Blake FruitsDietsNeotropical 1992
Blake FruitsDietsNeotropical 1992
Blake FruitsDietsNeotropical 1992
JSTOR is a not-for-profit service that helps scholars, researchers, and students discover, use, and build upon a wide
range of content in a trusted digital archive. We use information technology and tools to increase productivity and
facilitate new forms of scholarship. For more information about JSTOR, please contact [email protected].
Your use of the JSTOR archive indicates your acceptance of the Terms & Conditions of Use, available at
https://about.jstor.org/terms
Association for Tropical Biology and Conservation is collaborating with JSTOR to digitize,
preserve and extend access to Biotropica
Department of Biology, University of Missouri-St. Louis, 8001 Natural Bridge Road, St. Louis, Missouri
63121, U.S.A.
ABSTRACT
We used fecal samples collected from birds captured in mist nets to determine the occurrence of fruit in the diets
of neotropical migrant birds (species that breed in North America and winter in Central and South America) at six
study sites in Costa Rica. A total of 1016 migrants representing 30 species were captured. Fecal samples were collected
from 85 percent of all birds captured. Fruits (seeds, pulp) occurred in 58 percent of all fecal samples; the proportion
of fecal samples containing fruit ranged from 0 percent (9 species) to over 90 percent (9 species). Individual fecal
samples contained seeds from as many as 4 species of plants. At least 98 plant species from at least 29 plant families
were represented in fecal samples. Species of Melastomataceae accounted for 37 percent and those of Rubiaceae for
16 percent of all species recorded. Number of fruit species in diets of individual migrant species ranged from 0 (9
species) to 63 (Swainson's Thrush, Catharus ustulatus). Our data demonstrate that some migrants may rival permanent
residents in the diversity of fruits they consume. A shift to increased frugivory in the late dry season, as birds were
preparing to migrate north, was noted.
A MAJORITY OF BIRDS THAT BREED in temperate North reported on diets of migrants in the tropics have
America spend most of the year in the tropics ("neo- tended to focus on relatively few species or on use
tropical migrants" hereafter; Rappole et al. 1983). of one specific plant species by different birds (e.g.,
Despite much recent attention given to apparent Morton 197 1, Greenberg 198 1). A comprehensive
declines in breeding populations of many of these analysis of migrant diets from any location, similar
species (e.g., Terborgh 1989, Askins et al. 1990, to the analysis of all fruit-eating birds at Montever-
Hagan & Johnston, in press), the habitat require- de, Costa Rica, by Wheelwright et al. (1984), has
ments and diets of most species are poorly known not been presented previously.
during nonbreeding seasons (Keast & Morton 1980). We have been studying habitat use and diets
Yet understanding habitat and diet requirements of of migrant (and nonmigrant) birds in several hab-
neotropical migrants throughout the year is neces- itats in Costa Rica since 1985 (Loiselle 1987b,
sary if we are to understand population dynamics Blake et al. 1990, Blake & Loiselle 1991, Loiselle
of these species and develop adequate plans for their & Blake 1991). Elsewhere (Blake & Loiselle, in
conservation. Population declines of neotropical mi- press) we report on patterns of habitat use by neo-
grants are, for example, most likely a consequence tropical migrants. Here we summarize data collected
of events on both breeding and wintering grounds on diets of migrants at our study sites.
(Hutto 1989, Terborgh 1989, Askins et al. 1990,
Blake et al., in press) (e.g., breeding habitat frag-
mentation, tropical deforestation). STUDY AREA
Most neotropical migrants feed primarily on
Study sites were on the Caribbean slope of the
insects during the breeding season; many, however,
Cordillera Central in northeast Costa Rica. Lowland
eat much fruit while in the tropics (Morton 1971,
habitats were 5-10 yr second-growth (one site), 2 5-
1980; Leck 1972a, b; Howe & DeSteven 1979;
35 yr second-growth (one site), and primary (un-
Martin 1985b; Martin & Karr 1986; Blake et al.
disturbed) forest (two sites) at the La Selva Biolog-
1990; Loiselle & Blake 1990) and may be impor-
ical Station (10025'N, 84001'W), a field station of
tant seed dispersers for some plants (Greenberg
the Organization for Tropical Studies. We also sam-
1981). Despite the apparent importance of fruit as
pled diets of birds in primary forest at 500 m (one
a resource, little quantitative information on the
site at 10'20'N, 84?04'W) and at 1000 m (one
occurrence of fruit in the diets of most overwintering
site at 10'16'N, 84?05'W) in Braulio Carrillo Na-
migrants is available. Previous studies that have
tional Park, about 15 and 20 km south of La Selva,
respectively. Detailed descriptions of these study
sites are in Blake and Loiselle (1991) and Loiselle
I Received 10 December 1990, revision accepted 15 July
1991. and Blake (1991).
