diversity of diet of white-tailed deer in a mexican tropical forest

8/7/2019 Diversity of diet of white-tailed deer in a mexican tropical forest

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Diet diversity of white-tailed deer(Odocoileus virginianus) in a tropicaldry forest in Mexico

Glo ria ARCEOSalvador MANDUJANOSon ia GALLINADepartment of Biodiversity and Animal Ecology , Instituto de Ecologia A. C.,Km 2.5 Carret. Ant. a Coatepec No. 351, Xalapa 91070, Veracruz (Mexico)

Luis Alfredo PEREZ-JIMENEZInstitut e of Biol ogy , Universidad Nacional Aut6noma de Mexico,

Apartado Postal 70-233 (Mexico)

Arceo G , Mandujano S.. Gallina S. & Perez-J imenez L. A. 2005. - Diet divers ity o f whitetailed dee r (Odocoi/eus virginianus) in a tropical dry forest in Mexico. Mamm :.l/ia 69 (2) :159-168 .

KEYWORDSdiet divcrsitv,seasonal var iatio n,Odocoileus vi rgi nid1J/fS,tro pical Io rcsr,Mexico .

ABSTRACTW/e report on diversity in rhe d iet ofwh i tr tai led deer (OducoiLcu, virginian li S)in a Mexican tropical dr y forest . Diet was estima ted using the microhisio logical analysis of plant epid ermis in deer pellet-groups. T he sam pling was fromJ993 ro 1994. From rhc rainy [Q dry season, rhc richn ess and diversiry of rhe[un ilies and specks in the diet increased and bot ani cal composition changed.Deer selected 82 planr species from 20 famili es; however, 12 species rep rescn rcd 50% of the annua l di et. f lowers and fruit s of arboreal species wereimpo rranr duri ng the dry season, as were young leaves of sh rub and vinespecies during the rainy season. Desp ite the small size of white-railed deer inrhis tropi cal dr y foresr study area, foraging st rategies there were sim ilar rochose observed in remperare forests.REsUMEDi uersirc de l'alimentation du w f de Virginie (OJocoileus virginianus) dan'une JO rh tropical au Mrxique.Cc r a rric lc prcscntc [a d ivcrs itc de l' a l i rncn rnr io n du ccrf de i : . i (Odocoilem virgil/ianus) dans une foret rropica lc de I'f:tar de Mexiquc . Leregime alime nraire a ere csrimc par l'analysc rnicro-hisiologiquc de l'cpid crrncdes planres da ns lcs f'eces de certechanrillou necs penda nt la saison des pluies,la sa iso n de transit ion er la saison seche en 1993-1 () ') 4 . I.a richessc cr la diversire des fami lies er especcs dan s le regime alirnenraire a augrnenu ' de la saisonseche a la saison des plui es, avec un changcmcnr de la com position botaniquc,

tv1AMMALIA 2005 69 (2) Publica tions Scientifiques du Museum national d' Histoire naturelle. Paris. 159


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MOTS CLEsdiversire alirncnraire,variation saiso nniere,Odocoilcus " j rgit/ il ll / lIs,fo ret tropicale.Mexique.

Les ce rFs o nr cbois i 82 es pcccs app;lrrenan, a 20 Famil ies, J 2 cspece:seulernen r consriruanr 50 % du regime a lirnc n ra irc. Flcurs, fruits cr l l e d 'arbres craicnr irnpo rranrs pendant la saison sechc, jeunes fC lIilles de buissoner especes rampanres pendant [a saison des pluics. Malgrc la raille corporcllcplus perire du cerFd e: Virginic da ns la Fo rer rropicale scchc de l'crude, lesstrategies alirnenraires apparaisscnr asscz semblables a celles des population des forers tcmp erces,

