surprisingly,Ameivachrysolaema had a mean weight of only4.2 ? 0.5 g and a mean massratio of 10.8%(range 4.8-28.6%, n = 15). Most of thelargest prey items taken by U. frenatuswere from satellite island localities (Fig.2).Uromacer oxyrhynchus is the mosttrophicallyspecialized member of the ge-nus: Anolis comprised over 80% of its dietby frequency and volume, and A. cybotesand A. distichus combined comprisednearly 60%of its diet. As in U. frenatus,those U. oxyrhynchuseating Ameiva hadgreater mean SVL than those eating otherlizards(Anolis and Leiocephalus). Amonganoles taken by U. oxyrhynchus, the A.

cybotes were significantly larger than theA. distichus (ANOVA, F = 16.45, P 0.05).Among prey species that were eaten bymore than one species of Uromacer, onlyAnolis coelestinus showed a significantdifference in size being exploited: U. fre-natus ate significantly larger individualsthan U. catesbyi (ANOVA, F = 6.75, P 0.05).

DISCUSSIONAll three species of Uromacer are diur-nal, spending most of their time in lowbushes and trees. Various aspects of be-havior and ecology vary among the threespecies. Available evidence indicates thatU. oxyrhynchusis the most arboreal,rare-ly descending to the ground, exploiting

very slender perches, and feeding almostexclusively on scansorialprey (Hendersonet al., 1981, 1982). Uromacercatesbyi willmakeoccasional ong forays on the ground(Henderson, unpublished data; Hender-son et al., 1982), and by day, U. frenatushas been collected on the ground as oftenas in trees or bushes (Henderson, unpub-lished data).The prey species exploited reflect theforaging strategy of the three species ofUromacer. Uromacer frenatus and U.oxyrhynchus are exclusively sit-and-waitstrategistsand are therefore dependent onactive prey (e.g., Anolis, Leiocephalus,Ameiva) approaching near them. Once alizard is nearby, their elongated, stream-lined shapesallow them to move unobtru-sively over very slender branches with lit-tle disturbance while stalking. Likewise,their attenuated heads facilitate a muchwider field of binocular vision (Walls,1942) for accurate stalking and striking(Ulrich and Ford, 1985).Uromacercatesbyi,like U.frenatus andU. oxyrhynchus, utilizes a sit-and-waitforaging mode, but unlike the other two


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246 HERPETOLOGICA [Vol. 43, No. 2species, U. catesbyi also at times exhibitsan active foraging strategy. Very unusualmovements were observed in U. catesbyi:on six occasions at three different locali-ties, U. catesbyi was seen ascending intoor descending from the crowns of Cocospalms. These crowns provide cool, dampretreats, but perhaps more importantly,they provide daytime shelters for noctur-nal tree frogs. This active strategyenablesU. catesbyi to locate diurnally quiescentOsteopilus dominicensis. Uromacercatesbyi has a snout that is more atten-uated than most colubrids(HendersonandBinder,1980), and it does frequentlycatchactive prey, but not with the accuracy ofits more specialized congeners (UlrichandFord, 1985). In light of inefficient stalk-ing/capturing of active prey (anoles),for-aging for diurnally quiescent tree frogsbyU. catesbyi is, in terms of energy and ben-efits, a useful specialization.Heavy exploitationof tree frogsfor foodsets Uromacercatesbyi apart from its twocongeners. Osteopilus was a frequentsource of food for U. catesbyi, and onethat, on the average, was the largest andmost frequently-eaten prey species withthe highest mass ratio taken by any Uro-macer. Anoles taken by the heavier-bod-ied U. catesbyi were almost exclusivelytrunk-groundspeciesrather than the moredelicate grass anoles. Although it makesprolonged movements on the ground, U.catesbyi feeds less frequently on ground-dwelling lizards than either U. frenatusor U. oxyrhynchus, indicating that terres-trialforaysare not for foraging.Tree frogsare patchily distributed,so the effort thatmay be expended in locating one shouldbe frequently rewarded. Ascending intothe crown of a Cocos palm must, moreoften than not, result in food being locat-ed. The size of that prey item may notcorrelatestronglywith snake size, becausethe snake will attack a larger-than-opti-mum frog as long as it is physically ingest-ible.Uromacercatesbyi is considered, on thebasisof skull morphology and internal to-pography, to be the least specialized andmost primitive species of Uromacer (Mag-lio, 1970; Martinez et al., 1985). The

