BlOTROPlCA 32(3): 523-532 2000

Quetzal Abundance in Relation to Fruit Availability in a Cloud Forest

in Southeastern Mexico’
Sofia Solorzano’
Laboratorio de Ecologia, Facultad de Ciencias, UNAM. Circuito Exterior, Ciudad Universitaria. Mexico D.F.,
04510, Mexico
Silvia Castillo
Laboratorio de Ecologia, Facultad de Ciencias, UNAM. Circuito Exterior, Ciudad Universitaria. Mexico D.F.,
04510, Mexico
Teresa Valverde
Laboratorio Especializado de Ecologia, Facultad de Ciencias, UNAM. Circuito Exterior, Ciudad Universitaria,
Mexico D.F., 04510, Mexico
Lourdes Avila
lnstituto de Historia Natural, Tuxtla Gutierrez, Chiapas, 29000, Mexico

ABSTRACT
The Resplendent Quetzal (Pbaromacbrus mocinno) is an altitudinal migrant that nests in high elevation cloud forests
and migrates toward lower areas during the summer rainy season. It has been suggested that its migratory movements
are related to the abundance of ripe Lauraceae fruits. We studied the quetzal diet during two consecutive years, as
well as changes in fruit abundance of the plant species on which the bird feeds at El Triunfo Biosphere Reserve,
southeastern Mexico. The quetzal was obscrved feeding on 32 plant species; of these, 24 are new records in its diet.
We chose 20 of these 32 species and studied their fruit phenology for two years in order to describe the relationship
between fruit and quetzal abundance. Our results showed that quetzal abundance in the breeding area was correlated
with the total number of fruiting species, whereas the correlation between quetzal abundance and the number of
fruiting Lauraceae species was only marginal. Additionally, a correlation test showed that quetzal abundance was
marginally correlated with total fruit availability (total no. of fruits per month); however, the correlation between
quetzal abundance and the number of fruits in the Lauraceae was not significant. Our results suggest that the dynamics
of food resources may be playing a major role in the quetzal’s migratory behavior. Knowing the birds diet may aid
in characterizin the type of habitat adequate for its conservation. Our observations in this respect suggest that
conservation effgorts to preserve this bird species should concentrate on the protection of its habitat, including both
breeding and nonbreeding (migration) locations.

RESUMEN
El Quetzal (Pbaromacbrus mocinno) es una especie migratoria altitudinal que anida en las partes altas de 10s bosques
de niebla y migra hacia areas mas bajas en el verano, durante la estaci6n de Iluvias. Se ha sugerido que sus movimientos
altitudinales estin relacionados con la abundancia de frutos maduros de la familia Lauraceae. Estudiamos la dieta del
quetzal durante dos afios consecutivos, asi como 10s cambios en la abundacia de frutos de las especies de las cuales
10s quetzales se alimentan en la reserva de la biosfera El Triunfo, en el sureste de Mexico. Los quetzales se observaron
alimentindose de 32 especies de plantas, de las cuales 24 son nuevos registros en su dieta. Durante dos afios estudiamos
la fenologia de frutos de 20 de a t a s 32 especies, para describir la relaci6n entre la abundancia de frutos y la de
quetzales. Nuestros resultados muestran que la abundancia de quetzales en el irea de reproducci6n est6 correlacionada
con el nhmero total de especies que fructificaron, mientras que la correlaci6n entre la abundacia de quetzales y el
nlimero de especies de Lauraceae con frutos fue marginal. Otra analisis mostr6 que la abundancia de quetzales est6
marginalmente correlacionada con la disponibilidad total de frutos (n6mero total de frutos por mes); sin embargo, la
correlacibn entre abundancia de quetzales y nhmero de frutos de Lauracea no fue significativa. Nuestros resultados
sugieren que la dinimica de 10s recursos alimentarios puede estar jugando un papel importante en la conducta
migratoria del quetzal. El conocimiento de la dieta del quetzal puede ayudar en la caracterizacibn del hibitat que es
adecuado para su conservaci6n. Nuestras observaciones sugieren que 10s esfuerzos de conservaci6n para preservar esta
especie deben concentrarse en la protecci6n de su hibitat, tanto de las ireas de reproducci6n como de migraci6n.