INTRODUCTIONBodysize is an impor tant factor in rhe dererrn ina-tion of ruminant ecology (Dernrnenr & Van Soesr1985; Hanley 1997). The white-t ailed deer(Odocoilcus virginianus) has a wide geographicaldistriburion from Sou thern Ca nada ro Norrh ernBrazil and Central Bolivia (H all 1981; Eisenberg1989); in consequence, size and body weighr varynota bly (Smith 1991) . In co n trast ro remperatehabirars in the northern region, where male andfemale mean weight (\X') is 100 kg and GO kg,respectively, in tropical habitarsweighrcan be 50%lower (Geist 1998). As me tabo lite requi remen tsscale co \ '(10.7 5 ;JoJ rumen volume ro W U ) (Shorreta!' 1969 ), rhen smallerdeershould require higherqual ity d iet rhan those inhabiring remperare area(Hanley 1997). Among ruminants, the whi terailed deer is considered an oppor tuni stic concenrrare feede r (Mysrerud 1998 ), selecring a greard iversiry offood and choosi ng relatively nutririous,highly d igesrib le plan ts and plane parrs such asyoung leaves, twigs, and saplings (Vangilder et 11.1982; Henke eta!' 1988).I n trop ica l habi tats, white-tailed de er inhabitqui te var ied vege ra rio n rypes such as t rop icalsava nnah, mangroves, flood for est, mounrainhabirars ar upper elevations, tropical d ry forest,seco ndary vegerarion bord ering rainforest , o ldseconda ry for est, ope n woodland s, an d fields(Brokx 1984; Mend ez 1984) . Unlike temperarehab i ra t s whe re rh is deer spec ies fac es seve rescarcity offood in win ter, in several tropical habirats the cr it ical pe riod is th e dr y season , whenplant abundance and quality decrease. In consequ ence, it has been assumed tha t dee r compen-

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sa rc for rh is seasonal limirarion by select ingspecies and plant communiries rhar offer the bestqu ality of food. However , few studies of whi terailed deer dicr have been und ertaken in tropicalfor est s (e.g., Branan et a!' 1985; Granado 1989;DiMar e 1994 ). Because this species is an imporra n r food reso urc e for loca l pe o ple (e .g . ,Mandu jano & Rico-Gray 199 I ) , ir is irnpo rran rco define managemenr policies rhar mainrain susrai nab le po pula tions in th e tropi cs (We ber &Gonzalez 2003) . Therefore, rhe study of foragingstrategies is key co und ersranding deer survival introp ical habitats.T he white-railed deer inhabi ts seasonally tropicaldr y fo rests on rhe Pacific Coast o f th e stare ofj ali sco , Mexico, wirh popularion densities ranging from 10 ro 14 deer per km2 (Mandujano &Ga llina 1995). T his fores t is characterized by amarked variation in plane phenology, due principally ro seasonality in rain fall parrern s that dividerhe year in to a rainy and dry season, each lasting6 ro 7 months (Lorr eta !' 1987; Bullock & 501isMagallanes 1990). In parti cular, plane richness,b iomass, and quality vary notably in the und ersto ry rhroughout rhe year (Silva-Villalobos et 1 1.1999) . To berrer understand rhe pop ulation ecology of rhe deer (Ma ndujano et al. 2002 ), we condu cted rhe presen r study, focusing on how th issmall subspecies (40 kg of body weighr) faces seasona l and spatial variations in food resources.The objective of the presenr paper is ro add newinformation about irs diet in tro pical forest andto discuss the relat ionship between feed ing srrategies and rhe body size of white-railed deer comparing wit h o rhe r st ud ies in north and so uthlatitud e.

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F,G. 1. - Loca lization of the "Chamela" Biological Stat ion in thePacific coast of the Mexican state of Jalisco . Tile shady rectangle of the inferior map represents the study place.

STUDY AREATh e s t udy was co nduc ted a t t he C hame laBiological Station, ope ra ted by the UniversidadNacional Autonoma de Mexico. T he stat ion is3 ,600 ha and located on ]alisco southwest coast(Fig. 1). Fieldwork was restricted to the 500 ha ofth e station rese rved for research . Mean annualtemperature was 25C (Bullock 1986) . Hottestrnonrhs arc [rom May to September. From 1977CO J997 , the me an an n ual precipitati on was748 mm (SD = 119) , with 80% of the rain fallingbetween Ju ly and O ctober. T he dom inant vegetation is trop ical dry forest located on hilly ter rainwith th in so ils and low wa ter reten t ion. Manytree and sh rub species lose their leaves during thedry seaso n. Common trees are Cordia alliodora,Lonchocarp us [anceo latus, a nd Caesalp in iaeriostachys (Lorr et al. 1987) . T he stat ion also hastropi ca l semi- evergreen forest , covering 2'1% of

MAMMALIA 2005 69 (2)

the area mainly along rhe major streams in deepsoil with high water retent ion. In th is habitat lesstree species lose their leaves dur ing the dry seasoncompare to th e dry fore st habitat. The most common rrees in thi s type of for est a rc Astron iumgraveofens, Brosimum alicastrum, and Sideroxyloncaplri.

