nutritional value of plants consumed by the white-tailed deer in a tropical forest of mexico

8/7/2019 Nutritional value of plants consumed by the white-tailed deer in a tropical forest of Mexico

1/5

NUTRITIONAL CHARACTERISTICS OF PLANTS CONSUMED BY THEWHITE-TAILED DEER IN A TROPICAL FOREST OF MEXICO

GABRIELA S/LVA-VILLALOBOS, C.U.C.BA, Universidad de Guadalajara, Apartado Postal 39-82, Zapopn 45110,Jalisco, Mxico. (Tel y Fax: 3-682-0230)SALVADOR MANDUJANO, GLORIA ARCEO, SONIA GALLINA, Departamento de Ecologia y ComportamientoAnimal, Instituto de Ecologa A. C., Apartado Postal 63, Xalapa 91000, Veracruz, Mxico. (Te/: 28-421800, Ext.4110; Fax: 28-187809; ENVIAR CORRESPONDENCIA A Correo-E: [email protected])LUIS ALFREDO PREZ-JIMNEZ, Estacin de Biologa Chamela, Universidad Nacional Autnoma de Mxico.Apartado Postal 70-233, Mxico, D. F. 04510, Mxico. (Tel: 5-622-5692; Fax:5-550-1760; Correo-E:[email protected])ABSTRACTThe objective 01' this s tudy was to determine the nutri tional value 01'sorne plant species con sumed by white- tailed deer (Odocoi!eusvirginianus), as well as o ther spec ie s abundan t in the understory 01't ropica l deciduous fores t (TDF) and tropica l semi-evergreen fores t(TSF). bu t not consumed by deer. The study was carried out at theCharnela Biologica l Sta tion on the coast 01' Ja li sco , Mxico. FromOctober, 1994 to August, 1995, bimonthly determinations were made 01'the percentages 01'crude protein, nitrogen-free extract(NFE). crude fiberand total polyphenols present in 11 species 01' plants consumed by thedeer, and 21 unconsumed species (10 from TDF and 11 from TSF) . Wefound that plants consumed by deer had more NFE, protein andpolyphenols and less f iber than unconsumed plants. TDF plants had ahigher nutritional value dur ing the rainy season than dur ing the dryseason. TSF plants were higher in f iber and lower in prote in and NFEthroughou t the year . F iber was correl at ed negat ively wi th NFE andprotein.KEY WORDS: Mxico, nutritional value, Odocoileus virgimanus,seasonal variatlOn, tropical deciduous forest, tropicalsemi-evergreen forest, white-tailed deer.RESUMENEl objetivo del estudio fue determinar el valor nutriclOnal de algunasespecies que consume el venado cola blanca (Odocoileus vlrgInzanus)y de especies no consumidas pero que son abundantes en el sotobosquedel bosque tropIcal caducifolio (BTC) y del bosque tropical subperennifolio (BTS) de la Estacin de Biologa "Chamela" en la costa del estadode Jalisco, Mxico. Se determin bimensualmente, de octubre de 1994a agosto de 1995, los porcentaj es de protena cruda, estracto libre den it rgeno (ELN) , fib ra cruda, y pol ifenoles to tal es de 11 especiesconsumidas por el venado y 21 especies no consumidas (10 del BTC y11 de BTS) Encon tr amos que las plantas que consume el venadotuvieron ms protena, ELN y polifenoles, y menor fibra que las plantasno consumidas. Las plantas del BTC tuvieron un valor nutricional altoA herbivore's diet is l imited, among other factors, bynutrient availability, toxin content, ingestion and digestionrates and by the time invested in searching, consuming andruminating food (Hanley 1997). The white-tailed deer(Odocoileus virginianus) is c1assified among those ruminantsthat select very nutritious plants which are high in cellularcontent, ferment rapidly, and are easily digested (Van Soest1982). Since its rumen is small compared to body size, deer

38

durante la poca de l luvias y menor durante la poca seca. Las plantasdel BTS tuvieron mayor porcentaje de fibra y menor de protena y ELNdurante todo el ao. Seencontr correlacin negativa signficativaentreel porcentaje de fibra con el porcentaje de proteina y ELN de las plantas.PALABRAS CLAVE: bosque tropical caducifolio, bosque tropicalsubperennifolio. Mxico, Odocoi/eus virginia-

nus. valor nutricional, variacin estacional.venado cola blanca.