Key words: altitudinal migration; Cbiapm; cloud forest; El fiiunfo; JFuitpbenology; Pharomachrus mocinno; Resplen-
dent Quetzal.

’ Received 12 August 1998; revision accepted 2 June 1999.

Current address: Instituto de Ecologia, UNAM. Campus Morelia. Laboratorio de Ecologia GenCtica y Molecular,
Antigua Carretera a Pitzcuaro no. 8701. Ex.-Hacienda de San JosC La Huerta, Morelia Michoacin, 58190. MCxico;
e-mail: slsofia@ate.oikos.unam.mx.

523

524 Solorzano, Castillo, Valverde, and Avila
QUETZAL
THERFSPLENDPNT (Pharomachrus mocin-
no) has been classified as a highly threatened species
(King 1981, Diario Oficial de la Federaci6n 1994),
mainly because illegal trade and habitat destruction
have caused a dramatic population decline during
the last decades (King 1981, Hanson 1982, Collar
1992). Its distribution ranges from southern Mex-
ico to Panama (Land 1970, Wetmore 1970, Peter-
son & Chalif 1994, Howell & Webb 1995), within
which two subspecies are distinguished: Pharo-
machrus mocinno costarkensis found in Costa Rica
and Panama, and l? m. mocinno from southern
Mexico to Nicaragua.
The Resplendent Quetzal has been considered
a specialisr frugivore; adults have never been ob-
served feeding on other dietary sources. Some small
animals such as lizards, frogs, snails, and insects,
however, are an important part of the nestling diet
(Skutch 1944, Wheelwright 1983, Avila et al.
1996). The adult has been observed feeding on 15
fruit species in southeastern Mexico (Avila et al.
1996), while Wheelwright (1983) reported 38 spe-
cies as part of the quetzal’s diet in Monteverde,
Costa Rica.
The Resplendent Quetzal is an altitudinal mi-
grant (Wheelwright 1983, Loiselle et al. 1989,
Powell & Bjork 1994). In southeastern Mexico (Si-
erra Madre de Chiapas), its reproductive period oc-
curs from January to June, during which this spe-
cies remains in cloud forest at elevations between
1800 and 2000 m (Sol6rzano 1995). The species
migrates to lower areas (900-1400 m; lower mon-
tane forest) in July and returns to the cloud forest
in early December (Sol6rzano 1995). Wheelwright
(1983) has suggested that the quetzal’s migratory
behavior is associated with the abundance of fruits
in the Lauraceae family (wild avocados), which are
known to be a basic element in the birds diet.
To determine the precise role of food abun-
dance on the migratory behavior of this bird spe-
cies, it would be necessary: (1) to detect all the food
sources on which the quetzal feeds; (2) describe the
changes in the abundance of these food sources
through time; and (3) experimentally determine
the quetzal’s food preferences. This study addressed
the first two questions through investigation of the
quetzal’s diet in southeastern Mexico and analysis
of the phenological patterns of the fruit species in-
volved. The results of such an approach could offer
a basis to investigate further questions related to
the precise effect of food abundance on the quet-
zal’s life cycle and migratory patterns. Thus, our
main objectives were to describe quetzal abundance
patterns during two consecutive years and to relate
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them to the availability of its food sources in one
of its breeding sites in a cloud forest of southeastern
Mexico.
MATERIAL AND METHODS
This study was carried out at El Triunfo Biosphere
Reserve on the Sierra Madre de Chiapas, south-
eastern Mexico. Our research site was one of the
quetzal’s breeding areas in the reserve (1800 m
elev.); this area of ca 10,000 ha includes a large
portion of cloud forest (Long & Heath 1991). The
climate at this site is temperate, humid with a
marked summer rainy season (Garcia 1988). An-
nual rainfall ranges from 2500 to 4500 mm. The
dry season lasts from December to early May.
Mean annual temperature varies from 14 to 18°C
(IHN 1993). The lowest temperatures (0-5°C)
usually are recorded during January or February
(Sol6rzano 1995).
QUETZAL
ABUNDANCE AND FEEDING HABITS.-TO
monitor variation in quetzal abundance at the
breeding site throughout the year, we established
six observation trails of 15 km total length. These
trails were at least 1 km apart, so that observations
on any one trail were independent of all others. We
visited the site each month from December 1991
to December 1993; during these visits we walked
along the trails (one day for each trail per month)
from 0600 to 0800 h. Quetzal abundance was reg-
istered along these trails by recording the number
of birds observed or heard during two-hour walks.
Monthly quetzal abundance was calculated as the
total number of quetzal sightings recorded during
these six-day observation periods.
To gather information on the quetzal’s food
sources at the study site, we carried out direct field
observations of feeding habits from December
1991 to December 1993 during daily walks (6 d/
mo) along the six observation trails at feeding times
(ca 0800-1000 and 1600-1800 h). From July to
October 1992 (the season in which the quetzal mi-
grates to lower areas), we also visited five different
migration sites (one 10-d visidsite) to record the
fruit species that the birds ate.
FRUII-rHENoLocx.--After the first three monrhs of
observations (December 1991, January and Feb-
ruary 1992) we had recorded the quetzal feeding
on 20 fruit species. We monitored the fruit phe-
nology of these species for the following two years
(December 1991-December 1993) in order to de-
scribe the quetzal’s yearly food availability at this
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Quetzal Abundance and Fruit Availability 525

