NUTRITIONAL CHARACTERISTICS OF PLANTS CONSUMED BY THE

WHITE-TAILED DEER IN A TROPICAL FOREST OF MEXICO

GABRIELA S/LVA-VILLALOBOS, C.U.C.BA, Universidad de Guadalajara, Apartado Postal 39-82, Zapopán 45110,
Jalisco, México. (Tel y Fax: 3-682-0230)

SALVADOR MANDUJANO, GLORIA ARCEO, SONIA GALLINA, Departamento de Ecologia y Comportamiento

Animal, Instituto de Ecología A. C., Apartado Postal 63, Xalapa 91000, Veracruz, México. (Te/: 28-421800, Ext.
4110; Fax: 28-187809; ENVIAR CORRESPONDENCIA A Correo-E: mandujan@ecologia.edu.mx)

LUIS ALFREDO PÉREZ-JIMÉNEZ, Estación de Biología Chamela, Universidad Nacional Autónoma de México.
Apartado Postal 70-233, México, D. F. 04510, México. (Tel: 5-622-5692; Fax:5-550-1760; Correo-E:
aperez@ibiologia.unam.mx)

ABSTRACT
The objective 01' this study was to determine the nutritional value 01'
sorne plant species consumed by white-tailed deer (Odocoi!eus
virginianus), as well as other species abundant in the understory 01'
tropical deciduous forest (TDF) and tropical semi-evergreen forest
(TSF). but not consumed by deer. The study was carried out at the
Charnela Biological Station on the coast 01' Jalisco, México. From
October, 1994 to August, 1995, bimonthly determinations were made 01'
the percentages 01' crude protein, nitrogen-free extract (NFE). crude fiber
and total polyphenols present in 11 species 01' plants consumed by the
deer, and 21 unconsumed species (10 from TDF and 11 from TSF). We
found that plants consumed by deer had more NFE, protein and
polyphenols and less fiber than unconsumed plants. TDF plants had a
higher nutritional value during the rainy season than during the dry
season. TSF plants were higher in fiber and lower in protein and NFE
throughout the year. Fiber was correlated negatively with NFE and
protein.
KEY WORDS: México, nutritional value, Odocoileus virgimanus,
seasonal variatlOn, tropical deciduous forest, tropical
semi-evergreen forest, white-tailed deer.
RESUMEN
El objetivo del estudio fue determinar el valor nutriclOnal de algunas
especies que consume el venado cola blanca (Odocoileus vlrgInzanus)
y de especies no consumidas pero que son abundantes en el soto bosque
del bosque tropIcal caducifolio (BTC) y del bosque tropical subperenni-
folio (BTS) de la Estación de Biología "Chamela" en la costa del estado
de Jalisco, México. Se determinó bimensualmente, de octubre de 1994
a agosto de 1995, los porcentajes de proteína cruda, estracto libre de
nitrógeno (ELN), fibra cruda, y polifenoles totales de 11 especies
consumidas por el venado y 21 especies no consumidas (10 del BTC y
11 de BTS) Encontramos que las plantas que consume el venado
tuvieron más proteína, ELN y polifenoles, y menor fibra que las plantas
no consumidas. Las plantas del BTC tuvieron un valor nutricional alto
durante la época de lluvias y menor durante la época seca. Las plantas
del BTS tuvieron mayor porcentaje de fibra y menor de proteína y ELN
durante todo el año. Se encontró correlación negativa signíficativa entre
el porcentaje de fibra con el porcentaje de proteina y ELN de las plantas.
PALABRAS CLAVE: bosque tropical caducifolio, bosque tropical
subperennifolio. México, Odocoi/eus virginia-
nus. valor nutricional, variación estacional.
venado cola blanca.
RESUMO
O objetivo deste estudo foi determinar o valor nutritivo de algumas
espécies de plantas consumidas pelo veado (Odocoileus virginianus).
assim como de outras espécies abundantes da floresta tropical caducifó-
lia (FTC) e floresta tropical subperenifólia (FTS), mas nao consumidas
pelos veados. O estudo foi realizado na Esta¡;;ao Biológica Chamela. na
costa do estado de Jalisco, México. Determinou-se bimensalmente, de
outubro de 1994 a agosto de 1995, as porcentagens de proteína crua,
extrato Iivre de nitrogénio (ELN), fibra crua, e polifenois totais de 11
espécies consumidas pelo veado e 21 espécies nao consumidas (10 de
FTC e 11 de FTS). Encontramos que as plantas consumidas pelos
veados continham mais proteínas. ELN e polifenois, e menor fibra que
as plantas nao consumidas. As plantas de FTC tinham um valor
nutritivo alto durante a época de chuvas e menor durante a época seca.
As plantas de FTS tinham maior porcentagem de fibra e menor de
proteína e ELÑ durante todo o ano. Encontrou-se correla¡;;ao negatIva
e significativa entre a porcentagem de fibra e a porcentagem de proteína
e ELN das plantas.
PALAVRAS-CHAVE floresta tropical caducifólia, tloresta tropical
subperenifólia, México, Odocoileus virgInia-
nus. valor nutritivo, varia¡;;ao estacional, veados
VIDA SILVESTRE NEOTROPICAL 8(1-2):38-42