200
La Selva receives about 3900 to 4000 mm rain frequently are hard to observe, so that visual records
annually (Hartshorn 1983, Organization for Trop- of fruit consumption often are incomplete or are
ical Studies, pers. comm.). The main dry season biased by observations at conspicuous plants with
lasts from January or February to March or Aprillarge crops.
with a shorter, less pronounced dry season in Sep-
tember and October. Climatological data are not
RESULTS
available for the 500 and 1000 m sites, but annual
rainfall probably exceeds 4500 mm at both. General We captured 1016 migrants representing 30 species
descriptions of La Selva and Braulio Carrillo are in (Table 1). The most common species induded Wood
Hartshorn (1983), Pringle et al. (1984), and Hart- Thrush, Swainson's Thrush, Ovenbird, and Ken-
shorn & Peralta (1988). tucky Warbler which together comprised 72 percent
of all migrants captured. We collected 864 fecal
samples. The number of fecal samples collected
METHODS
from different bird species ranged from 1 (6 species)
Mist nets (12-m, 4-shelf, 36-mm mesh) were used to 259 (Wood Thrush) (Table 1). All 30 migrant
to sample birds in the lower levels of each habitat species consumed insects, as indicated by presence
(Blake & Loiselle 1991, Loiselle and Blake 1991). of insect parts and uric acid in most (83%) fecal
Captured birds were banded, weighed, held for col-samples. Percentage of samples with insects ranged
lection of fecal samples, and released at the point from 11 percent (Eastern Kingbird) to 100 percent
of capture. We collected samples every 5 to 6 weeks (21 species).
from January 1985 through April 1986 and from Fruit seeds or pulp occurred in 58 percent of
December to April 1986-1987 and 1987-1988. all fecal samples. The percentage of species-specific
Diet analyses were based on seeds and pulp samples with fruit ranged from 0 (for 9 species) to
from feces or regurgitated material (hereafter re- over 90 (in 9 other species, including 4 represented
ferred to collectively as "fecal samples") from mistby > 5 fecal samples). All species represented by at
netted birds. We placed captured birds in plasticleast 5 samples consumed some fruit, except for
containers lined with filter paper for 5 to 15 min. Mourning Warbler and Canada Warbler (Table 1).
Using a dissecting microscope, we separated seeds Highly frugivorous species included Swainson's
from fecal samples and identified them to species Thrush, Gray-cheeked Thrush, Eastern Kingbird,
through comparison with a reference collection at Gray Catbird, and Traill's Flycatcher. More than
La Selva. Some seeds were lumped by genus because one fruit species often occurred in a single fecal
we could not distinguish species (e.g., Anthurium, sample (Table 1); Gray-cheeked Thrushes averaged
Ficus). Further details are in Loiselle and Blake two fruit species per sample, for example. The max-
(1990). imum number of fruit species represented in a single
The advantages and disadvantages of using fecaldropping was four.
samples to determine diets of birds have been dis- We recorded at least 98 plant species from at
cussed elsewhere (Wheelwright et al. 1984, Loiselleleast 29 families (Appendix) in samples collected
& Blake 1990); here we summarize previous dis- from 21 of the 30 species of neotropical migrants
cussions. Differential passage rates of small and large captured (Table 2). (The actual number of species
seeds (Levey 1986, 1987) may lead to an overes- represented is probably higher because seeds of some
timation of the importance of small-seeded fruits samples were identified only to genus.) Most plant
in diets of some bird species. Further, some birds species were either members of the Melastomataceae
may drop rather than ingest seeds of some fruits (35 species) or Rubiaceae (16 species), two of the
(Moermond & Denslow 1985, pers. comm.), lead- most common families in the understory of our
ing to an underestimation of those fruits in the diet.