TABLE 1. Plant species at which the quetzal was observed feeding The tree species included in the phenological study
were those f o r which a number of tagged individuals was reported. The right-hand column reports the elem-
tional range in which the birds were observed feeding from each species. Asterisks represent species present only
a t the nonbreeding sites. Species marked with (-) are reported for the jirst time as part of the quetzal? diet
in the study area.

No. of
tagFcd Elevational
Family Species individuals range (m)
Arecaceac -Chamaedorea sp. 0 1900
Actinidiaceae -Saurauia madrensis B. T. Kcller & Brcedlove 5 1900-2000
Celastraceac -Celastrus vulcanicola Donn. Sm. 0 2000
Lauraceae *Beilschmiedia sp. 0 1400-1760
-Cinnamomum zapatae Lorca-Hcrn. 6 1770-2000
Licaria excehu Kostermans 5 1900-2000
-L. glaberrima (Lundell) C. K. Allen 6 1800-2000
-Nectandra rudis C. K. Allen 9 1900-2000
-N.aff. salicina C. K. Allen 3 1900-1930
Ocotea acuminatissima (Lundell) Rohwer 5 1850-2000
-*O. brotantha Rhowcr 0 1400-1 630
-0. chiapensis (Lundell) Standl. & Steyerm. 40 1950-2000
-0. pkztyphylla (Lucndell) Rohwcr 5 1900-2000
Melastomataceae Conostegia volcanalis Standley & Steyerm. 11 1830-2000
Moraceae -'Ficus cookii Standley 0 1400-1 590
-*E costaricana (Licbm.) Miq. 0 1100
Morus insignes Bureau 11 1900-2000
-Gophis cuspidata Lundell 7 1900-1970
Myrsinaceae Ardisia compressa H.B.K. 16 1900-2000
-Synardisia venosa (masc) Lundell 4 1870-1 970
My ttaceae Eugenia captlli (Schl. Et Cham) Berget 5 1900-2000
I'odocarpaceac -Podocavpus matudue Lundell 0 1830-2000
Rhamnaceae -Rhamnus capraefolia var. grandifolia M. C. 8r L,. A. 5 1870-1970
Johnston
-*Rhamnus discolor (Donn. Smith) Rose 0 1240
Kosacear -Prunus tetradeniu Koehne 6 1900-1 930
-Prunus brachybotrya Zucc. 3 1870- 1950
Solanaceae -Lycianthes chiapensis (Brand) Standl 0 1870-2000
Symplocaceac -*Symplocos Jav ifolia Lundell 0 1100-1200
-Symplocos harwegii A. D .C. 0 1870-2000
Staphyleaceae -*Turpinia occidentalis (Sw.) G. Don subsp. occiden- 0 1200
talis
Theaceae Symplococurpon puvpusii (Brandegce) Kobuski 13 1800-1930
Verbenaceac Citharexvlum mocinnii D. Don. 5 1870-2000