A herbivore's diet is limited, among other factors, by
nutrient availability, toxin content, ingestion and digestion
rates and by the time invested in searching, consuming and
ruminating food (Hanley 1997). The white-tailed deer
(Odocoileus virginianus) is c1assified among those ruminants
that select very nutritious plants which are high in cellular
content, ferment rapidly, and are easily digested (Van Soest
1982). Since its rumen is small compared to body size, deer
must compensate for low rumino-reticular capacity by
choosing such high-quality foods (Henke el al. 1988, Short
el al. 1974). Nutritional value depends also on such factors
as 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 vegeta-
tion available to deer would allow a better understanding of
this herbivore's foraging strategies.

38

!
1
 The white-tailed deer is a major herbivorous species in Table 1. Species consumed and unconsumed during the rainy
tropical forests in the Pacific coast region of the state of and dry seasons inclued in the nutritional analysis (TDF-
tropical deciduous torest and TSF-tropical semiever-
Jalisco, Mexico, with population densities ranging from 10 green torest).
to 14 deer/km 2 (Mandujano and Gallina 1995a). In this
region, the precipitation pattern, as well as topographic and
Unconsumed Season
soil conditions (Bullock 1986) cause tropical deciduous
Consumed TDF TSF Ramy Dry
forest to predominate in hilly areas, while tropical semi-
evergreen {orest dominates riparian sites. These vegetation Dicliptera resupmala (Vahl) Juss. X X X
communities are different from each other in tloristic Caesalpinia eriostachys Benth. X X X
composition (Lott el al. 1987), phenology (Bullock and Acalypha sp. X X
Solis-Magallanes 1990), and biomass (Martínez-Yrizar el al. Croron sp. X X X
Spondias purpurea L. X X
1992). As a consequence, deer are faced with spatial and Heliocarpus pa/lidus Rose X X
temporal variation in the availability of food, water, and Coursetia caribaea (Jacq.) Lavin
cover (Mandujano and Gallina 1995b). vaL caribaea X X
This work is part of a research project aimed at discover- Desmodil/m procl/mbens
(MJ1!) Hitchc X X
ing individual and population strategies employed by white- TephroslO sp X X X
tailed deer in response to habitat heterogeneity in the tropical Serjania brachycarpa A Gray X X X
forest of Chamela, Jalisco, Mexico. An important factor to [,,"O.)'lml"'l alicastrum Sw X X X
Lasiaos rusclfolia (HBK.)
consider in investigating these strategies is the variation in
Hitchc vaL ruscifolia X X X
the nutritional value of vegetation, and how such variation Jacql/ll1la pl/ngens A. Gray X X X
might affect the selection of plants species included in the Trie/ullO trifolia L. subsp
diet (Arceo 1999), as well as intluence the activity and palmen (C DC) Pennington X X X
habitat use patterns of deer (Sánchez-Rojas el al. 1997, Lonchocarpus lanceolatus Benth X X X
E~vtrana Imbricara (Vahl) Pers. X X