RESUMOO objet ivo des te estudo foi determinar o valor nutri tivo de algumasespcies de plantas consumidas pelo veado (Odocoileus virginianus).assim como de outras espcies abundantes da floresta tropical caduciflia (FTC) e floresta tropical subpereniflia (FTS), mas nao consumidaspelos veados. O estudo foi realizado na Esta;;ao Biolgica Chamela. nacosta do estado de Jalisco, Mxico. Determinou-se bimensalmente, deoutubro de 1994 a agosto de 1995, as porcentagens de prote na crua,extrato Iivre de nitrognio (ELN), fibra crua, e polifenois totais de 11espcies consumidas pelo veado e 21 espcies nao consumidas (10 deFTC e 11 de FTS). Encon tramos que as p lan ta s consumidas pelosveados continham mais protenas. ELN e polifenois, e menor fibra queas plantas nao consumidas. As plantas de FTC tinham um valornutri tivo al to durante a poca de chuvas e menor durante a poca seca.As plantas de FTS tinham maior porcentagem de fibra e meno r deprotena e EL durante todo o ano. Encontrou-se correla;;ao negatIvae significativa entre a porcentagem de fibra e a porcentagem de protenae ELN das plantas.PALAVRAS-CHAVE floresta tropical caduciflia, tloresta tropicalsubpereniflia, Mxico, Odocoileus virgInia-

nus. valor nutritivo, varia;;ao estacional, veadosVIDA SILVESTRE NEOTROPICAL 8(1-2):38-42must compensate for low rumino-reticular capacity bychoosing such high-quality foods (Henke el al. 1988, Shortel al. 1974). Nutritional value depends al so on such factorsas plant parts, life form, and community type (Blair el al.1977, Vangilder el al. 1982, McCullough and Ullrey 1985).Therefore, knowledge of the nutritional value of the vegetation available to deer would allow a better understanding ofthis herbivore's foraging strategies.


2/5

METHOS

Table 1. Species consumed and unconsumed during the raand dry seasons inclued in the nutritional analysis (Ttropical deciduous torest and TSF-tropical semievgreen torest).

range from 10-25 m in height. The most common treesAslronium graveo/ens Jacq., Brosimum alicaslrum Sw., aSideroxy/on capiri A. DC. Mangroves , palm groves , athorny secondary scrub are also found surroundingstation. Field work was restricted to the 500 ha ofthe statreserved for research, and o f this area, over 75 % was TD

Due to time and cost considerations, we decided to wwith only 32 plant species, despite high species richnesthe understory ofthe study area and high species diversitdeers' diet (Arceo 1999). We selected some o f the plamost frequently consumed by deer (as determined by diobservation) as well as other unconsumed species that anevertheless, very abundant and found almost exclusively

Seaso

XXXXXXXXXXXXXXXXXXXXX

X XX XX XX XX XX XX XX XXX

TS F Ramy

XX

XX

XXXXXXXXXX

UnconsumedConsumed TDF

Dicliptera resupmala (Vahl) Juss. XCaesalpinia eriostachys Benth. XAcalypha sp. XCroron sp. XSpondias purpurea L. XHeliocarpus pa/lidus Rose XCoursetia caribaea (Jacq.) LavinvaL caribaea X

Desmodil/m procl/mbens(MJ1!) Hitchc X

TephroslO sp XSerjania brachycarpa A Gray X[,,"O.)'lml"'l alicastrum Sw XLasiaos rusclfolia (HBK.)Hitchc vaL ruscifoliaJacql/ll1la pl/ngens A. Gray

Trie/ullO trifolia L. subsppalmen (C DC ) Pennington

Lonchocarpus lanceolatus BenthE ~ v t r a n a Imbricara (Vahl) Pers.ApoplalleslO panicl/lta PreslHvballtlllls mexicanus GingO'oton sl/berosl/s HBKDecachaeta haenkeana DCEl/patoril/m solidagil1l{olil/m A. Gray.Thollll1ldl1lm decandrum