site. It is important to mention that during these
two years of fieldwork, we were able to observe the
birds feeding on a larger group of species (32 to-
tal); however, we could not include all these species
in the phenological study because most of these
records were obtained toward the end of the study.
In any case, the 20 species recorded during the first
three months of observation appear to be the most
important elements of the quetzal's diet. With the
aim of monitoring the fruit phenology of these 20
species, we tagged a minimum of five individual
trees per species, as recommended for tropical trees
by Frankie et al. (1974). We chose those trees in
which at least 80 percent of the canopy could be
observed from the forest floor. The number of trees
tagged per species (Table 1) was determined sub-
jectively in relation to the apparent relative abun-
dance of each species in the forest. We could not
use more objective criteria to determine the num-
ber of trees to be tagged per species in relation to
their abundance per unit area because no infor-
mation on community composition and structure
was available for the study sites. A total of 170 trees
of 20 species were included in our phenological
observations. All ripe fruits in each tagged tree were
counted monthly with the aid of binoculars. For
trees in which the whole of the crown was not
observable from the forest floor, the observed frac-
tion was estimated, and the number of fruits re-
corded was extrapolated to the whole crown.
~ L A T I O N S H I P BETWEEN FRUIT AVAILABILITY AND
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526 Solorzano, Castillo, Valverde, and Avila

QLlF,TZAI, ABUNDANCE.-we estimated the total

number of fruits available to the quetzal within the
170 tagged trees each month as the sum of all *
counted fruits. Although this does not represent an
absolute measure of fruit availability, we believe it *
accurately reflects relative fruit abundance within
*
the observation trails where the birds were moni-
tored. We carried out a Spearman’s correlation test *
between the number of fruits and quetzal abun-
dance (as defined above). Since it has been sug- *
gested that the abundance of Lauraceae fruits are
particularly important in determining the quetzal’s *
migratory behavior (Wheelwright 1983), we car-
ried out another Spearman’s correlation test be- *
tween quetzal abundance and the abundance of
* *
fruits in the Lauraceae only. Finally, to test whether
quetzal abundance was related to the number of *
species bearing fruits, or to total fruit availability
as suggested by Wheelwright (1986), we correlated *
quetzal abundance against the number of species
bearing fruits (for all of the studied species and the
Lauraceae only).

RESULTS *
QLJF
1LAL ABUNDANCF AND FEEDING HABITS.-The
*
number of quetzal sightings recorded throughout
the year along the trails at the study site demon- *
strated that the quetzal was most abundant in this
area from January to June, which coincided with *
the breeding season (Table 2). During these
months, the bird was observed engaging in court- *
ship activities, as well as nesting and chick rearing.
x
Note that a few birds were observed in the study
site during the nonbreeding season between July
*
and November (Table 2).
During the two years of monthly observations * *
at both the breeding and nonbreeding sites, we saw
the quetzal feeding at a total of 32 tree species * *
(Table 1). Plants of the Lauraceae and Moraceae
made up a large proportion of these species (31.5 i *

and 12.5%, respectively). O f the species reported

in Table 1, most of them (78%) corresponded to t

cloud forest species, while seven were present only

i
at the nonbreeding sites. O f the 32 species record-
ed, 24 (75%) had never been reported as part of
the quetzal’s diet at the study site (Avila et aL.
1996).

FRUITPI4ENOLOGY.-MOSt of the observed tree spe-

cies produced fruits only during a specific period
a
of the year, which was as short as three months in
Cinnamomum zdpatae and Morus insigne5 or as long
as seven to ten months in CitharexyLum mocinnii,