Mandujano 1999). The specific objectives of the present ApoplalleslO panicl/lta Presl X X
study were: a) to determine the nutritional values of plants Hvballtlllls mexicanus Ging X X
consumed, or not consumed, by deer in tropical deciduous O'oton sl/berosl/s HBK X X
and semi-evergreen forests, and b) account for any seasonal Decachaeta haenkeana DC X X
El/patoril/m solidagil1l{olil/m A. Gray. X X
variations or differences in these values between plant Thollll1ldl1lm decandrum
species. (Humb & Bonpl) Radlk X X x
Cvnolllelra oaxacana Brandegee X X X
Forchhalllmena pal/ida Liebm X X X
STUOY AREA ROl/rea glabra HBK X X X
Cupal1la denlata DC X X X
This study was conducted at the Chamela Biological C~vtosfollla binafllm (Thunb.) Sandw. X X X
Station, operated by the Universidad Nacional Autónoma de Adenoca~l'Illa inl/ndatum Mart. ex DC X X X
México, and located on the southwestern coast ofthe state of Cappar¡s verrl/cosa Jacq. X X X
Jalisco, Mexico. The station covers 3,600 ha and has COlllbretumfruticosl/m (Loetl) Stuntz X X
Podopterus cordi{oll1ls Rose & Stand! X X
elevations ranging from 30 to 500 m. The region is charac- Gl/aplta Illacrocarpa Miranda
terized by an irregular topography. Mean annual temperature
is 25°C, with May through September being the hottest
months. From 1977 to 1984, maximum and mínimum range from 10-25 m in height. The most common trees are
temperatures ranged from 29-32°C and 15-23°C, respec- Aslronium graveo/ens Jacq., Brosimum alicaslrum Sw., and
tively. Mean annual rainfall is 748 mm (SD= 119 mm), with Sideroxy/on capiri A. DC. Mangroves, palm groves, and
80% falling between July and October. Rains normally occur thorny secondary scrub are also found surrounding the
from July to November, while the dry season lasts from station. Field work was restricted to the 500 ha ofthe station
December to June (Bullock 1986). reserved for research, and of this area, over 75% was TDF.
The dominant vegetation at the station is tropical dry
forest (TDF), located on hilly terrain with thin soils. Forest
METHüOS
height varies between 4 and 15 m, and a well-developed
understory is usually present. Most tree species lose their Due to time and cost considerations, we decided to work
leaves during the dry season (Lott el al. 1987). Some ofthe with only 32 plant species, despite high species richness in
most common trees found are Cordia a/fiodora [Ruiz & the understory ofthe study area and high species diversity in
Pav.] Oken, Lonchocarpus /anceo/alus Benth., and Caesa/pi- deers' diet (Arceo 1999). We selected some of the plants
nia erioslachys Benth. The station also has tropical semi- most frequently consumed by deer (as determined by direct
evergreen forest (TSF), which is typically found along observation) as well as other unconsumed species that are,
streams and in protected places with deep soils. These forest nevertheless, very abundant and found almost exclusively in

