Diet diversity of white-tailed deer

(Odocoileus virginianus) in a tropical

dry forest in Mexico

Glo ria ARCEO

Salvador MANDUJANO
Son ia GALLINA
Depa rtmen t of Biodiversi ty and Ani ma l Ecology , Inst itu to de Ecologia A. C.,
Km 2.5 Carret. Ant. a Co atepec No . 35 1, Xalapa 9 1070, Veracruz (Mexi co)

Luis Alfredo PEREZ-JIMENEZ

Inst itut e of Biol ogy , Universidad Nac io nal Aut6no ma de Mexico ,
Apartado Post al 70-233 (Mexico)

Arceo G , Mandu jano S.. Gallina S. & Perez-J imen ez L. A. 2005. - Diet divers ity o f white -
tailed dee r (Odocoi/eus virginianus) in a tropic al dry forest in Mexic o. Mamm :.l/ia 69 (2) :
159-168 .

ABSTRACT
W/e report on div ersity in rhe d iet of wh i tr tailed deer (OducoiLcu, virginian li S)
in a Mexican tropical dr y forest . Diet was estima ted using the m icrohisio logi-
cal ana lysis of plant epid ermis in deer pellet-groups. T he sam pling was from
J993 ro 1994. Fro m rhc rai ny [Q dry seaso n, rhc richn ess and diversiry o f rhe
[ un ilies a nd specks in the diet increased and bot ani cal co m posi tio n changed.
D eer selected 8 2 pla nr species from 20 famili es; however , 12 species rep re-
KEYWO RDS scn rcd 50% o f the an n ual di et. f lowers and fruit s of arboreal species were
diet divcrsitv, impo rranr duri ng the dry seaso n, as wer e yo u ng leaves o f sh ru b a nd vine
seasonal var iatio n , species durin g th e rainy season. De sp ite the sma ll size o f white-railed deer in
Odocoileus vi rgi nid1J/fS,
tr o pical Io rcsr, rhis tropi cal dr y for esr study a rea, for aging strategies th ere were sim ilar ro
Mexico . chose observed in rem perare fo rests.

REsUME
Di uersirc de l'alimentation du w f de Virginie (OJ oco ileus virg in ian us) dan'
une JO rh tropical» au Mrxique.
Cc r a r ric lc prc sc nt c [a d ivcrs it c d e l' a li rnc n rnr io n du ccrf de V iq:.~ i n ie
(Odocoilem virgi l/ianus) dans une foret rropi calc d e I' f:tar de M exiquc . Le
regim e alime nraire a ere csrimc par l'analysc rnicro-hisiologiquc de l'cpid crrnc
des planres da ns lcs f'eces de certec ha nrillo u necs penda nt la saiso n des pluies,
la sa iso n de transit ion er la saiso n seche en 1993-1 ()')4 . I.a richessc cr la div er-
sire des fami lies er especcs dan s le regime alirnen raire a augrnenu' de la saiso n
seche a la saiso n des plui es, avec un changcmcnr de la com position bo tani qu c,

tv1AM MALIA • 2005 • 69 (2) © Publica tions Sc ientifiques d u M useum na tional d' Histoire nature lle. Paris.
159
I Arw , C . ,·r,,1

Les ce r Fs o nr cbois i 82 es pcccs ap p;lrr e na n, a 20 Famil ies, J 2 cs pece:

seulernenr consriruanr 50 % d u regime a lirnc n ra irc. Flcurs, fru its cr Fe ll i ll e~
MOTS CLEs d 'arbres craicnr irnpo rran rs pe ndant la saiso n sechc, jeun es fC lIilles de buisson-
diversire alirncnraire, er especes ramp anr es pendant [a saison d es pluics. Ma lgrc la raille corporcllc
variatio n saiso nn iere, plu s perire du cerF d e: Virgin ic da ns la Fo rer rropicale scchc de l'crud e, les
Odocoi lcus " jrgit/ illl/lIs,
fo ret tro picale. strategies alirnenr aires apparaisscnr asscz semblables a celles des pop ulation -
Mexique. des forers tcmp erces,