study sites at La Selva and Braulio Carrillo (Loiselle
Similarly, seeds of some fruits are regurgitated rather
1987a).
than defecated by some bird species; regurgitation Number of identified fruit species in diets of
often occurs, however, while the birds are being bird species ranged from 1 (4 bird species) to 52
held. Despite these problems, collection of fecal (Wood Thrush) and 63 (Swainson's Thrush). Over
samples is an effective way to gather data on fruit half (58) of all fruit species were recorded in the
consumption (Wheelwright et al. 1984, Loiselle & diets of only one bird species; 27 plant species oc-
Blake 1990). Fecal samples are not subject to manycurred only in samples collected from Swainson's
Thrushes and 22 species only from Wood Thrushes.
of the biases inherent in direct observations of fruit
consumption. Birds foraging in forest understory Plant species consumed by the greatest number of
TABLE 1. Number of captures and number offecal samples collected from migrants. Number and pe
of samples with insect parts or with fruit seeds or pulp are indicated. Mean, SD, and m
plant species represented in a fecal sample are given for samples with fruits.
Total
Total fecal
Bird
Bird cap- sam-tcap- scam Samples with Plant sp
Bird species code tures ples Insects Fruits Mean SD Max
TABLE 1. Continued.
Total
E M L E M L E M L E M L
DISCUSSION 1 985 1 986 1 987 1 988
D RY SEASON SAMPLES
Many species of migrants are known to consume
fruit while on their nonbreeding grounds (Rappole
FIGURE 1. Percentage of fecal samples from all species
et al. 1983). Over 80 species of neotropical migrant and from Wood Thrush separately that contained fruit
birds occur at La Selva Biological Station and in during early (E: late December-January), middle (M:
Braulio Carrillo National Park (Blake et al. 1990, February-mid March); and late (L: late March-April) dry
seasons. Data are pooled from all habitats and study sites.
Blake & Loiselle, in press). Most neotropical mi-
t M;{ > o -
_7 rq
0 Iq -4 Iq - r 4r- N . 0 -4- nc f,'-- c qc qC4-4- C . ' , ~r n'
II~ 0
>, | bt _~i
>4
0
2 b | i :; - : t (
( |WN I
~~ tr\ ~ ~ r
et b z # 0i
il!
4 5
00 -4
U
44 D
.4
u
z
0 W
71
P4 '.4 Cd
0 a)
Cf) 00 Z
z U
0
2pu O Cd
u 'o
(U 11)
r4 O o
Cd (N
u
Z 0
0 U
u
r14 en O rq r- r\l 0 cd m
rq vl 0 04 4u --4
u
D Cd U W D
cd ri) U I - 0
(j
oo
C\ 't 0 00
u (U W4 P-4
-0 u 4 4
cd
0
75
-10
.!z
u >1 P4 V) r4
U Z
(U
Cd u
uuuo
.4 U
0
r,14
>
cd
,d C-4 C\
tn ZP4 00
U td
0 U
ti u k42
P4
v J: C\ C\
> 0 0
> m 0 W 0
C14 \D z V) cd V
-':v P4 0 7ZL
0 u O
C: U -.- W
Qi 4-1 v
Z 4
C)
0- & u
V) 0 E
Z
U u H uU) W-4
n
P4 P4 ,, W-4
O\ C\ '.4 M.4
m u 0 0 O. > P4 0-4 W z
t4 Cl U C:
TABLE 3. Number offecal samples (N), percent with insects (% I), and percent with fruit (% F) co
in different habitats. Bird species codes in Table 1.