METHODSMICROHISTOLOGICAL ANAl.YSISFresh pell et gro ups were obtain ed during therainy (August and O ctober of 1993), transition(N ovember and December of 1993), and dry seasons (April and May of 1994 ). They were collected along fou r tropical dry Iorest transects andtwo in tropi cal semi-evergreen forest. Each transec t was 400 In long ; ar eve ry J0 rn, a circularplot mea su rin g 10m ! was marked oFf (Manduj ano & Gallina 199 '5 ). A to tal of 243 faecalgroups were collected. Anthony & Sm ith (1974)indicate a sample size of 15 to 50 pellet groups asthe sugges ted minim um required to determi nedier on a seasonal basis.T he use of chloral hyd ra te to di scolo r and setsamples is, acco rdi ng to Middleton & Sanc hezRojas (1994 ), the best rechn iqu e to de rermineherbivo re di et. C hloral hyd rat e is, however, aresrricred drug in Mexico. In the present study,th is substance was therefore replaced, following as lig h tl y modified ve rs io n of rhc rec bn iqu cdescribed by Quin tanilla et al. (19 88). It co nsisred of placing the sample in sod ium hydroxidero '5%, boiling it fo r 5 minutes, rinsing it wirhdistilled water, and add ing sod ium hypochlor ite;ir was th en lefr to stand unr il complete d iscoloration had taken place. Th e sample was rinsedwi th distilled wat er and placed in alcohol so lutions, each with d ifferent concentrations, and lefrto stand for 20 minutes until complete dehydratio n had occurre d. T he samp le was rhen mixedwieh Xy lol in o rd e r co st a in t he r iss ues .Subsequently, a da b of Canad ian balsam mixedwith a porrion of rhe treated rnare rial was placedon each slide. For eac h plant species , va rioussl ides were pr epared in an effort to garher th e

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38% between rainy and dry, and 36% betweenrransirion and dry seasons .The annual diet was composed of 82 speciesCrable 1). Dier dive rsity increased significantlyfrom rlie rainy ro dry season (Fig . 2, t 4 .06,d .f. = 60, p 0 .00 I) . Dier similarity was 22%between rhe rainy and rransitio n seasons , 16%between the rainy and dry seasons , and 26%between rhe t rans ition and dry seasons . In partie-ular, of rhe 4\ species consumed during the rainysea son, .36 species were rare 5% comb ined ) ,while only five (Acalypha langiana, Cardia-spcrmum halicarabum, Coursetia caribaea, andtwo underermined Co nvo lvul acca e andLegurninosae) constituted 57 % of th e diet duringrhis season. Of rhe 65 species consumed duringrhe transition season, 59 were represenred by lessthan 5% of planr epidermis in th e sample, wh.ileonly six species (Acalypha langiana, Croton sp.,Abutilon sp., ,iyenia rnicranta and two underer-

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min ed Convolvulaceac and Leguminosae) consri-ruted 49% of rhe d ier in rhis sea son . Of th e61 species co ns u m e d durin g rhe dry se a so n ,57 made up < 5% of the p lanr ep id e rm is in rhesample , while only four species (Spondias pur-pu rea, Rourca tlabra , Cardiosperm um haLica-cabum, and one und ererrnincd Am ar.mth aceae)constituted 38% of the diet in rhis season .Life forms were signiflcanrly differenr among


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Imporra nr in the rainy Season, while the fr uitsan d th e leaves of trees an d bu shes were irn porrant in the d ry seaso n. U nfor runa rely, sp ecificdat a o n the co rporal weigh t of deer are no t provided in each analyzed study, thus was no t possib le to mak e a detailed analysis in th is top ic. Inconcl us io n, the overall re sults sugges t th atdespite the relat ively sma ll size o f wh ite- taileddeer in th e stud ied trop ical dry fo rest, thi s deerbehaves as oppo rtunis tic co ncentrate se lectorwhich is simi lar to the forage stra tegy in temperate hab itats.

