Blake FruitsDietsNeotropical 1992

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Fruits in the Diets of Neotropical Migrant Birds in Costa Rica

Author(s): John G. Blake and Bette A. Loiselle


Source: Biotropica , Jun., 1992, Vol. 24, No. 2, Part A (Jun., 1992), pp. 200-210
Published by: Association for Tropical Biology and Conservation

Stable URL: https://www.jstor.org/stable/2388674

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BIOTROPICA 24(2a): 200-210 1992

Fruits in the Diets of Neotropical Migrant Birds in Costa Rica1

John G. Blake and Bette A. Loiselle

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

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Diets of Neotropical Migrants 201

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

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202 Blake and Loiselle

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

Yellow-bellied Flycatcher EMFL 4 1 1 (100) 0 (0)


Empidonax flaviventris
Acadian Flycatcher EMVI 40 23 23 (100) 6 (26) 1.0 0.0 1
E. virescens
"Traill's" Flycatcher EMTR 24 20 17 (85) 14 (70) 1.0 0.0 1
Empidonax sp.a
Great Crested Flycatcher MYCR 10 10 10 (100) 6 (60) 1.17 0.41 2
Myiarchus crinitus
Eastern Kingbird TYTY 9 9 1 (11) 9 (100 1.56 0.88 3
Tyrannus tyrannus
Veery CFUS 1 1 1(100) 1(100) 2
Catharus fuscescens
Gray-cheeked Thrush CAMI 24 23 20 (87) 22 (96) 2.0 0.98 4
C. minimus
Swainson's Thrush CAUS 206 199 105 (53) 196 (98) 1.41 0.70 4
C. ustulatus
Wood Thrush HYMU 303 259 240 (93) 154 (59) 1.29 0.62 4
Hylocicbla mustelina
Gray Catbird DUCA 47 43 27 (63) 41 (95) 1.34 0.53 3
Dumetella carolinensis
Red-eyed Vireo VIOL 2 2 1 (50) 2 (100) 1.0 0.0 1
Vireo olivaceus
Blue-winged Warbler VEPI 3 3 3 (100) 1(33) 1
Vermivora pinus
Golden-cheeked Warbler VECH 4 4 4 (100) 0 (0)
V. cbrysoptera
Chestnut-sided Warbler DEPE 45 44 44 (100) 8 (18) 1.0 0.0 1
Dendroica pensylvanica
Magnolia Warbler DEMA 1 1 1 (100) 0 (0)
D. magnolia
Blackburnian Warbler DFUS 2 2 2 (100) 1 (50) 1
D. fuscus
Bay-breasted Warbler DECA 3 3 3 (100) 2 (67) 1.0 0.0 1
D. castanea
Black-and-white Warbler MNVA 3 3 3 (100) 0 (0)
Mniotilta varia
Worm-eating Warbler HEVE 19 14 14 (100) 3 (21) 1.0 0.0 1
Helmitheros vermivorus
Ovenbird SEAU 113 81 81 (100) 18 (22) 1.17 0.38 2
Seiurus aurocapillus
Northern Waterthrush SENO 6 2 2 (100) 0 (0)
S. noveboracensis
Louisiana Waterthrush SEMO 5 4 4 (100) 0 (0)
S. motacilla
Kentucky Warbler OPFO 109 86 86 (100) 10 (12) 1.0 0.0 1
Oporornis formosus
Mourning Warbler OPPH 11 9 9 (100) 0 (0)
0. philadelphia
Hooded Warbler WICI 4 1 1 (100) 0 (0)
Wilsonia citrina
Canada Warbler I 6 6 6 (100) 0 (0)
W. canadensis
Summer Tanager PIRU 8 7 6 (86) 4 (57) 1.25 0.50 2
Piranga rubra
Scarlet Tanager PIOL 2 2 2 (100) 2 (100) 1.5 0.71 2
Piranga olivacea

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Diets of Neotropical Migrants 203

TABLE 1. Continued.

Total

Total fecal Samples with Plant species


Bird cap- sam-
Bird species code tures ples Insects Fruits Mean SD Max

Rose-breasted Grosbeak PHLU 1 1 1 (100) 1 (100) 1


Pheucticus ludovicianus
Northern Oriole ICGA 1 1 1 (100) 1 (100) 1
Icterus galbula

a Mostly Empidonax traillii.