Conostegia volcanalis, Nectandra aff. salicina, and
Nectandra rudis (each of which peaked during a
particular season of the year; Table 3). Some spe-
cies, however, such as Ardisia compressa, Licaria ex-
ceha, L. glaberrima, Ocotea cbiapensis and Prunus
tetradenia, bore fruits all year long, mainly due to
the asynchrony in fruit production within individ-
uals of the same species (Table 3). The low number
of fruits per tree recorded in some months on these
species can be accounted for by the fact that only
a small proportion of the tagged trees bore fruits.
Note that 4 out of 20 species produced fruits in
only one of the yearly periods studied: C. zapatae
produced fruits only in 1992, I! bracbybotya and
7: cuspidata fruited only in 1993, and S. purpusii
only in 1991 and 1993 (Table 3).
At our study site, 50 percent in 1992 and 45
percent in 1973 of the studied species produced
their highest fruit numbers during the dry season
(December-May), whereas 40 percent in 1992 and
35 percent in 1993 did so during the rainy season
(June-November). Each month there were at least
six species bearing ripe fruits. Months with the
highest number of species in fruit were February,
March, and April in 1992, and March, April, and
June of 1993. The months with the lowest number
of species bearing fruits were December of 1991
and 1992, and October and November of 1993
(Fig. 1).
RELATIONSHIP BETWEEN FRUIT' AVAILABILITY A N D
QUETZAL ABuNDANcE.-In general, the months in
which the quetzal was abundant at the breeding
site were those corresponding with the highest fruit
availability (Fig. 2). Total fruit availability (i.e., the
total no. of fruitdm0 in the tagged trees of the 20
spp.) and quetzal abundance (no. of sightingslmo),
were marginally correlated ( r = 0.43, P = 0.08, N
= 17 monthly observations). Additionally, quetzal
abundance and the total number of species bearing
fruits were significantly correlated ( r = 0.49, P =
0.04, N = 17). O n the other hand, the correlation
between quetzal abundance and the total number
of fruits in the Lauraceae was not significant (r =
0.35, P = 0.16, N = 17), while quetzal abundance
was only marginally correlated with the total num-
ber of fruiting Lauraceae species ( r = 0.44, P =
0.08, N = 17). These results showed that quetzal
abundance at the study site appeared to be associ-
ated mainly with the number of species bearing
fruits, and less clearly to total fruit availability and
to the number of Lauraceae species bearing fruits.
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Quetzal Abundance and Fruit Availability 527
DISCUSSION
We recorded 32 species as actual food sources for
the quetzal; of these, only 8 had been reported be-
fore as part of the birds diet in the study area (Avila
et al. 1996). Thus, as has been suggested, the quet-
zal is a frugivorous species using a wide range of
fruit types (McKey 1975; Wheelwright 1983,
1986; Wheelwright et al. 1984; Avila et al. 1996).
More detailed experimentation would be necessary
to assess if the quetzal prefers any particular fruit
type.
Although our results suggest that quetzal abun-
dance is associated with the number of species
bearing fruits at the study site, it would be neces-
sary to assess quetzal and fruit abundance on a per-
area basis, both in the breeding and nonbreeding
sites, before the effect of food availability on mi-
gratory behavior can be ascertained more firmly.
The evidence available does not allow us to con-
clude that food availability is the causal factor that
determines migration in this species. Our results,
however, clearly suggest that the dynamics of food
resources may be playing a major role in this re-
spect.
As has been noted by other authors (Skutch
1944, Wheelwright 1983, Avila et al. 1996), fruits
of plants in the Lauraceae were found to be an
important element of the quetzal's diet (Table 1).
Our data, however, indicate that quetzal abundance
at the study site was not significantly correlated
with the total availability of fruits in this plant fam-
ily; instead, it was associated with the total number
of fruiting species in the forest during the different
seasons of the year. Yet, it is important to note that
our phenological study included only the first 20
species from which the quetzal was observed feed-
ing; nonetheless, after two years of observations we
had recorded a total of 32 species as part of the
bird's diet. Thus, a more complete assessment of
the quetzal's food distribution in space and time
would be necessary before reaching firm conclu-
sions about the role of food availability in its mi-
gratory behavior. For example, plants in the genus
Ficus and Symplocos (not studied), which had a
large number of ripe fruits during the nonbreeding
season, were the main food sources at the five mi-
gration sites visited. O n the other hand, fruits of
Celastrus uulcanicola and Qciantbes cbiapensis (also
not studied) were important food sources for the
birds that remained in the breeding site during the
nonbreeding season.
Regarding the fruit phenology of the plant spe-
cies studied, we observed that although different
 TABLE 3. Number o f ripe fruits per species from December I991 to December 1993. Asterisks represent months in which observations were not made. The right hand coluwin (N)
indicates the number of trees tagedper species. The rows in italics indicate the number of trees fruiting each month of the total number of trees follouedper species.
~~