(Humb & Bonpl) RadlkCvnolllelra oaxacana BrandegeeForchhalllmena pal/ida LiebmROl/rea glabra HBKCupal1la denlata DCC ~ v t o s f o l l l a binafllm (Thunb.) Sandw.A d e n o c a ~ l ' I l l a inl/ndatum Mart. ex DCCappars verrl/cosa Jacq.COlllbretumfruticosl/m (Loetl) StuntzPodopterus cordi{oll1ls Rose & Stand!Gl/aplta Illacrocarpa Miranda

The white-tailed deer is a major herbivorous species intropical forests in the Pacific coast region of the state ofJalisco, Mexico, with population densities ranging from 10to 14 deer/km 2 (Mandujano and Gallina 1995a). In thisregion, the precipitation pattern, as well as topographic andsoil condit ions (Bul lock 1986) cause tropical deciduousforest to predominate in hilly areas, while tropical semievergreen {orest dominates riparian sites. These vegetationcommunities are different from each other in tloristiccomposition (Lott el al. 1987), phenology (Bullock andSolis-Magallanes 1990), and biomass (Martnez-Yrizar el al.1992). As a consequence, deer are faced with spatial andtemporal variation in the availabil ity of food, water, andcover (Mandujano and Gallina 1995b).

This work is part of a research project aimed at discovering individual and population strategies employed by whitetailed deer in response to habitat heterogeneity in the tropicalforest of Chamela, Jalisco, Mexico. An important factor toconsider in investigating these strategies is the variation inthe nutritional value of vegetation, and how such variationmight affect the selection o f plants species included in thediet (Arceo 1999), as well as intluence the activity andhabitat use patterns o f deer (Snchez-Rojas el al. 1997,Mandujano 1999). The specific objectives of the presentstudy were: a) to determine the nutri tional values of plantsconsumed, or not consumed, by deer in tropical deciduousand semi-evergreen forests, and b) account for any seasonalvariations or differences in these values between plantspecies.STUOY AREAThis study was conducted at the Chamela Biological

Station, operated by the Universidad Nacional Autnoma deMxico, and located on the southwestern coast ofthe state o fJalisco, Mexico. The station covers 3,600 ha and haselevations ranging from 30 to 500 m. The region is characterized by an irregular topography. Mean annual temperatureis 25C, with May through September being the hottestmonths. From 1977 to 1984, maximum and mnimumtemperatures ranged from 29-32C and 15-23C, respectively. Mean annual rainfall is 748 mm (SD= 119 mm), with80% falling between July and October. Rains normally occurfrom July to November, while the dry season lasts fromDecember to June (Bullock 1986).The dominant vegetation at the station is tropical dry

forest (TDF), located on hilly terrain with thin soils. Forestheight varies between 4 and 15 m, and a well -developedunderstory is usually present. Most tree species lose thei rleaves during the dry season (Lott el al. 1987). Some ofthemost common trees found are Cordia a/fiodora [Ruiz &Pav.] Oken, Lonchocarpus /anceo/alus Benth., and Caesa/pi-nia erioslachys Benth. The station also has tropical semievergreen forest (TSF), which is typically found alongstreams and in protected places with deep soils. These forestVIDA SILVESTRE NEOTROPICAL 8(1-2):1999 DEER FORAGE QUALlTY IN TROPICAL FOREST Silva-Villalabas et al.


3/5

Table 2. Percentages of nutri t ional characterist ics of sorneconsurned and unconsurned plants bywhite-tailed deerduring the rainy and dry season in the tropical forest ofCharnela, Jalisco, Mxico. TDF ( t ropical deciduousforest), TSF (tropical sernievergreen forest). Mean (standar error).