IN TRODU CTIO N sa rc for rh is seaso na l limirarion by selec t ing

Bod y size is an important factor in rh e dererrn ina-
tion of ru mi nant ecology (Dernrnenr & Van Soesr
1985; H anley 199 7) . The white-t ailed deer
(Odocoilcus virgin ianus) has a wide geog raphica l
distriburion from Sou thern Ca nada ro N orrh ern
Brazil an d Central Bolivia (H all 1981 ; Eisenberg
1989); in consequence, size and bod y weighr vary
nota bly (Sm ith 1991) . In co n trast ro rem perate
habir ars in the nort hern region, where male and
fem ale mean weight (\X') is 100 kg and GO kg,
respectively, in tropical ha bitars weigh r can be 50 %
lower (Geist 1998 ). As me tabo lite requi remen ts
scale co \ '(10.7 5 ;J oJ ru men volume ro W U ) (Shorr
eta!' 1969 ), rhen smaller deer should require higher
qual ity d iet rhan those inhabiring rem perare area
(H anley 1997). Amo ng rumin ant s, the whi te-
railed deer is co nside red an opportuni stic concen-
rrare feede r (M ysrerud 1998 ), selecring a grear
d iversi ry o ffood and choosi ng relatively n utririous,
highly d igesrib le plan ts and plane parrs such as
young leaves, twigs, and saplings (Van gilder et £11.
1982; H enk e eta!' 1988).
I n tr op ica l habit at s, w hi te- ta iled de er inhabit
q ui te var ied vege rario n ryp es suc h as t rop ical
sava n na h, man g ro ves, flood for est, mounrain
habir ars ar upper elevations, tropical d ry for est,
seco nda ry vegerarion bord ering rai nforest , o ld
seco nda ry for est , ope n woodland s, an d field s
(Brokx 1984; M end ez 1984) . U nlike tem perare
h ab i ra t s w he re rh is d eer sp ec ies fac es seve re
scarcity offoo d in win ter, in several tropi cal habi-
rats the cr itical pe riod is th e dr y sea son , whe n
plant abundanc e and quality decrease. In conse-
qu ence, it has been assumed tha t dee r compe n-
species and plant co mmu niries rhar offer the best
qu ality o f food . However , few st ud ies of w hi te-
railed deer dicr have been und ert aken in tropical
for est s (e.g., Branan et a!' 1985; Gran ado 1989;
D iMar e 1994 ) . Because this sp ecies is an i mpor-
ra n r f o od reso urc e for loc a l pe o ple (e. g. ,
Mandu jano & Rico-Gr ay 199 I ), ir is irnpo rran r
co de fine man agem enr po licies rhar mainrai n sus-
rai nab le po p ula tio ns in t h e tr opi cs (We ber &
Go nzalez 200 3) . Th erefore, rhe study of foraging
strategies is key co und ersrandin g deer su rvival in
trop ical habit ats.
T he white-railed deer inh abi ts seasonally tropi cal
dr y fo rests o n rhe Pacific C oast o f th e stare o f
j ali sco , Mexico, wirh po pu larion densities rang-
ing from 10 ro 14 deer per km 2 (Ma nd ujano &
Ga llina 199 5). T his fores t is cha racterized by a
marke d variation in plane phenology, due prin ci-
pally ro seasonality in rain fall parrern s that divide
rhe year in to a rainy and dry seaso n, each lastin g
6 ro 7 month s (Lorr eta!' 1987 ; Bullo ck & 501is-
M agallan es 1990) . In parti cular, plane richn ess,
b iom ass, and qu ality vary not ably in the und er-
sto ry rhr oughout rhe year (Silva-Villalobos et £11.
1999) . To berrer u nderstand rhe pop ulation ecol-
ogy of rhe deer (Ma ndujan o et al. 20 02 ), we con-
du cted rhe p resen r study , focusing on how th is
small subspecies (40 kg of body weighr) faces sea-
so na l a nd spatial va riatio ns in foo d resources.
Th e objective of the presenr paper is ro add new
infor matio n abo ut irs diet in tro pical forest and
to discuss the relat ionship between feed i ng srrate-
gies and rhe bod y size of white-railed deer co m-
parin g w it h o rhe r st ud ies in north a nd so ut h
lat itud e.