Month D J F rvi A M J J A S O N D
Year 199 1 * 1992
Ardisia compressa 181 326 462 124 3126 269 1111 68 118
9 I2 I3 IG I6 I2 I4 7 4
Cinnamomum zapatae 32782 48506 666
5 5 5
Citharexylum mocinnii 200 8800 724 1 4030 16374
I I I 1 4 4
Conostegia volcanalis 11527 1326 3252 4616 43 1 155
4 G 5 8 7 5
Eugenia capuli 2125 194 4 134 234 72
4 3 1 1 3 2
Licaria excelsa 6 3167 4280 34 3340 154 1008 8020 1720
I I 2 3 4 4 4 4 4
L. glabewima 8 75 750 1150 3900 245
I I 2 2 3 2
Morw insignes 7485 3544 875
4 5 1
Nectandra aE. salicina 140 775 403 1917 1243 683 36
I I 3 3 3 3 I
N. rudis 47 282 176 18 20 3 98
I I 2 I I I I
Ocotea acuminatissima 6 168 174 56 28 18
I 3 4 2 2 2
0. chiapensis 16 206 27 143 13 10 31 17
17 13 12 5 I 9 I8 13
0.plathyphylla 9162 93 532 77 22
2 2 I I I
Prunus brachybotiya

I? tetradenia 15597 40644 17867 988 5523 13614 1214

6 7 7 7 6 1 I
Rhamnus capraefolia var. grandifolia 113 125 10716 17256 22708 1866
3 3 3 5 5 5
Saurauia madrensis 249 64 17 82
I I I 1
Symplococatponputpusii 110 3827 464 2686
6 4 5 13
Synardisia venosa 3575 2502 1912
4 4 4
Trophis cuspidata

3 4 5 3

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 TABLE 3. Extended.

Monrh J F M A M J J A S 0 N D
Year 1993 * * * N
Ardisia compressa 233 1158 3892 198 208 47 33
12 9 9 14 I 13 15 16
Cinnamomum zpatae
5
Citbareqlum mocinnii 1620 16732 5479 1764 22518 3012 214
2 4 5 5 4 1 I 5
Conostegia volcanalis 1212 8921 3358 2442
4 7 7 6 8
Eugenia capuli 34 42 234 16 1128
I 2 2 I 2 5
Licaria excelsa 5 1220 120 3640 568 3696
I 3 1 4 2 5 5
L. glaberrima 125 575 825 1475 765
1 1 2 2 2 6
Morus insignes 24562 15925 225
10 5 I 10
Nectandra aK. salicina ~~
163 233 12 3467 2007
3 3 3 2 2 13
N. rudis 114 28 48 446 238 14 484 636 22
1 2 2 2 I I 2 3 2 9
Ocotea acuminatissima 146 528 332 32 6
2 5 2 I I
0. cbiapensis 45 112 563 98 182
12 15 15 17 15 21
0. platbypbylla 522 1175 1232 912
3 1 1 2 2 2 5
Prunus bracbybottya 260 78 C
2 3
2 Q
I! tetradenia 1607 273 1 543 1567 91 270 3943 I
3 4 6 5 1 5 6 6 0
(D
Rbamnus capraefolia var. grandgolia 24752 23584 17664
4 5 5 5
I
Q
Saurauia madrensis 16 284 83 -n
7
I I 4 5 5.
Symplococarponpurpusii 26 697 768 -
2 4 7 I3 D
<
Synardisia venosa E.
4 5
F.
-
-.
Tropbis cuspidta 7467 2507 1017
-7 u"
/
4 4 3 5 5 2 5
u1
N
(0

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 17447429, 2000, 3, Downloaded from https://onlinelibrary.wiley.com/doi/10.1111/j.1744-7429.2000.tb00498.x by Universidad Veracruzana, Wiley Online Library on [16/08/2023]. See the Terms and Conditions (https://onlinelibrary.wiley.com/terms-and-conditions) on Wiley Online Library for rules of use; OA articles are governed by the applicable Creative Commons License
530 Solorzano, Castillo, Valverde, and Avila