UnconsumedNutritional Content Season Consumed TDF TSFCrude protein rainy 17.0 (0.3) 16.4 (O 1) 152 (0.1)

dry 11.4 (15) 126 (14) 12.9 (0.7)Nitrogen-free-extract ramy 48.9(07) 47.3 (06) 419(0.3)

dry 492 (13) 45.3 (09) 44.3 (0.3)Crude fiber rainy 210 (09) 23.4 (1.4) 29.3 (2.0)

dry 27.3(4.3) 30.3 (3.6) 29.4 (29)Total poliphenols ramy 2.5 (O 1) 1.7 (02) 16 (0.1)

dry 3.0 (O 1) 15 (01) 2.1 (02)NutritlOnal mdex ramy 3.1 2.7 2.0(protein+NFE)/fiber dry 2.2 1.9 2.0

the TDF or TSF understory (Table 1).We sampled various plants of each species a long deer

trails and dirt roads ofthe biological station. Only leaves andtender branches of plants were collected. Two hundredgrams of each species were collected, and samples wereoven-dried at 50C for 2 to 5 days. To prevent heat fromvolatizing sorne of the secondary compounds, 50 g of eachsample were air-dried. Later, they were ground separately ina Wiley mili with a 20-mesh screen, tagged and stored inbags.Sampling was carried out in October of 1994 (mid-rainy

season), December of 1994 (late rainy season), February of1995 (early dry season), April of 1995 (mid-dry season), Juneof 1995 (late dry season) and August of 1995 (early rainyseason). Each plant species from each location, sampledduring each season, was homogenized and analyzed for theirnitrogen-free extract (NFE), crude protein, and crude fiberontents, using A.O.A.C. (1975) methods. Total polyphenolswere determined by the method ofLesli-Harti and Johnstone1989). AII determinations were done in triplicate except forpolyphenols, which were done in duplicate.We used the arcsin function to transform data which wereinally expressed in percentages (Sokal and Rohlf 1969).frequency distribution of transformed data was normal

d variances were homogeneous, therefore we used paraetr ic tests. For each one of the nutritional determinationsrude protein, NFE, crude fiber and polyphenols), we testedr seasonal differences among the three groups of plantsunconsumed plant species from TDF and TSF, as well asant species consumed by deer). We applied analyses ofriance (Sokal and Rohlf 1969) and for significant resultse employed the Student-Newman-Keuls a posterior iltiple comparison test to determine differences between

groups. To determine possible associations between nutritional var iables , we performed simple linear regressionanalyses for all pairwise combinations of variables.RESULTSAnual mean percentage of crude prote in was 14.2% in

plants consumed by deer, 14.5% for plants from the TDF and14.0% for plants from the TSF. These differences were notsignificant (F=O.13, df=2, 16, P=0.88). However, there wasa significant difference in mean percentage of proteinbetween seasons (Tab le 2, F=33.8, df=l, 16, P


4/5

34 ~ - - - - - - - - - - - - - - - - - - - - - , fiber content of plants increased. This can be explainenoting that as plants matured, their cell wall constituiincreased and their digestibi li ty decreased (Short e1974). The fiber content (mainly lignin) is known tgreater in leaves of evergreen species than in leavedeciduous species (Prudhomme 1983). This suggestsbecause many species in the TSF keep their leaves yround, they have developed mechanisms to reducepossibility ofattack by herbivores. Therefore, during theseason forage quality and abundance is low in both vegetnal communities, and deer must modif)r their forastrategies (Arceo 1999).The biochemical content of plant species consumeddeer during the rainy season was higher in nutritional vashowing a greater percentage of NFE and protein conand less fiber, than in unconsumed species. This suggthat in order to allow the accumulation of fat reserves fomating season and for the critical dry se::son, deer must seplants which are more nutritious (Vangilder el al. 1982).estimates of crude protein contents (10-17%) could potially cover the requirements (6-16%) for the growthdevelopment of fawns and adults , as well as for the gro