MAMr:' AI IA • 20U,; • 69 (?\

160

F,G. 1. - Loca lization o f the "C hamela" Biolog ical Stat ion in the
Pacific coast of the Mexican state o f Jalisco . Tile shady rect an-
gle o f the inferior map rep resents the study place.
STUDY AREA
Th e s t u dy was co n d uc ted a t t he C ha me la
Biolo gical Sta tio n, o pe rated by the U niversida d
Na cional Auton oma de Mexico . T he stat io n is
3 ,600 ha and located on ]alisco so uthwest coas t
(Fig. 1). Fieldw ork was restricted to the 500 ha of
th e sta tio n rese rved for resear ch . M ean a n nua l
temperature was 25°C (Bu llock 1986) . Hottest
rnonrh s arc [rom M ay to September. From 1977
CO J 99 7 , the me an a n n ual p recipitati on was
748 mm (SD = 119), with 80% of the rain falling
betwee n Ju ly and O ctob er. T he do m inant vege-
tat ion is trop ical dry forest locat ed on hilly ter rain
w ith th in so ils a nd low wa ter retent ion. M an y
tree and sh rub species lose their leaves d uring the
dry seaso n. Co mmon trees are Cordia alli odora,
Lonc hocarp us [a nceo latus, a n d Ca esalp in ia
eriostachys ( Lo rr et al. 1987) . T he station also has
tropi cal se mi-evergree n for est , covering 2'1% o f
MAMMALIA • 2005 • 69 (2)
the area mainly along rhe major streams in deep
soil with high water retent ion. In th is habitat less
tree species lose their leaves dur ing the dry season
com pa re to th e dr y fore st habitat. Th e most com-
mon rrees in thi s typ e of for est a rc Astronium
graveofens, Brosimum alicastrum, and Sideroxylon
caplri.
METHODS
MI CRO H ISTOL O GI C AL ANAl.YS IS
Fr esh pell et gro u ps wer e obtain ed durin g the
rainy (Aug ust and O ctober of 1993) , tr ansiti o n
(N ove m be r and December of 199 3), and dr y sea-
so ns (A pril a nd M ay of 1994 ). The y were col-
lected along fou r trop ical d ry Ior est transects and
two in tropi cal semi-evergreen forest. Each tran-
sec t was 4 00 In lo ng ; ar eve ry J 0 rn, a circ ular
pl ot mea su rin g 10m! was m ar ked oFf (M a n-
duj an o & G allin a 199 '5 ). A to tal of 243 faecal
groups were co llected . Anthon y & Sm ith (1974)
indi cate a sam ple size of 15 to 50 pellet groups as
th e sugges ted minim um required to deter mi ne
dier o n a seaso nal basis.
T he use of c hlo ral hy d ra te to di scol o r a nd se t
sam p les is, acco rdi ng to M idd leton & Sanc hez-
Rojas (1994 ), the best rechn iqu e to de re rmi ne
herbiv o re di et. C h lo ral hyd rat e is, how ever , a
resrricred d rug in M exico. In the present study,
th is substa nce was th erefore replaced , followin g a
s lig h tl y m odified ve rs io n of r h c r ec bn i qu c
de scribed by Quin tanilla et al. (19 88). I t co n-
sisred of placing the sam ple in sod ium hyd roxide
ro '5%, boiling it fo r 5 minutes, rinsin g it wirh
distilled water, and add ing sodium hypochlor ite;
ir was th en lefr to sta nd unr il co m plete d iscol-
oration had taken place. Th e sam ple was rinsed
wi th dist illed water and pl aced in alco ho l so lu-
tion s, each with d ifferent co nce ntra tions, an d lefr
to stand for 20 m inu tes u n til co mplete dehydr a-
tio n had occ urre d. T he samp le was rhen mixe d
wieh Xy lol in o rd e r co st a in t h e r i ss u es .
Su bsequ e ntl y, a da b of C anad ian balsa m mixed
with a porrion of rhe treated rnare rial was placed
on ea ch slide. For eac h pla n t species , va rio us
sl ides wer e pr epared in an effort to ga rher th e
161 I
I Arceo G . <I al.