heavier body and blunt snout of U. cates-byi conform to a more "typical"colubridshape, whereas U. frenatus and U. oxy-rhynchus exhibit body and head mor-phology that is shared by only two othercolubrid genera (Ahaetullaand Oxybelis).Maglio (1970) suggested an Alsophis- orHypsirhynchus-like ancestor for Uroma-cer. Uromacer catesbyi is very similar inskull structure to Hypsirhynchus and isAlsophis-like in foraging mode and diet.It is more catholic in diet than long-snout-ed congeners, taking, like Alsophis, morethan one prey type (frogs, lizards). Ba-hamian and Greater Antillean Alsophisspp. are even more generalized, preyingon all groups of terrestrial vertebrates(frogs,lizards, snakes,turtles, birds,mam-mals). Uromacer catesbyi shareswith Al-sophis the trait of taking multiples of Os-teopilus spp. froglets (up to six by U.catesbyi and 26 by A. vudii; Henderson,unpublishedobservations).Just as the high frequency of Osteopi-lus in the diet of U. catesbyi sets it tro-phically apart from its congeners, so toodoes the high incidence and volume ofAmeiva chrysolaemain the diet of U. fre-natus set it apart, but only in one sectionof the latter species' range. The high in-cidence of Ameiva in U. frenatus stom-achs is, in our sample, restricted to Ile dela Gonave, Haiti (cf. Hendersonand Horn,1983, for a detailed analysisof the diet ofU. f. dorsalis). Likewise, most of the U.frenatus that had eaten Anolis olssoniwere also taken on Ile de la Gonave. Onthe Hispaniolanmain island, U. frenatuspreys primarily on Anolis (and to a lesserdegree on Leiocephalus and Ameiva), soits diet here is very similar to that of thelargely allopatric U. oxyrhynchus.Although large U. frenatus take largerprey items than small U. frenatus (a com-monplace phenomenon in snake feeding)and, at least on Gonave, make a radicalshift (expansion) in diet from scansorialanoles to ground-dwelling Ameiva, themean mass ratio of Anolis to Ameiva issimilar (11.5% vs. 10.8%). Thus there islittle or no change in mass ratio with in-creasing snake size. We suspect that thisis not an unusual occurrence in snake


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June 1987] HERPETOLOGICA 247feeding ontogeny. Whether prey handlingtime increases with larger prey (as we sus-pect it does with anoles versus large treefrogstaken by U. catesbyi) and affects netenergy gain is unknown. Greene's (1984)excellent discussion touches upon this.Anoles, however, because of their taperedand streamlinedshape, would presumablybe easier to consume even if their masswere equal to that of a medium-sized Os-teopilus. All three species of Uromacertake prey that (except in the case of preyspecies that are small as adults: e.g.,Sphaerodactylus, Anolis olssoni) is gen-erally much smaller than large adults ofparticularprey species.For example, largeOsteopilus dominicensis may reach aweight of 65 g and a volume of 75 cm3;Anolis cybotes 10.5 g and 12 cm3; A. coe-lestinus 17g and 16 cm3;Ameiva chryso-laema 48 g and 52 cm3. Some, like largeOsteopilus, A. coelestinus, and A. chryso-laema, may be too large to ingest by aUromacer, or even dangerous to attack(e.g., Ameiva);or large individuals of preyspeciesmay utilize partsof the habitat dif-ferent from smaller conspecificsand fromforaging Uromacer.The evidence available suggests thatUromacer evolved from a diurnal, semi-arboreal, euryphagic ancestor, probablyclose to Alsophiscantherigerus. Extant U.catesbyi is the most morphologically, be-haviorally and ecologically generalizedUromacer (Horn, 1969; Maglio, 1970).Both morphologies (i.e., heavy-bodied U.catesbyi and streamlined U. frenatus andU. oxyrhynchus) impose behavioral andecological constraints. Uromacer catesbyimay be prohibitedfrom using the slenderperch diameters exploited by its conge-ners,and branch diameter becomes a lim-iting factor when stalkingalert,active preysuch as anoles. On the other hand, theslender, streamlined morphology of U.frenatus and U. oxyrhynchus may imposeconstraints on prey size and type. Al-though U. catesbyi exploits a food re-source that apparently takes more energyto locate, the energy gained per averageprey item probablyoutweighs the energyspent. Henderson (1984) concluded that,among Hispaniolancolubrids,there was a

trend toward trophic specialization, ndthat same trend appearswithinthe genusUromacer.Uromacer atesbyi s the mosttrophically generalized in taking bothfrogs and lizards; U. frenatus exploitedthe most prey species, but they were alllizards i.e., it is a lizardgeneralist);Uro-maceroxyrhynchuss a lizard specialist,preyingpredominantly ponAnolis.Onthe mainislandof Hispaniola,Uro-macer renatusand U. oxyrhynchus reyon anoles (Anolis chlorocyanusand itssouth slandallospeciesA.coelestinus,andA. cybotesandA. distichus) hat are geo-graphicallyand ecologicallywidespread.Although population densities are notavailable, ll of theseanolesarehighlyvis-ible and abundant; hey are, in fact, themost common anoles on Hispaniola, ustas 0. dominicensis s themostwidespreadand most frequently-seenarboreal an-uran.Speciesof Uromacer,whetherspe-cializedor not, eat thosevertebratepreyitemsthat are mostreadilyavailable.Sim-ilar evidence of opportunisticpredationhas been observed n Hispaniolanboids(Henderson t al., 1986b)and other His-paniolan colubrids (Henderson andSchwartz, 986;Henderson t al., 1986a).