Habitat N %I %F N %I %F N %I %F N %I %F
Primary forest
Lowland (50 m) 158 86 63 3 67 100 27 41 100 115 96 59
Foothill (500 m) 84 87 60 20 70 95 59 92 51
Premontane (1000 m) 24 88 71 13 77 100 2 100 100
Habitat N %I %F N %I %F N %I %F N %I %F
Loiselle & Blake 1991). A higher degree of frugivo- (e.g., Raven 1988); conversion of second-growth
ry during migratory periods may have important habitats to pasture will have negative effects on
consequences for seed dispersal; birds may be more many species as well. Effects may be felt not only
likely to move greater distances at this time than by second-growth species but also by forest species
during the winter, when many are on winter ter- that use second-growth on a seasonal or irregular
ritories. basis. Thus, it is appropriate to incorporate consid-
Abundant supplies of fruit in many second- eration of second-growth habitats when discussing
growth habitats may be a partial explanation for strategies for habitat management and species pres-
ervation (Gilbert 1980).
the use of such habitats by many species of migrants
(Martin 1985b), as well as by many permanent
residents (Blake & Loiselle 1991). Fruit often is
ACKNOWLEDGMENTS
abundant in second-growth habitats throughout the
year (Levey 1988, Blake & Loiselle 1991) and may, We thank the Organization for Tropical Studies for per-
mission to work at La Selva and David and Deborah
as a consequence, be an important resource for many
Clark for facilitating our research there; Servicio de Par-
animals when fruit abundance is low in forests (Ter-ques Nacionales, particularly F. Cortes and J. Doblez, for
borgh 1985, Levey 1988, Blake & Loiselle 1991). permission to work in Parque Nacional Braulio Carrillo;
Many species, including both residents and mi- and Russ Greenberg and an anonymous reviewer for con-
grants, move from forest to second-growth during structive comments on the manuscript. Primary support
for this project was provided by: Jessie Smith Noyes
periods when fruit is low in abundance in forests
Foundation Fellowship (Organization for Tropical Stud-
(Martin 1985b, Martin & Karr 1986, Blake & ies); Michael H. Guyer Fellowship (Department of Zo-
Loiselle 1991). ology, University of Wisconsin); National Geographic
The adverse consequences to many species of Society; and the Douroucouli Foundation.
LITERATURE CITED
ASKINS, R. A., J. F. LYNCH, AND R. GREENBERG. 1990. Population declines in migratory birds in eastern North
America. Curr. Ornithol. 7: 1-57.
BLAKE, J. G., AND B. A. LOISELLE. 1991. Variation in resource abundance affects capture rates of birds in three
lowland habitats in Costa Rica. Auk 108: 114-130.
, AND B. A. LOISELLE. In press. Habitat use by neotropical migrants at La Selva Biological Station and
Braulio Carrillo National Park, Costa Rica. In J. Hagan and D. W. Johnston (Eds.). Ecology and conservation
of neotropical landbird migrants. Smithsonian Institution Press, Washington, D.C.
, G. J. NIEMI, AND J. M. HANOWSKI. In press. Drought and annual variation in bird populations. In J.
Hagan and D. W. Johnston (Eds.). Ecology and conservation of neotropical landbird migrants. Smithsonian
Institution Press, Washington, D.C.
, F. G. STILES, AND B. A. LOISELLE. 1990. Birds of La Selva: habitat use, trophic composition, and migrants.
In A. Gentry (Ed.). Four neotropical rain forests, pp. 161-182. Yale University Press, New Haven, Connecticut.
GILBERT, L .E. 1980. Food web organization and the conservation of neotropical diversity. In M. E. Soule and B.
A. Wilcox (Eds.). Conservation biology, pp. 11-33. Sinauer Associates, Inc., Sunderland, Massachusetts.
GREENBERG, R. 1981. Frugivory in some migrant tropical forest wood warblers. Biotropica 13: 215-223.
HAGAN, J., AND D. W. JOHNSTON (Eds.). In press. Ecology and conservation of neotropical landbird migrants.
Smithsonian Institution Press, Washington, D.C.
HARTSHORN, G. S. 1983. Plants: introduction. In D. H. Janzen (Ed.). Costa Rican natural history, pp. 118-157.
University Chicago Press, Chicago, Illinois.
, AND R. PERALTA. 1988. Preliminary description of primary forests along the La Selva-Volcan Barva
altitudinal transect, Costa Rica. In F. Almeda and C. M. Pringle (Eds.). Tropical rain forests: diversity and
conservation, pp. 281-296. California Academy of Sciences, San Francisco, California.
HOWE, H. F., AND D. DESTEVEN. 1979. Fruit production, migrant bird visitation, and seed dispersal of Guarea
glabra in Panama. Oecologia 39: 185-196.
HuTTo, R. L. 1989. The effect of habitat alteration on migratory land birds in a West Mexican tropical deciduous
forest: a conservation perspective. Conserv. Biol. 3: 138-148.