grearest possible microsco pic hisrological ch ar ac-
rer isric s. The sa me procedure was app lied for the
elabo rat io n of th e reference co llec rio n in order ro
become fami liar wirh rhe h istological cha racre ris-
tics of different plant species.
To facilitare rhe co m pa riso n o f reference slides as
a me an s of id enti fyin g th e species pre sent in the
diet. we use d a photographic collecrion of th e
vario us str uc tu res. In order to id enrify and quan-
ti fy t he pl anr s pec ies present in rhe diet, fiv e
m ixed p repa rat ion s of pellet groups collecr ed
du rin g on e sea son were prepared following th e
Spa rks & Malecheck (1968) rechnique . In each
pr eparati on a grid co ns ist in g of 20 microscopi c
field s was used to co u nt pl ant fragments. Thus.
each field uf observation represented a sa m p le
un it, wirh a total of 100 fields per seaso n. To esti -
mate th e a mo u nt of sam ple un its rhar might pro-
vi de acc e p t a ble rcpresenrat iou of the seas o na l
d ier, a regis ter was kept of th e new plant species
appe arin g in e ac h a ccumul at ing m icro scopi c
field . Th e mi croscopi c sam p ling W <lS carried out
b y reg isrerin g rhe frequenc y of identifiabl e plant
fr agm ent s p er field (Pe fi a & Hab ib 19 80) .
S u bseque n rly, spec ies frequ enc y w as di vid ed
b etw een th e total frequen ci es o f a ll s pec ies and
mult ipl ied by 100 in order ro o b ta in rhe percent-
age of eac h planr spec ies in th e di et.
D IET D IVERS IW
To a na lyze d ee r fora g in g st ra teg ies in further
d er ail , two di versit y indi ces we re est im at ed: one
;H spec ies lev el (pe rce n tage of e p ider m al fr ag-
m enrs per spec ies) a nd anorh er ar fa m ily lev el
( n u m be r o f s pec ies per fa m ily) . Di versir y wa s
esrirna red usin g rhe Sha n no n-Wie ne r index (H' )
a n d c o m p a r e d bet ween seaso ns u sing the
S t ude nt 's z-r esr (Z a r 198 4). Furth ermore, we
ren cy of leav es (d ec id uo us and eve rg ree n). Later,
we cl assi fie d as co nc en t ra te forage type herbs,
yo u ng le aves o f tr ees , fruit s of tr ee and sh ru b
spec ies wirh d eciduous young leav es: and fibrou s
forag e ryp e was co ns id e re d to be g ras,es, a nd
leaves/bran ch es of tr ees a nd sh ru bs with per en -
nial leav es. Becau se we did not CO ITecr for diffe r-
ential digesribiliry of each co nsu me d spec ies (see
(, ill et al. 1983 ), a potenti al bi as ex ists in ho w
mu ch of a particul ar spec ies wa s ea te n bu r nor in
se as o na l di er d iver sir y. H o w e ver, a n easi ly
di gesribl e concentrat e food like fruir co uld disap -
pear in fec al a nd ye r be und er esrim ar ed in t he
seaso na l d ier.
RE SULTS
T wenty bo tanica l fam ilies co nstit u ted t he a n nua l
di et (T able I ) . Th e famili es Euph orb iac ea e ,
Legu minosae, Co nvo lvu lacea c, a nd Sap indaceae
accounred fo r 80% , 64% , and 37% durin g th e
rai ny , r ran siri o n and d ry se as o n resp ecri vel y .
T hese fam ilies we re repr ese nt ed by three to re n
s p ec ies. Oth er imp orranr fam ilies in d iffe re n t
seaso n s w e re Ma lva cea e , S rer c u liaceae, a nd
A naca rd iaceae. In particular, during th e dry sea-
so n 63% of rh e fami lies pr esent wer e rep resented
by o nly one spec ies. Fam ily r ich ne ss and d ive rsity
in cr eased s ig nifica ntly fro m th e rain y ro dry sea-
so n (Fig. 2; r = 2.10, d .f. = 68 , P = 0.0 3), D iet
similarity was 22% berween rain y and tran sition ,
g
III
~ 40
c
70
60 -
50
1