VIDA SILVESTRE NEOTROPICAL 8(1-2):1999 DEER FORAGE QUALlTY IN TROPICAL FOREST • Silva-Villalabas et al. 39
 Table 2. Percentages of nutritional characteristics of sorne groups. To determine possible associations between nutri-
consurned and unconsurned plants by white-tailed deer
tional variables, we performed simple linear regression
during the rainy and dry season in the tropical forest of
Charnela, Jalisco, México. TDF (tropical deciduous analyses for all pairwise combinations of variables.
forest), TSF (tropical sernievergreen forest). Mean ±
(standar error).
RESULTS
Unconsumed Anual mean percentage of crude protein was 14.2% in
Nutritional Content Season Consumed TDF TSF plants consumed by deer, 14.5% for plants from the TDF and
14.0% for plants from the TSF. These differences were not
Crude protein rainy 17.0 (0.3) 16.4 (O 1) 152 (0.1)
dry 11.4 (15) 126 (14) 12.9 (0.7) significant (F=O.13, df=2, 16, P=0.88). However, there was
a significant difference in mean percentage of protein
Nitrogen-free-extract ramy 48.9(07) 47.3 (06) 419(0.3)
dry 492 (13) 45.3 (09) 44.3 (0.3) between seasons (Table 2, F=33.8, df=l, 16, P<O.OOI).
During the rainy season, all vegetation had a higher percent-
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) age of proteins than during the dry season. There was no
significant interaction between season and group of plants
Total poliphenols ramy 2.5 (O 1) 1.7 (02) 16 (0.1)
dry 3.0 (O 1) 15 (01) 2.1 (02) (F=2.2, df=2, 16, P=O. 16).
Anual mean percentage of NFE was 49.0% for plants
NutritlOnal mdex ramy 3.1 2.7 2.0
(protein+N FE )/fiber dry 2.2 1.9 2.0 consumed by deer, 46.3% for plants ofthe TDF and 43.1%
in species of the TSF. These differences were significant
(F=33.2, df=2, 16, P<O.OOI). Although there was no differ-
ence between rainy and dry seasons (Table 2, F=0.17, df=l,
the TDF or TSF understory (Table 1). 16, P=0.67), there was a significant interaction between
We sampled various plants of each species along deer season and the particular group of plants (F=4.23, df=2, 16,
trails and dirt roads ofthe biological station. Only leaves and P=0.04). Plants from the TSF had the lowest NFE and plants
tender branches of plants were collected. Two hundred consumed by deer had the highest NFE.
grams of each species were collected, and samples were Anual mean percentage of fiber was 24.2% for plants
oven-dried at 50°C for 2 to 5 days. To prevent heat from consumed by deer, 26.9% ofthe TDF and 29.3% ofthe TSF.
volatizing sorne of the secondary compounds, 50 g of each These differences were significant (F=5.9, df=2, 16, P=0.02).
sample were air-dried. Later, they were ground separately in Species consumed by deer had less fiber. Percentage offiber
a Wiley mili with a 20-mesh screen, tagged and stored in in vegetation was lower during the rainy season compared to
bags. the dry season (Table 2, F=12.3, df=l, 16, P<0.005). There
Sampling was carried out in October of 1994 (mid-rainy was no significant interaction between season and group of
season), December of 1994 (late rainy season), February of plants (F=3. 1, df=2, 16, P=0.09).
1995 (early dry season), April of 1995 (mid-dry season), June Anual mean percentage of polyphenols was 2.6% for
of 1995 (late dry season) and August of 1995 (early rainy plants consumed by deer, 1.6% in the TDF and 1.8% in the
season). Each plant species from each location, sampled TSF. These differences were significant (F=46.7, df=2, 16,
during each season, was homogenized and analyzed for their P<O.OO 1), and plants consumed by deer had the highest
nitrogen-free extract (N FE), crude protein, and crude fiber percentage of polyphenols. There was a slight difference in
contents, using A.O.A.C. (1975) methods. Total polyphenols the percentage of polyphenols between seasons (Table 2,
were determined by the method ofLesli-Harti and Johnstone F=5.1, df= 1, 16, P<0.05). There was a significant interaction
(1989). AII determinations were done in triplicate except for between seasons and plant groups (F=4.1, df=2, 16, P<0.05).
polyphenols, which were done in duplicate. Crude fiber content ofplants consumed and unconsumed
We used the arcsin function to transform data which were by deer was negatively correlated with crude protein content
originally expressed in percentages (Sokal and Rohlf 1969). (r=-0.76, F=20.0, df=I,16, P<O.OOI), as well as with NFE
The frequency distribution of transformed data was normal (r=-0.71, F=15.0, df=l, 16, P<0.005) (Fig. 1). In general,
and variances were homogeneous, therefore we used para- plants from TSF had higher percentages of fiber and lower
metric tests. For each one of the nutritional determinations percentages of NFE and protein. In contrast, plants con-
(crude protein, NFE, crude fiber and polyphenols), we tested sumed by deer had lower fiber and higher NFE and protein
for seasonal differences among the three groups of plants contents. The nutritional index [(protein + NFE)/ fiber] was
(unconsumed plant species from TDF and TSF, as well as higher for consumed species than for unconsumed species
plant species consumed by deer). We applied analyses of during the rainy and dry seasons (Table 2). The nutritional
variance (Sokal and Rohlf 1969) and for significant results index was lowest for the TSF plants.
we employed the Student-Newman-Keuls a posteriori
multiple comparison test to determine differences between