KEAST, A., AND E. S. MORTON (Eds.). 1980. Migrant birds in the neotropics: ecology, behavior, distribution, and
conservation. Smithsonian Institution Press, Washington, D.C.
LECK, C. F. 1972a. Seasonal changes in feeding pressures of fruit- and nectar-eating birds in Panama. Condor 74:
54-60.
1972b. The impact of some North American migrants at fruiting trees in Panama. Auk 89: 842-850.
LEVEY, D. J. 1986. Methods of seed processing by birds and seed deposition patterns. In A. Estrada and T. H.
Fleming (Eds.). Frugivores and seed dispersal, pp. 147-158. Dr. W. Junk, Publishers, Dordrecht, The
Netherlands.
1987. Seed size and fruit-handling techniques of avian frugivores. Am. Nat. 129: 471-485.
1988. Spatial and temporal variation in Costa Rican fruit and fruit-eating bird abundance. Ecol. Monogr.
58: 251-269.
LOISELLE, B. A. 1987a. Birds and plants in a neotropical rain forest: seasonality and interactions. Ph.D. Dissertat
University of Wisconsin, Madison, Wisconsin.
1987b. Migrant abundance in a Costa Rican lowland forest canopy. J. Trop. Ecol. 3: 163-168.
AND J. G. BLAKE. 1990. Diets of understory fruit-eating birds in Costa Rica: seasonality and resource
abundance. Stud. Avian Biol. 13: 91-103.
, AND . 1991. Resource abundance and temporal variation in fruit-eating birds along a wet forest
elevational gradient in Costa Rica. Ecology 82: 494-500.
MARTIN, T. E. 1985a. Resource selection by tropical frugivorous birds: integrating multiple interactions. Oecologia
66: 563-573.
1985b. Selection of second-growth woodlands by frugivorous migrating birds in Panama: an effect of
fruit size and plant density? J. Trop. Ecol. 1: 157-170.
, AND J. R. KARR. 1986. Temporal dynamics of neotropical birds with special reference to frugivores in
second-growth woods. Wilson Bull. 98: 38-60.
MOERMOND, T. C., AND J. S. DENSLOW. 1985. Neotropical avian frugivores: patterns of behavior, morphology,
and nutrition, with consequences for fruit selection. Ornithol. Monogr. 36: 865-897.
MORTON, E. S. 1971. Food and migration habits of Eastern Kingbirds in Panama. Auk 88: 925-926.
1980. Adaptations to seasonal changes by migrant land birds in the Panama Canal Zone. In A. Keast
and E. S. Morton (Eds.). Migrant birds in the neotropics: ecology, behavior, distribution, and conservation,
pp. 437-453. Smithsonian Institution Press, Washington, D.C.
PRINGLE, C., I. CHACON, M. GRAYUM, H. GREENE, G. HARTSHORN, G. SCHATZ, G. STILES, C. GOMEZ, AND M.
RODRIGUEZ. 1984. Natural history observations and ecological evaluation of the La Selva Protection Zone,
Costa Rica. Brenesia 22: 189-206.
RAPPOLE, J. H., E. S. MORTON, T. E. LOVEJOY, AND J. L. Ruos. 1983. Nearctic avian migrants in the neotropi
U.S. Department Interior, Fish and Wildlife Service, Washington, D.C.
RAVEN, P. H. 1988. Our diminishing tropical forests. In E. 0. Wilson (Ed.). Biodiversity, pp. 119-122. National
Academy Press, Washington, D.C.
TERBORGH, J. 1985. Five New World primates: a study in comparative ecology. Princeton University Press, Princeton,
New Jersey.
1989. Where have all the birds gone? Princeton University Press, Princeton, New Jersey.
WHEELWRIGHT, N. T. 1985. Fruit size, gape width, and the diets of fruit-eating birds. Ecology 66: 808-818.
, W. A. HABER, K. G. MURRAY, AND C. GUINDON. 1984. Tropical fruit-eating birds and their food plants:
a survey of a Costa Rican lower montane forest. Biotropica 16: 173-192.
APPENDIX. Plant species recorded in the diets of birds captured in mist nets.
APPENDIX. Continued.