.-- /-
.' •
4

3.5 ~

3 ~
e
compared s im ila riry in borh family and s pec ies ii 0 2,5 .~

.-
30

•
'c 20 "0
co m po s irio n be tw een seaso ns using rhc Bray- 0
2
C u rti s d issim ila rity ind ex. 10
i
0 1.5
T o de term in e w het he r deer co ns umed more con- Rainy Transitio n Dry
ce nrrare o r fibrou s fo rage types, we first sepa ra ted
..... R famil y __ R spec ies --e--- H' family - - H' species
rhe p lan r fragments regisrer ed in microhisrologi-
ca l a na lys is into th e followi ng ca te go ries: plant FIG. 2, - Estima tion of the number of fam ilies and species (R,
pa n s (lea ves, br an ch es, flowe rs, and fruits) , life ric hnes s) and d iversit y (H', Shannon index) of the wh ite - tailed
deer d iet dur ing t he rainy, tran siti on and dry sea sons in a
fo rm (trees, sh rubs, herbs, and g rass), and persis- Me xican tropi cal forest.

MAMMALIA • 2005 • 69 (2)

162

1
 D ie( d iversity o f whi te-railed deer (OdoroilCII' ";'1';11;"'11(1) in a tro pical d rv fo resr in M exico I

TABLE 1. - Contribution of plant species (leaves or fruits, expressed as % count in fece s) to the d iet of th e wh ite-t ailed deer in the
trop ical d ry forest of Chamela. Mexico.

Season
Family Species
Rainy Transition Dry

Amaranthaceae Iresine pac ifica 0 .3

2 spp. 0 .6
2 spp. 8.5
Anacardiaceae Spondias purpurea (fruit) 13 .1
Bignoniaceae 1 sp. 2.6
Bombacaceae Ceiba sp . 03
Compositae Zinnia maritime 0 .3
Connaraceae Rourea glabra 0.6 0 .3 4.8
Convol vulaceae Ipomoea ampul/a cea 2.0
Ipomoea quamoc/it 0. 3
Ipomoea trttiae 0 .3 2.2
Ipomoea sp . 0.6 0.8 0 .3
Ipomoea sp . 0.9 0.5 2.0
1 sp . 13 .8 11 .3 3.7
1 sp . 1.5 2.6
Euphorbiaceae Acalypha langiana 23 .4 14.8 1.6
Acalypha shiedeana 1 .6 0.6
Acalypha sp . 1.2 2.8 0.6
Acalypha sp. 2.2
Acalypha sp. 0.5 0. 3
Acalypha sp . 0.5
Acalypha sp. 0. 3
Croton sp. 4.4 6.5 0.6
2 spp . 0.8
Graminae 1 sp . 0.5 1.0
Leguminosae Apoplanesia panicula/a 0.5 0.6
Brongn iartia sp . nov. 0.6
Caesalpinia coriaria 0.3
Coursetia caribaea 5 .8
Cro/alaria sp . 0.5
Desmodium pro cumbens 2 .5
Haema/oxylum brasilelto 0.5
Tephrosia leiocarp a 3 .8 0.8
Zapoteca formo sa sub. Rosei 1.2 0.3
1 sp . 1.8 0.8 2.6
1 sp . 69 5 .5 2.0
1 sp . 0 3 0 .3
Malvaceae AbutiJon mcvaughii 0 .3 0 .3
Abutilon sp . 0 .3 6.2 1.3
Abu/ilon sp. 2.1 1.9 1.0
Briquetia spicata 0.5
Malvastrum coromande lianum 0.3 1.4
Sida glabra 2.2
1 sp , 0.6 0 .3
3 spp . 2.1
Moraceae Brosimum alicastrum (fruit) 1.0
Nyctaginaceae Pisonia aculea/e 0 .3
Olacaceae Ximena pubescens 0 .3 0.6
Sapindac eae Cardio spermum halicacabum 6.6 4. 0 12 .1
Thouinia pauciden/a/a 1.5 2.6 3 .3
Serjania brachy carpa 1.5 0.8 1.0
l sp. 1.5 2.6 0 .3
Sapotaceae Sidero xylon sp . 0 .3
Slerculiaceae Ayenia micranta 1.2 4.9 2.6
Theophrastaceae Jacquinia pungens (fruit) 1.6

MAMM ALIA • 2005 • 69 (2)