N = 20
Migration Migration Fleming 1991, Loiselle & Blake 1991). Our study
'61 Reproduction ~
__
Reproduction
~ ~~
~ ~

supports this hypothesis and represents an example

I) J Y M A M I J A s o N u J F M A M J J A s
Y e a t 9 1 92 93
Months
FIGURE 1. Number of species bearing fruits each
month from December 1991 to December 1993 at the
quetzal's breeding site. Asterisks indicate months in which
no observations were made. The length of the different
phases of the quetzal's yearly cycle is indicated above the
bars in each graph.
fruits were available all year long (Table 3), fruit
availability was highest during the dry season (Figs.
1 and 2). This pattern has been described for plants
in other cloud forests (Wheelwright 1986, Seres &
Ramirez 1993) and appears to be characteristic of
other tropical ecosystems in general (Frankie et al.
1974). It has been suggested that this maximum in
fruit production coincides with the highest frugi-
vore abundance in these habitats (Terborgh 1986,
of the type of information that can be attained
from a combined analysis of fruit phenology and
bird abundance.
Our observations support the idea that the
quetzal's reproductive cycle is closely coupled with
general fruit availability (Wheelwright 1986); the
main reproductive activities (courtship, nesting,
chick rearing) occur when fruits are most abundant
in the cloud forest (Table 2; Fig. 2). This may allow
o N u
the birds to face this high energy-demanding phase
when food abundance is highest, which may in-
crease the probability of successful chick rearing.
Later, when the rainy season starts, most members
of the species migrate to lower areas.
If quetzal populations are to be maintained, de-
forestation and habitat loss must be stopped, not
only in the main breeding areas within the cloud
forest, but also in its different migration habitats.
At El Triunfo Biosphere Reserve, the core areas are
occupied mainly by cloud forest, while the buffer
zones are lower montane forest. Several farming
and agricultural activities are allowed in these buff-
er zones; as a result, the vegetation at these sites is
highly disturbed and contains some grassland areas
and coffee plantations. The abundance and diver-
sity of the quetzal food sources are declining at
these sites, which constitutes an important threat
to its persistence in this area. Additionally, in most

120000
- -0- -Total o f fruits 80

70
100000 +Lauraceae fruits

60
-0- No.of Quetzals
80000
50 3
m
c

$ 60000
9 40
8
W

8
8

z 40000
30 2
20
20000
10

0 0
D F M A J A S N D J F M A J J O N D
91 92 93
Months

FIGURE 2. Variation in quetzal abundance (no. of quetzal sighting per month on the observation trails), total
fruit abundance (total no. of fruits in the tagged trees of the 20 species studied at the breeding site) and fruit abundance
in the Lauraceae (no. of fruits in the tagged trees of the Lauraceae) during two years. Asterisks represent months in
which no observations were made.
 17447429, 2000, 3, Downloaded from https://onlinelibrary.wiley.com/doi/10.1111/j.1744-7429.2000.tb00498.x by Universidad Veracruzana, Wiley Online Library on [16/08/2023]. See the Terms and Conditions (https://onlinelibrary.wiley.com/terms-and-conditions) on Wiley Online Library for rules of use; OA articles are governed by the applicable Creative Commons License
Quetzal Abundance and Fruit Availability 531

of the quetzal’s range, habitat fragmentation has
been extensive during recent decades. As a result,
quetzal populations are declining and their degree
of isolation is increasing rapidly (Powell & Bjork
1994). Conservation activities must be coupled
with development; the protection of natural areas
should consider both the needs of the local human
communities and the urgency for wildlife conser-
vation. Therefore, we must make the best of our
conservation efforts through the adequate design of
nature reserves, considering the creation of reserve
networks linked through wildlife corridors (Harri-
son 1994, Powell & Bjork 1994, Wilcove 1994).
In the case of the quetzal, these corridors would
increase its persistence probabilities by providing
adequate breeding and migration sites and decreas-
ing the isolation of populations.
ACKNOWLEDGMENTS
We are grateful to the Instituto de Historia Natural dc
Chiapas for allowing us to carry out fieldwork at El
Triunfo. We thank Rafael Solis, Ismael Gilvez, and Laura
Noble for their assistance in different field activities, Mar-
co Romero for technical support, and Guillermo Ibarra
and Francisco Lorea for taxonomic assistance. We also
wish to thank G. Powell and J. Shopland for their en-
couragement and support in the development of this pro-
ject. This study was supported by Rare Center for Trop-
ical Conservation and the Nature Conservancy. The com-
ments of H. G6mez de Silva, J. F. Ornelas, and N.
Wheelwright greatly improved the quality of this paper.

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