Acknowledgments. -We are grateful to museumpersonnelwho have allowedus to examinespeci-mens:RichardG. Zweifel AMNH), os6P. Rosadoand Pere Alberch (MCZ), Peter Meylan and WalterAuffenbergUF),andW.RonaldHeyerandGeorgeZug USNM).We benefittedromcomments n ear-lierversions f thispaperby GaryS. Casper,HenryS. Fitch,MaxA. Nickerson nd RichardA. Sajdak.Most f Henderson'secent ieldworkon Hispaniolahasbeen undedbyFriends f the Milwaukee ublicMuseum. raigDethloff ssistedwithdissectionsndChristineGenich ypedseveraldraftsof the manu-script, and their contribution is appreciated.

LITERATURE CITEDGREENE, H. W. 1984. Feeding behavior and dietof the eastern coral snake, Micrurus fulvius. Pp.147-162. In R. A. Seigel, L. E. Hunt, J. L. Knight,L. Malaret, and N. L. Zueshlag (Eds.), VertebrateEcology and Systematics-A Tribute to Henry S.

Fitch. Mus. Nat. Hist., Univ. Kansas Spec. Publ.10.HENDERSON, R. W. 1982. Trophic relationships andforaging strategies of some New World tree snakes(Leptophis,Oxybelis, Uromacer).Amphibia-Rep-tilia 3:71-80.. 1984. The diets of Hispaniolan colubrid


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snakes. I. Introduction and prey genera. Oecologia62:234-239.HENDERSON, R. W., AND M. H. BINDER. 1980. Theecology and behavior of vine snakes (Ahaetulla,Oxybelis,Thelotornis,Uromacer):A review. Con-

trib. Biol. Geol. Milwaukee Pub. Mus. 37:1-38.HENDERSON, R. W., M. H. BINDER, AND R. A. SAJ-DAK. 1981. Ecological relationships of the treesnakes Uromacer catesbyi and U. oxyrhynchus(Colubridae) on Isla Saona, Republica Dominicana.Amphibia-Reptilia 2:153-163.

HENDERSON, R. W., B. I. CROTHER, T. A.NOESKE-HALLIN, A. SCHWARTZ, AND C. R.DETHLOFF. 1986a. The diet of the Hispaniolancolubrid snake Antillophis parvifrons. J. Herpe-tol.: In press.

HENDERSON, R. W., J.F. DROUGHT, AND A. D. KATZ.1982. Daily activity patterns of the Hispaniolantree snakes Uromacercatesbyiand Uromaceroxy-rhynchus (Colubridae). Ann. Mtg. Soc. StudyAmphib. Rept. and Herpetol. League (Abstract),p. 78.

HENDERSON, R. W., AND H. S. HORN. 1983. Thediet of the snake Uromacer frenatus dorsalis onIle de la Gonave, Haiti. J.Herpetol. 17:409-412.HENDERSON, R. W., T. A. NOESKE-HALLIN, J. A.

OTTENWALDER, AND A. SCHWARTZ. 1986b. Onthe diet of a Hispaniolan boa, Epicrates striatus,with notes on E. fordi and E. gracilis. Amphibia-Reptilia:In press.HENDERSON, R. W., AND A. SCHWARTZ. 1984a.

Uromacer frenatus. Cat. Am. Amphib. Rept.357.1-357.2.1984b. A guide to the identificationof theamphibiansand reptilesof Hispaniola.MilwaukeePub. Mus. Spec. Publ. Biol. Geol. 4:1-70.1986. The diet of the Hispaniolancolubridsnake Darlingtonia haetiana. Copeia 1986:529-531.HORN, H. S. 1969. Polymorphismand evolutionofthe Hispaniolansnake genus Uromacer(Colubri-dae). Breviora324:1-23.MAGLIO, V. J. 1970. West Indian xenodontinecol-ubridsnakes:Theirprobableorigin,phylogeny,andzoogeography.Bull. Mus.Comp. Zool. 141:1-54.MARTiNEZ, D. R., J. A. LUCIO, AND A. SCHWARTZ.1985. Topografiainterna de las culebras del ge-nero Uromacer(Colubridae).Caribaea1:48-58.SCHWARTZ, A. 1979. The statusof Uromacerfre-natus and Uromacer dorsalis (Reptilia: Serpentes:Colubridae).Herpetologica35:207-215.ULRICH, N., AND N. B. FORD. 1985. The feedingstrike of three species of vine snake, Uromacer.Soc. Study Amphib. Rept. and Herpetol. League(Abstract),p. 81.WILLIAMS, E. E. 1961. Notes on Hispaniolanher-petology.3. The evolutionand relationshipsof theAnolis semilineatus group. Breviora 136:1-8.WALLS, G. L. 1942. The VertebrateEye and ItsAdaptiveRadiation.Bull. Cranbrook nst. Sci. 19:1-785.

Accepted: 18 March 1986Associate Editor: Arthur Dunham

henderson etalk 2011 league food habits of three colubrid tree snakes (genus uromacer) on hispaniola

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