I A rceo , ; . rt "I.
38% between rainy and dry, and 36% b etween
rransirion and dry seasons .
The annual diet was compos ed of 82 sp ecies
Crable 1). Dier dive rsity increased significantly
from rlie rainy ro dry seaso n (Fig. 2, t ~ 4 .06,
d .f. = 60, p ~ 0 .00 I) . Dier similarity was 22%
b etween rhe rainy and rransitio n seasons , 16%
between the rainy and dry seasons , and 26%
between rhe transition and dry seasons . In partie-
ular, of rhe 4\ species consumed during the rainy
sea son, .'3 6 species w ere rare « 5% co m b ine d ) ,
while only five (Acalypha langia na, Card ia-
spcrmum halicarabum, Cou rsetia caribaea, and
two underermined Co nvo lvul acca e and
Legurninosae) constituted 5 7 % of th e diet during
rhis season. Of rhe 65 spec ies consumed during
rh e tr ansition season, 59 were represenred by less
than 5% of planr epidermis in th e sample, wh.ile
only six species (Acalypha lang iana, Croton sp.,
Abutilon s p., ,iyenia rnicranta and two underer-
11.64
min ed Convolvulaceac and Leguminosae) consri-
r uted 49% of rhe d ier in rhis season . Of th e
61 species c o n s u m e d durin g rhe dry se aso n ,
57 made up < 5% of the planr e p id e rm is in rhe
sample, while only four s p ec ies (Spondias p ur-
pu rea, Rourca tlabra , Cardiospe rm um haLica-
cabum , and one und ererrnincd Am ar.mth aceae)
constituted 38% of the diet in rhis seaso n .
Life forms were signiflcanrly differenr among <ca-
so n s (Table 2 , X 2 = 49.9, d.f. = 9, p = 0.001) .
Shrub (principally Euphorbiacea e) and vine
(Ccnvol vulaceae) specie s were the rno sr impor-
tant throughout rhe year, bur in the dry seaso n
tree s (Anacardiaceae) were al so i m p o rt a nr .
Deciduou s leaf spec ies had the highesr percem
values in rhe diet throughout rhe year, alrhough
r h c consumprion of e ve r g ree n leaf spe c ie s
increased during th e dry seaso n (X 2 = 4.1, dJ. = 2,
P . 0 .13) . Conccnrrare forage type mad e up th e
highesr percenrage of rhe diet during the year ;
MAM MALIA • 2005 • 69 (2)
T ABLE 2. - Seasonally variation of the catego ries of plant s
former. The nutritional quality hypoth esis pr e-
consumed (expressed as % count in feces) by the white-tailed
deer in the tropical forest of Chamela, Mexico. di cts th at anim al s will b e less se le ct ive durin g
per iod s of hi gh resource a b u n da n ce becau se a
Season w id e variety of high -quality food resou rces a re
av a ila b le . In co n tr ast , the for agin g abundance
Category/
Rainy Transition Dry hypothesis p redi ct s th at when th e abund ance of
Characteristics
food resources in cre ases, an im als ca n a ffo rd to be
Life form :
more selective becau se the ir in cr eased fora gi ng
trees 5 18 38
shrubs 50 39 29 effici ency allows th em to se lec ti vely con sum e
herbs 10 18 4 pl ants that ar c m ore nurri riou s. Plants in th e
vines 35 21 25 s tudy area ha ve higher co ncent ra t io ns carbohy-
grasses 0 4 4
dra res and protein a n d lower fib e r during th e
Lea ve persistency :
evergreen 6 8 14 rain y seaso n, w h ile during th e dry sea son th e
deciduous 94 92 86 nurriri on al qualit y and abundan ce d im in ishes
Parts: (Si lva- V illa lo bos et at. 1999) . Thi s later is al so
lea ve s 86 89 70
cha ra crer iz e d by rhe hi gh e st d i cr diver sit y ,
fruits 14 11 30
according to th e foraging abundan ce h ypothesis.
Th e white -tail ed d eer con sumed concentrate
foods suc h as yo u ng leav es o f s hru bs (principally
fruits were i m po r ta n t during th e dry se as o n Acalypha, E u p ho rb iaceac) a nd vines iLpomoea.
(T able 2, X 2 = 13.9 , d .f. = 2, P = 0.001) , gr asses Corivolvulaccac) dur ing the rain y seaso n . In par-
we re poorly eaten a ll year round. ticul a r, perenni al species wirh d eciduou s leaves
were t he m ost importa nt in th e diet. This co in-
cides w ith findings reponed in other st ud ies indi-
DIS CUSSIO N caring rh ar during rhe ra iny seaso n , yo u ng leaves
ge ne rally represenr a resou rce o f higher nutritious
Th e w hite- ta iled de er con sum ed less spec ies dur- quality and les s se c ond ar y compounds th an
in g the rainy seaso n and increased its d iet di ver- mature lea ves the qu ality o f w h ic h declines as
s i t y during th e dry seas o n . How e ver , only th ey age (Fo rd et ill. 1994). Fruits p ro ved to be a n
12 spec ies from six famili es wer e the most irnpor- important reso u rce in the d eer di et d u rin g th e
ranr in thi s tropical dry fore st. Freeland & Jan zen dry season . T h is prefer ence mu sr relate to rh e
(I (74) expl a in that the majority of herbivore peak of Flower and fruit produ ction obse rved
m amm al diets co ns ist prim ordiall y of rel atively durin g th e dry period in rhe stu d y ar ea as well as
few plant s pe c ies . While a hi ghly di verse diet ro a de c rease in hi gh-qu aliry leav es (Bulloc k &
allows them to o bta in nec essal) ' nutrients and to So lis-Magallanes 1990). In tr opical for est s, rumi-
redu ce the po ssibl e e ffec ts o f seco n d a ry com - nants select a high diversity o f spec ies, and fruits
pounds in plants. Empiri cal data and th eoretical ar e an important d ietary elem enr (e.g., Dubose
fora ging models p redicted rhar, as th e w inter sea- 1984; Bodmer 1989). In ge nera l, fru it is rich in
so n advances, an ungula t e may compen sate for non-structural ca rbo hydrates and fats and has rel-
decl ining food abundance by widening it s ali - atively low level s of crud e fib er , representing all
m ent;:lIY br eadth (c.g. , Shorr et al. 1974; O we n- i m po rra n t so u rc e o f ene rgy (V angild er et al .
Smirh & Novelli e 1982; IlIius & Go rd o n ] 993). 1982) . However, fruit co ns u m p tio n is limited by
In particul ar , W eckerly & Kennedy (1992) com- seas o na l pr odu ction, terrestri al availa biliry, a nd
pared the nutritional quality and for aging ab u n- inter-individu al tr ee production variation. Thus .
d an ce h ypotheses as rhe y apply to w h ite - ra iled it represents a patch y resource for concentrate
de er feeding strategies in a remperate fore st habi- se lec to r ruminanrs. Granado (1989) reporrs that
tar , and found rhar foraging parrerns support rhc fruit constirurcd 49% of th e d eer diet during the