AclmowledgementsThe Consejo Nacio na l C ie nc ia y T ecnologi a(CONACYT) p rovided study grant to G . Arceo.T he proj ects CONACYT (P 2 2 0 CCOR 892 154) , CONACYT (P020CCOR-903 703 ) ,SE P (D G TCSA -90246 7 ) , and CO NACYT(0327 N9 107 ) provided financial support. T heCharnela Bio logical Sta tion, Insriruro de Biologiao f the Universidad Naciona l Auto no rna de Mexico, provided logistic suppon an d facilities. W ealso thank to G . Silva-Villalobos, R. E. Sanche zManti lla, C. M. Ga lan , C. Pacheco and S. LeonTorr es fo r t heir help durin g t he fie ld an dlabo rato ry work . T he fol lowing peo ple co llaborated in the identification of herba rium samples:t= . C hiang , M. Sousa, O . Tel lez and ]. L. Vi llasenor. Also, we thank to C . Aguilera-Reyes andG . Gomez-Belt ran their help d uring the microhyscological analysis. Early versio n of this manusc r i p t was read by A. Go n za le z- Ro me ro ,V. San chez-Cord ero , Y. Hoerrelano, L. Leon,and J. Murcia. Lastly , two anonymo us reviewersprov ided sugges t ions rh a t im proved th e fi nalversion of this paper.

REF ERENCESANTHONY R. G . & SMITH N. S. 1974 . - Comparisonof rumen and fecal analysis (Q describe deer diets .I iu: Manage. 38: 535 -540 .BODMER R. E. 1989. - Frug ivo ry in AmazonianArt iod act yla: ev ide nce for th e evolu t io n of theruminant stomach. j. Zool. Land. 219: 457167 .

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I Arceo C . r t a].HANLEV T. A. 1997. . A nurririona l view of under-sranding and comp lexity in rhe problem of d ierselecrion by deer (Cervidae). Olleos 79: 209 -218.H ENKE S. E., DEMA JUS S. & PFISTER ] . A. 1988.

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1.01"1 ' E. ] ., BULLOCK S. H . & SOLis-MACI\I.I .JINI.S j. A.1987. - Florisric d iversi ty and str uctu re of uplandand ar royo Forests in Coa sta l ] alisco . Biotropica 19:228-235.MANDUJANO S. & l L S. 1995. - Com parisonof deer cens using me thods in a tropical d ry Forest.Wilell. Soc. Bull. 23: J80- 186.tvlANDUjl\1\O S. & RJell-GRAY V. 1991. - Hunri ng,use an d know ledge of th e b iology of the whi retailed deer , Odocoilcus uirginianus (Hays) , by rhemaya of cen tral Yucatan , Mexico. )' Etbnob iol. I I :175-183 .MANDUJANO S., GALLINA S. & BULLOCK S. H. 1994 .- Frugivo ry an d d ispe rsal of Spondias purpurea(Anaca rdiaccac) in a t ropical dry fo rest of Mexico .Rev. Biol. trop. 42: 105-1 12.MIINDuJflNo S., GALLI!',\ S. , ARCEO G., SANCHEZROJAS G . & S ILVA-V1Ll.ALOBOS G . 2002 . Vc nado cola b lanca (Odocoilew I'irginianus sinaloae]. A. Allen , 1903), in NOCUEI0\ F. A., AVAI )\ R. &GARciA-Aw fU :TE A. N . (cds), Histo ria Natu ral rleChamelet. Univcrsid ad Nacio na l Auro norn a d eMexico, Mexico, D . F.: 4 15-422.MENIH:Z E. 1984 . - Mex ico and Central America, inH ALL (ed.), Vl hite-Tailed Deer: Ecology and Management. Srackpole Books, Harr isburg: 'i 13-524 .M ILJ Dl.ETON B. A. & SANCHEZ-ROJAS E. 1904 . Microhisrological analysis of the Food habits of herbivores ill the tro pics. Videt Silvestre Ncotropical :3:4 1-47

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MVSTERUD A. 1998. - T he relative roles of body sizan d Feed ing ryp e on acrivi t y rime o f rempe ra r

diversity of diet of white-tailed deer in a mexican tropical forest

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