grants at La Selva subcrustulata


bird species included Conostegia depend primarily on insects for
(eaten
foodaffinis
by 16 species), Miconia but about 20 percent
(13 feed primarily
species), or exten- and
Clidemia dentata (10 species). All three species are sively on fruit (Blake et al. 1990). This compares
in the Melastomataceae and produce small, juicy to approximately 30 percent of the total avifauna
berries. at La Selva (363 total species, excluding accidentals)
Overall, a higher proportion of samples from that rely primarily on fruit. Two-thirds of the 30
forest habitats contained fruit (Table 3), probably migrant species captured during this study ate some
because many more generally nonfrugivorous species fruit and fruit was represented in over 50 percent
(e.g., Ovenbird, Kentucky Warbler) were captured of all fecal samples collected.
in second-growth habitats (see Blake & Loiselle, in The importance of fruit in the diets of migrants
press). Within individual migrant species, several varies widely among species and is influenced by a
(Table 3) were more frugivorous in the young sec- variety of factors, including morphology and for-
ond-growth site, where fruit was more abundant
(see also Blake & Loiselle 1991).
Seasonal variation in frugivory (based on num-
100 ALL SPECIES
ber of fecal samples containing fruit) was apparent
(Fig. 1). When data from all species and sites were H-80
pooled, an increase in frugivory from early to late
of 60
dry season was apparent (X2 > 14, P < 0.001, all
years; chi-square based on number of samples with T 40
and without fruit). The increase in fruit use late in
the dry season corresponded to the period when
many highly frugivorous migrants, particularly
Swainson's Thrush, pass through La Selva on their
way north. Other migrants, including winter resi- <E10 WOOD THRUSH
dents, also may increase consumption of fruit prior
to migration. The Wood Thrush, a common winter 20

resident at La Selva, showed a statistically significant


0~
increase in frugivory from early to late dry season
during 1987 and 1988 (Fig. 1; 1985: x2 = 0.65,
P > 0.50; 1986: x2 = 3.4, P < 0.20; 1987 and
1988: x2 > 16, P < 0.001).

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-

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204 Blake and Loiselle

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Diets of Neotropical Migrants 205

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206 Blake and Loiselle

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.

All species CAMI CAUS HYMU

Habitat N %I %F N %I %F N %I %F N %I %F

Young second-growth 475 81 57 16 88 94 100 49 99 62 89 65


Old second-growth 123 83 53 3 100 100 39 54 97 21 90 67

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

EMVI DEPE SEAU OPFO

Habitat N %I %F N %I %F N %I %F N %I %F

Young second-growth 13 100 38 32 100 22 72 100 24 54 100 13


Old second-growth 4 100 0 12 100 8 8 100 0 23 100 13
Primary forest (all) 6 100 17 9 100 0

aging behavior (Martin


ole), and 2 (Solitary Vireo1985a, Moerm
Vireo solitarius) species
low 1985). Gape width of the bird affects the size of fruits. Number of samples (fecal samples, ob-
range of fruits that birds consume (Wheelwright servations) collected from each species was not pre-
1985) and may influence patterns of habitat selec- sented by Wheelwright et al. (1990), so direct com-
tion (Martin 1985b). Most warblers encountered parisons by species with the present study are not
during this study have relatively small gapes and possible. Keeping this in mind, we recorded a similar
were primarily insectivorous. Many eat fruit infre- number of fruits in the diet of Summer Tanagers
quently and are perhaps most likely to opportunisti- at La Selva (5 species; 6 samples) but found many
cally consume small, juicy berries of the Melasto- more species of fruits (63) in the diet of Swainson's
mataceae (e.g., Conostegia subcrustulata and Miconia Thrush; 52 species were recorded from Wood
affinis eaten by Kentucky Warbler and Ovenbird). Thrushes. The species with the greatest diversity of
In contrast, migrant thrushes, mimids, and some fruits in its diet at Monteverde (95 species) was the
flycatchers have larger gapes than warblers and con- Emerald Toucanet (Aulacorhynchusprasinus), a per-
sumed substantial amounts of fruit, often of many manent resident. Our data thus demonstrate that
different species. some migrants may rival permanent residents in the
Fruit handling technique also affects the number number of fruits consumed. The diversity of fruits
and types of fruits consumed. Mimids and thrushes in the diets of some migrants suggests that they,
commonly swallow fruits whole ("gulpers" of like many resident frugivores, may disperse the seeds
Moermond & Denslow 1985); fruit selection is like- of many plant species. The relative importance of
ly to be constrained by gape size. Birds that man- migrants as seed dispersers will also depend on the
dibulate fruits in their bills ("mashers"; e.g., Piran- probability that seeds are disseminated to suitable
ga tanagers), may be less constrained. Although we sites for survival and growth (Greenberg 1981).
obtained relatively few samples from such migrants Few data are currently available to assess this point.
(see Table 2), those samples we did obtain suggest Although many species may eat fruit through-
that these birds also feed on a wide variety of fruits. out the nonbreeding season (e.g., Eastern Kingbird;
Many of the species that consumed fruit at La Morton 1971), temporal shifts in degree of frugivo-
Selva have been reported to consume fruit elsewhere ry also occur (Greenberg 1981). Wood Thrushes
in the tropics (Leck 1972b, Morton 1980, Green- are more frugivorous during the late dry season
berg 1981, Martin 1985b). Wheelwright et al. (spring) than earlier (winter). A shift to greater
(1984) presented a survey of fruit use by 70 species frugivory by migrants during the later dry season
of birds at Monteverde, Costa Rica, a mid-elevation in Panama has been noted by previous authors (Leck
site (1350-1550 m). Four species of neotropical 1972b, Morton 1980, Martin & Karr 1986). Such
migrants were reported to consume 13 (Swainson'sincreases in frugivory likely relate to increased need
Thrush), 4 (Summer Tanager), 4 (Northern Ori- to accumulate fat for migration (Martin 1985b,