M AMM ALIA • 2005 • fi9 (2)

165[
I Arceo G . (I ,Ii.

TABLL 3. - Comparison of white-tailed deer foraging strategy in diHerent habitats from South to North America.

Latitude gradient
Characteristics 6' N 9' N 19' N 23' N 28 N 40' N

Habitat type Tropical forests, Tropical Tropical Temperate Mesquite and Temperate
plains, and dry forest dry forest deciduous and chaparral-mixed coniferous and
cultivated areas mixed forest grass deciduous forest

Families:
total 33 23 20 41
principals 11 6 5
Species:
total 125 45 82 135 39 23
principals 16 12 7 4 4
Diversity' :
Rainy season 1.1 low 3.0 1.9 0.9 0.75
Dry season 0.7 high 3.5 2.5 1.1 0.70
Principal food items:
Rainy season? herbs herbs leaves herbs herbs fruits
grasses grasses vines leaves leaves
herbs
Dryseason leaves fruils leaves leaves leaves herbs
fruils leaves vines grasses
grasses
fruits
Method of diet analysis fecal fecal fecal fecal rumen rumen
Country Venezuela Costa Rica Mexico Mexico USA Canada
Reference Granado 1989 DiMare 1994 This study Gallina et al. Kieel al. 1980 Skinner & Teller
1981 1974

- no specific data: 1 when available we estimated the Shannon diversity index using data of Ihe author; ! rainy season includesspring,
summer and fall in temperate region, and dry season includes winter.