ofmale antlers (Asleson el al. 1996, UlIrey el al. 1967).low fiber content and high percentage of NFE in the pspecies consumed throughout the year suggest thatprefer species of high cel lular content. Deer obtain a hpercentage of digestible energy from cellular content (Amann el al. 1973, Thompson el al. 1973), especially duthe dry season when food availability is very low.Due to the astringent and indigestible nature of tan(Robbins el al. 1987), we expected deer to select specieswere low in polyphenols. At this site, however, deer seleplants with a higher percentage of polyphenols, wcompared with the content in unconsumed plants. Howethis percentage covers total polyphenols, and includinconsiderable amount of other phenol ic compounds whichdifferent from tannins. In particular, it will be necessarquantif)r the percentage of hydrolyzable and condentannins. Because the former hydrol ize under gastricconditions and liberate trapped vegetable proteins (Robel al. 1987). However , deer are adapted to tanninferfodders, having tannin binding proteins in their saliva (AlIel al. 1989). These aspects will be necessary to study infuture in this tropical forest.MANAGEMENT IMPLlCATIONSThe dry season is the critical per iod to manage wh

tailed deer in this t ropical forest, due to the low qualityscarcity offood, as well as lack ofwater. The actual poption density (l0-14 deer/km2 ) is below carrying capa(Mandujano 1999); therefore, there is no apparent needprovide additional food under normal circumstanHowever years ~ i t h low rainfall result in reduced forproduction. Therefore, it is necessary to implement habmanagement practices. One possibility would be to com

50

44 46 48nitrogen-free-extract ( % )

unconsumed TDF .. . unconsumed ~ S f J

22 .-0 - ' - -


5/5

ment native food sources with nutritionally-balanced food aswell as sufficient water, at specific places within the forest,principally when the winter rains are scarce. This is especially import an t because diminished winter rains usuallyresult in a lower standing plant biomass and less availablewater in subsequent dry season (Mandujano y Gallina1995b). Another important management practice in this typeof habitat is the maintenance and encouragement of thosetree species whose fruit are preferred by deer, because theyrepresent an important source of energy, protein and waterduring the dry season.ACKNOWLEDGMENTS

The fol lowing people provided valuable assistance andlabonitory analysis of samples: L. Barrientos-Ramrez, M.Barri en tos-Ramos, M. Cas te ll anos , G. Lpez-Ortiz , L.Villarino, and M. Torres. The Consejo Nacional Ciencia yTecnologa (CONACYT) provided S. Mandu jano and G.Arceo with a grant for postgraduate studies. The CONACYT(P220CCOR-892154), CONACYT (P020CCOR-903703),SEP (DGICSA-902467), and CONACYT (0327N9107)projects provided financial support. The Biological Stationof "Chamela" provided all the facilities for this project.L1TERATURE CITEDAMMANN, A. P., R. L. COWAN, e. L. MOTHERSHEAD and B. R.BAUMGARDT. 1973. Dry matter and energy intake in relationto digestibility in white-tailed deer. Journal of WildlifeManagement 37:195-201.AOA.C 1975. Official methods of analysis of the Association

of Official Analytical Chemists. 12th ed. Association ofOfficial Analytical Chemists. Washington, D.e., USA. 1094pp.ARCEO, G. 1999. Hbitos alimentarios del venado cola blancaen el bosque tropical caducifolio de Chamela, Jalisco. Tesisde Maestra. Facultad de Ciencias, Universidad NacionalAutnoma de Mxico, Mxico, D.F., Mxico. 66 pp.ASLESON, M. A., E. e. HELLGREN and L. W. VARNER. 1996.Nitrogen requirements for antIer growth and maintenance inwhite-tailed deer. Journal of Wildlife Management 60:744752.AUSTIN, P. J., L. A. SUCHER, e. T. ROBBINS and A. E. HAGERMAN.1989. Tannin-binding proteins in saliva of deer and theirabsence in saliva of sheep and cattle. Journal of ChemicalEcology 15:1335-1347.BLAIR, R. M., H. L. SHORT and E. A. EpPs. 1977. Seasonalnutrient yield and digestibility of deer forage from a youngpine plantation. Journal of Wildlife Management 41 :667676.BULLOCK. S. H. 1986. Climate of Chamela, Jalisco, and trendsin south coastal region ofMexico. Archives for Metereology,Geophysics, and Bioclimatology 36:297-316.. and J. A. SOLs-MAGALLANES. 1990. Phenology of~ p trees of a tropical deciduous forest in Mexico.Biotropica 22:22-35.HANLEY, T. A. 1997. A nutritional view of understanding andcomplexity in the problem of diet selection by deer (Cervi-