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Diets of Neotropical Migrants 207

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.

loss of tropical forests has received much attention

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lowland habitats in Costa Rica. Auk 108: 114-130.
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Hagan and D. W. Johnston (Eds.). Ecology and conservation of neotropical landbird migrants. Smithsonian
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, F. G. STILES, AND B. A. LOISELLE. 1990. Birds of La Selva: habitat use, trophic composition, and migrants.
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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.
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Diets of Neotropical Migrants 209

APPENDIX. Plant species recorded in the diets of birds captured in mist nets.

Family/Species Code Family/Species Code

Araceae M. barbinervis MICBAR


Anthurium sp. ANTHUR M. brenesii MICBRE
Dieffenbachia sp. DIEFFE M. centrodesma MICCEN
Haemodoraceae M. graci/is MICGRA
M. multispicata MICMUL
Xiphidium coeruleum XIPCOE M. nervosa MICNER
Heliconiaceae M. serrulata MICSER
Heliconia wagneriana HELWAG M. simplex MICSIM
He/iconia wagneriana HELWAG M. aff. smaragdina MICSMA

Smilacaceae M. sp. MICSPP


Smilax sp. SMILAX M. Sp. MICI 10
Araliaceae M. sp. MIC036
Dendropanax arboreus DENARB M. sp. MIC280
Schefflera systyla SCHSYS Ossaea macrophylla OSSMAC
Cecropiaceae 0. robusta OSSROB
Cecroplaceae Ossaea sp. ~~~~~~~OSSSPP
Cecropia insignis CECINS Ossaea sp. TOP175
C. obtusifolia CECOBT Topobea sp. TEL075
Compositae Melastomataceae sp. MEL073
Melastomataceae sp. MELI 18
Clibadium asperum CLIASP Melastomataceae sp. MEL108
Dilleniaceae Melastomataceae sp. MEL103
Doliocarpus dentatus DOLDEN Menispermaceae
Pinzona coriacea INCissamPeos sp.
Dilleniaceae sp. DILLEN Monimiaceae

Ericaceae Siparuna tonduziana SIPTON


Ericaceae sp. ERI2 52 Moraceae

Euphorbiaceae Ficus sp. FICCOL


Alchornea costaricensis ALCCOS Myrsinaceae

Gesneriaceae Ardisia auriculata AURAUR


Besleria columneoides BESCOL A. nigropunctata AURNIG
Drymonia pilifera DRYPIL Nyctaginaceae
Guttiferae Neea psychotrioides NEEPSY
Clusia palmana CLUPAL Passifloraceae
Clusia sp. CLU 113
C/usiacea sp. CLU13Passiflora auriculata PASAUR
Lauraceae
Nectandra membranacea NECMEM Phytolaccaceae
Phytolacca rivinoides PHYRIV
Malvaceae Trichostigma polyandrum TRIPOL
Hampea appendicu/ata HAMAPP Piperaceae

Marcgraviaceae Piper sp. PIP007


Marcgravia affinis MARAFF Rubiaceae
Melastomataceae Bertiera guianensis BERGUI
Clidemia crenulata CLICRE Cephaelis elata CEPELA
C. densiflora CLIDNS Hamelia patens HAMPAT
C. dentata CLIDEN Psychotria brachiata PSYBRA
C epiphytica CLIEPI P. copensis PSYCOP
C. disco/or CLIDIS P. grandis PSYGRA
Conostegia bracteata CONBRA P. grandistipulata PSYGST
C. subcrustulata CONSUB P. orchidearum PSYORC
Conostegia sp. CON162 P. officinalis PSYOFF
Henriettea tuberculosa HENTUB P. pittieri PSYPIT
Leandra sp. LEANDR P. psychotriifolia PSYPSY
Miconia affinis MICAFF Psychotria sp. PSY099
M. appendiculata MICAPP Sabicea villosa SABVIL
M. "attenuate' MICATT Sabicea sp. SABO6O

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210 Blake and Loiselle

APPENDIX. Continued.

Family/Species Code Family/Species Code

Sabicea sp. SABO94 Vitaceae


Rubiaceae sp. RUB390 Cissus pseudosicyoides CISPSE
Solanaceae Unknown
Cestrum racemosum CESRAC Unknown sp. UNK090
Lycianthes sp. LYCO84 Unknown sp. UNK106
Witheringia asterotricha WITAST Unknown sp. UNK046
Urticaceae Unknown sp. UNK041
Urera baccifera UREBAC Unknown sp. UN0855
U. caracasana URECAR Unknown sp. UN1641
Unknown sp. UN2441
Verbenaceae Orange pulp ORPULP
Vitex cooperi VITCOO Juice, pulp (not orange) PULP
Verbenaceae sp.? VER009

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