dr y seaso n i11 a tro pical foresr in Ven ezue la. In
o the r hab itat cypes, frui t and flower are imp or-
ran r food fo r deer (Ford et al. 19 9 3 ; Di Ma re
1994 ; J ohn son ct al. 19 9 5). In th e st udy area ,
rhey satisfy the need for food and water by con -
su rn ing pri ck ly pea rs i Opuntia excelsai, ramon
(Brosimum alicastrum i, ficus iFicus spp.) and red
morn bim (Spondias purpurea) fruits d ur ing t his
seaso n (M a nd uja no et al. 1994). In pa rt icular ,
rhe fru it of S, purpurea. a species also reponed by
DiMare ( 199IJ) in a Co sta Rica n trop ical dry for-
esc. Plan r pam th at are easily d igest ed and ch ar
have low cellu lar walls are usually und erestim ated
in faeces co m pa red [Q their propo rtions in t he
real diet (Vavra & Holecheck 1980). It is there-
fore probab le char t he cont rib ution of flower s
and [ruirs (Q rhc d iet was higher (ha n rhe percent-
age reported herein.
A non exhaus tive com parison on data of forag-
ing of the wh ite-railed deer in a latitud ina l gradi -
e n t , showe d th a t r h is s p ec ies h as a si m i lar
srt'ategy in d ifferenr types o f tropical, temperare
an d sem i-a rid hab itats (Ta ble 3 ). Based on rhe
pe rce nt age o f co n rr ibu tio n o f each species in
rh e d ie t, it wa s ob serve d th at o f rhe [O ra l o f
consumed plants, 12-33 % o f the fam ilies and
5- 17% of rhe spec ies were the most irnporra nt .
Also , in 67 % of the analyzed pa pers the dive rsity
of th e d ier was sma ller during t he rainy season.
Although the pe rce n t:lge of fooel ite ms va ried
sea sona lly am o ng hab ira rs . in ge n e ral i t was
ob se rved that t he herbs and gr:.tsses were mo re

MAMM ALIA • 2005 • 69 (2)

166

1
Imporra nr in the rainy Seaso n, while the fr uits
an d th e leaves of trees an d bu sh es were irn po r-
rant in the d ry seaso n. U nfor runa rely, sp ecific
dat a o n the co rporal weigh t of deer are no t pro-
vid ed in each ana lyzed st udy, thus was no t possi-
b le to m ak e a d et ailed analysis in th is top ic. In
concl us io n, the over all re sults s ugges t th at
despite t he relat ively sma ll size o f w h ite- ta iled
deer in th e stud ied trop ical dry fo rest, thi s deer
be haves as oppo rtunis tic co nce ntrate se lector
whic h is simi lar to the forage stra tegy in temp er-
ate hab ita ts.
Aclmowledgements
The C o ns ej o Nacio na l C ie nc ia y T ec nol o gi a
(C O N AC YT) p rovided study grant to G . A rceo.
T h e proj ects CONACYT (P 2 2 0 C C O R -
8 92 154 ) , CO NA C YT (P020CCOR-903 703) ,
SE P (D G TCSA -902 46 7 ) , and CO NACYT
(0 3 27 N 9 10 7 ) p rovided financial support. T he
C ha rnela Bio logica l Sta tion, Insriru ro de Biologia
o f the U niversidad Nacional Auto no rna de M ex-
ico, provid ed logistic sup po n an d facilities. W e
also thank to G . Silva-Villalo bos, R. E. Sanche z-
Manti lla, C. M. Ga lan , C. Pacheco and S. Leon-
Torr e s for t h eir h elp durin g t he fie ld an d
labo rato ry work. T he fol lowing peo ple co llabo-
rated in th e identification o f herba rium samp les:
t= . C h ia ng , M. Sousa, O . Tel lez and ]. L. Vi l-
lasenor. Also, we th ank to C . Aguilera-Reyes an d
G . G omez - Belt ran their help d uring the micro-
hyscological analysis. Early versio n of this man u-
sc r i p t w as read by A . Go n za lez- R o m e ro ,
V . Sanch ez-Cord ero , Y. Hoerrelano, L. Leon,
an d J. M urcia . Lastly, two anony mo us reviewers
prov ided sugg es t ions rh a t im p roved th e fi nal
versio n of this paper.
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