40 DEER FORAGE QUALlTY IN TROPICAL FOREST· Silva-Villalobos et al. VIDA SILVESTRE NEOTROPICAL 8(1-2):1999
 fiber content of plants increased. This can be explained by
34 ~---------------------,
noting that as plants matured, their cell wall constituients
• increased and their digestibility decreased (Short el al.
1974). The fiber content (mainly lignin) is known to be
greater in leaves of evergreen species than in leaves of
deciduous species (Prudhomme 1983). This suggests that,
because many species in the TSF keep their leaves year-

.-
round, they have developed mechanisms to reduce the
possibility ofattack by herbivores. Therefore, during the dry
22
season forage quality and abundance is low in both vegetatio-
20 -'--<---+I---+--+----+--<-------,---+-,
8 10 12 14 16
.------,,----1
18
nal communities, and deer must modif)r their foraging
crude protein ( % )
strategies (Arceo 1999).
The biochemical content of plant species consumed by
deer during the rainy season was higher in nutritional value,
34 showing a greater percentage of NFE and protein content,
32 • and less fiber, than in unconsumed species. This suggests
that in order to allow the accumulation of fat reserves for the
_30
;;s¡ mating season and for the critical dry se::son, deer must select
~28 plants which are more nutritious (Vangilder el al. 1982). The
'"
-O
estimates of crude protein contents (10-17%) could poten-
""'" 26
"'O tially cover the requirements (6-16%) for the growth and
2
U 24- development of fawns and adults, as well as for the growth
22-
ofmale antlers (Asleson el al. 1996, UlIrey el al. 1967). The
low fiber content and high percentage of NFE in the plant
20
40 42 44 46 48 50
species consumed throughout the year suggest that deer
nitrogen-free-extract ( % ) prefer species of high cellular content. Deer obtain a high
percentage of digestible energy from cellular content (Am-
• consumed • unconsumed TDF ... unconsumed ~SfJ mann el al. 1973, Thompson el al. 1973), especially during
the dry season when food availability is very low.
Due to the astringent and indigestible nature of tannins
Figure 1. Correlation between crude fiber with crude protein and (Robbins el al. 1987), we expected deer to select species that
nitrogen-free-extract for consumed plants by white- were low in polyphenols. At this site, however, deer selected
tailed deer and unconsumed plants from tropical plants with a higher percentage of polyphenols, when
deciduous forest (TOF) and tropical semi-evergreen
compared with the content in unconsumed plants. However,
forest (TSF) during the rainy and dry seasons.
this percentage covers total polyphenols, and including a
considerable amount of other phenol ic compounds which are
DISCUSSION different from tannins. In particular, it will be necessary to
The white-tailed deer face a temporal (from rainy to dry quantif)r the percentage of hydrolyzable and condensed
season) and spatial (between TDF and TSF) variation in the tannins. Because the former hydrolize under gastric acid
forage quality in the tropical forest of Chamela. During the conditions and liberate trapped vegetable proteins (Robbins
rainy season, plants from the TDF were higher in nutritional el al. 1987). However, deer are adapted to tanninferolls
value than those from the TSF. This variation was due to fodders, having tannin binding proteins in their saliva (AlIstin
distinct phenologies ofthe plants in both communities. After el al. 1989). These aspects will be necessary to study in the
the first rain, plants from the TDF undergo a fast period of future in this tropical forest.
growth, and invest more energy in soft tissue growth rather
than in the development of woody structures. As a result, MANAGEMENT IMPLlCATIONS
plants are generally more palatable to deer (McCullough and
UlIrey 1985). During their growth phase, plants have a The dry season is the critical period to manage white-
higher concentration ofnitrogen compounds and are easier to tailed deer in this tropical forest, due to the low quality and
digest, and have lower fiber and lignin contents (Blair el al. scarcity offood, as well as lack ofwater. The actual popula-
1977, Short el al. 1974). Therefore, during the rainy season tion density (l0-14 deer/km 2 ) is below carrying capacity
the TDF offers deer greater species richness, higher produc- (Mandujano 1999); therefore, there is no apparent need to
tivity and availibility ofpalatable parts, and higher nutritional provide additional food under normal circumstances.
value of plants, when compared with plants of the TSF. However years ~ith low rainfall result in reduced forage
During the dry season, plants of both types of forest had production. Therefore, it is necessary to implement habitat
lower nutritional values. Protein and NFE decreased as the management practices. One possibility would be to comple-

VIDA SILVESTRE NEOTROPICAL 8(1-2):1999 DEER FORAGE QUALlTY IN TROPICAL FOREST· Silva-Villalobos et al. 41
ment native food sources with nutritionally-balanced food as dac). Oikos 79:209-218.
well as sufficient water, at specific places within the forest, HENKE, S. E., S. DEMARAIS and .J. A. PFISTER. 1988. Digestive
principally when the winter rains are scarce. This is espe- capacity and diets ofwhite-tailed deer and exotic ruminants.
cially important because diminished winter rains usually Journal of Wildlife Management 52:595-598.
result in a lower standing plant biomass and less available LESLl-HARTI, F. and F. H. JOHNSTONE. 1989. Análisis moderno
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