dac). Oikos 79:209-218.HENKE, S. E., S. DEMARAIS and .J. A. PFISTER. 1988. Digestivcapacity and diets ofwhite-tailed deer and exotic ruminantsJournal of Wildlife Management 52:595-598.LESLl-HARTI, F. and F. H. JOHNSTONE. 1989. Anlisis modernode los alimentos. Editorial Acribia, Zaragoza, Espaa.Lon, E. J., S. H. BULLOCK and 1. A. SOLs-MAGALLANES. 1987Floristic diversity and structure of upland and arroyo forestsin Coastal Jalisco. Biotropica 19:228-235.MANDUJANO, S. 1999. Ecologa del venado cola blanca en unbosque tropical caducifolio de Jalisco. Tesis Doctoral.Faculdad de Ciencias, Universidad Nacional Autnoma deMxico. Mxico, D.F., Mxico. 142 pp.___ . and S. GALLINA. 1995a. Comparison of deer censusing methods in a tropical dry forest. Wildlife SocietyBulletin 23:180-186.and . 1995b. Disponibilidad del agua para el---enado cola blanca en un bosque tropical caducifolio deMxico. Vida Silvestre Neotropical 4: 107-118.MCCULLOUGH, D. R. and D. E. ULLREY. 1985. Chemicalcomposition and gross energyof deer forage plants on theGeorge reserve, Michigan. Natural Resources TechnicalInformation. Research Report No. 465. Michagan StateUniversity, Agricultural Experiment Station, East Lansing,Michigan, USA. 19 pp.PRUDHOMME, T. 1. 1983. Carbon allocation to antiherbivorecompounds in a deciduous and evergreen subartic shrubspecies. Oikos 40:344-356.ROBBINS, e. T., T. A. HANLEY, A. E. HAGERMAN, O. HJELJORD, D.

L. BAKER, e. e. SCHWARTZ and W. W. MAUTZ. 1987. Roleof tannins in defending plants against ruminants: Reductionin protein availability. Ecology 68:98-107.SNCHEZ-RoJAS, G., S. GALLINA and S. MANDUJANO. 1997.reas de actividad y uso del habitat de dos venados colablanca (Odocoileus virginianus) en un bosque tropical de lacosta Jalisco, Mxico. Acta Zoologica Mexicana (nuevaserie) 72:39-54.SHORT, H. L., R. M. BLAIR and e. A. SEGELQUIST. 1974. Fibercomposition and forage digestibility by small ruminants.Journal of Wildlife Management 38: 197-209.SOKAL, R. R. and F. 1. ROHLF. 1969. Biometry. W. 1-1. Frccmanand Co., San Francisco, CA, USA. 832 pp.THOMPSON, e. 8. , J. B. HOLTER, H. H. HAYES, H. SILVER and W.E. URBAN, JR. 1973. Nutrit ion of white-tailed deer. l.Energy requirements of fawns. Journal of Wildlife Management 37:301-311.ULLREY, D. E., W. G. YOUATT, H. E. JOHNSON, L. D. FAyand B.L. BRADLEY. 1967. Protein requirement ofwhite-tailed deerfawns. Joumal of Wildlife Management 31 :679-685.

VAN SOEST, P. 1. 1982. Nutritional ecology ofthe ruminant. O& B Books, Inc., Corvallis, Oregon, USA. 374 pp.VANGILDER, L. D., O. TORGERSON and W. R. PORATH. 1982.Factors int1uencing diet selection by white-tailed deer.Journal of Wildlife Management 46:711-718.

Associate Editor: Juhani OjastiReceived: June 9, 1997

Accepted: September 3, 199942 DEER FORAGE QUALlTY IN TROPICAL FOREST Silva-Villa/abas et al. VIDA SILVESTRE NEOTROPICAL 8(1-2):1999

nutritional value of plants consumed by the white-tailed deer in a tropical forest of mexico

Documents