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Integrative Zoology 2013; 8: 57?62 doi: 10.1111/j.1749-4877.2012.00313.x
ORIGINAL ARTICLE
Viability of small seeds found in feces of the Central American 
tapir on Barro Colorado Island, Panama
Paula I. CAPECE1,  Enzo ALIAGA-ROSSEL1,2 and Patrick A. JANSEN3,4
1Ecology, Evolution and Conservation Biology Program, University of Hawaii at Manoa, Honolulu, HI, USA, 2Institute of         
Ecology, Higher University of San Andres, La Paz, Bolivia, 3Smithsonian Tropical Research Institute, Anc?n, Panama City, 
Panama and 4Center for Ecosystem Studies, Wageningen University, Wageningen, Netherlands
   
Abstract 
Tapirs are known as effective dispersers of large-seeded tree species, but their role in dispersing small-seeded 
plant species has yet to be established. Tapir feces have been reported to contain large numbers of small seeds, 
but whether these are viable has rarely been evaluated. We determined the abundance and viability of small 
seeds in feces of Central American tapir (Tapirus bairdii) on Barro Colorado Island, Panama. A total of 72 fe-
cal samples were collected opportunistically from 4 tapir latrine sites. Seeds were manually extracted from fe-
ces and classified by size. Seed viability was estimated by opening each seed and examining for the presence of 
at least 1 intact firm white endosperm. In total, we obtained 8166 seeds of at least 16 plant species. Small-seed-
ed species dominated, with 96% of all seeds found measuring <5 mm. The canopy tree Laetia procera was the 
most abundant species in the samples. Of all small seeds found, 69% contained an intact endosperm and ap-
peared viable. This suggests that small seeds, like large seeds, often pass through the digestive tract of T. bairdii 
intact. Thus, tapirs potentially serve as effective dispersers of a wide range of small-seeded plant species. 
Key words: Central America, megafauna, seed dispersal, seed size, tapir
Correspondence: Paula I. Capece, National Park Service, 1978 
Island Ford Parkway, Sandy Springs, GA 30350, USA. 
Email: paula_capece@nps.gov
INTRODUCTION
Seed dispersal by tapirs (Tapirus spp.) is an impor-
tant determinant of the spatial distribution of plants in 
Neotropical forests (Forget & Sabatier 1997; Fragoso et 
al. 2003). Tapirs are the largest extant terrestrial mam-
mals in Neotropical forests, are able to swallow large 
seeds and can ingest a large quantity of seeds while 
feeding (Terwilliger 1978; Naranjo 1995). Tapirs pos-
sess a ?primitive? digestive tract characterized by incom-
plete digestion; therefore, many seeds can pass through 
the tapir gut intact (Janzen 1984; Bodmer 1991). Long 
seed retention times in the gut (Janzen 1981) and large 
home-range sizes (Foerster & Vaughan 2002) contribute 
to the tapir?s ability to disperse seeds over long distanc-
es, in some cases to suitable sites beyond the range of 
seed-consuming larvae (Fragoso et al. 2003). 
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Tapirs are considered effective dispersers of plant 
species with large, thick-walled seeds; that is, they tend 
to deposit seeds unharmed to suitable germination sites 
away from the parent plant (see Schupp 1993 and Jorda-
no & Schupp 2000 for a detailed discussion of effective 
dispersal). For example, Tapirus terrestris (Linnaeus, 
1758) is an effective disperser of large-seeded palms 
(Rodrigues et al. 1993; Fragoso & Huffman 2000; 
Quiroga-Castro & Roldan 2001; Tobler et al. 2010) and 
Tapirus bairdii (Gill, 1865) is an effective disperser of 
large-seeded tree species, including Caesalpinia coriar-
ia (Janzen 1982), Guazuma ulmifolia (Janzen 1982) and 
Manilkara zapota (O?Farrill et al. 2006, 2012). How-
ever, tapirs also feed on fruits that contain many small 
seeds and are known to ingest small seeds while brows-
ing (Janzen 1984). Fragoso and Huffman (2000) report 
that seeds of 4 small-seeded species germinated after 
they were extracted from T. terrestris feces. Small seeds 
have been found in T. terrestris stomach contents (Hen-
ry et al. 2000) and feces (T?foli 2006; Zorzi 2009; To-
bler et al. 2010), as well as in T. bairdii feces (Naran-
jo 1995). Whether tapirs are effective dispersers of these 
small seeds is partially determined by the degree to 
which small seeds survive passage through the digestive 
tract intact, which is largely unknown. 
We examined the potential of the Central American 
tapir (T. bairdii) as a disperser of small-seeded plant 
species by estimating the viability of seeds found in ta-
pir feces on Barro Colorado Island, Panama. We also re-
corded the abundance and diversity of small seeds in fe-
ces. 
MATERIALS AND METHODS
Study site and species
Barro Colorado Island (BCI), Panama (9?9?N, 
79?51?W) is a 16 km2 island covered with tropical low-
land moist forest. BCI has an average temperature of 
27 ?C, an average yearly rainfall of 2600 mm, a rainy 
season from May through December and a marked dry 
season from January through April. The island has a het-
erogeneous landscape with numerous creeks, most of 
which are seasonally dry (Leigh et al. 1996). 
Tapirus bairdii is 1 of 4 species of the family Tapir-
idae of the order Perissodactyla and the only remain-
ing Pleistocene megafauna in Central America (Jan-
zen 1982). It is distributed intermittently from southern 
Mexico to northwestern Colombia in humid habitats 
from sea level to 3600 m (IUCN 2012). The diet of T. 
bairdii consists of a variety of plant species and plant 
parts, including leaves, stems and fruits (Terwilliger 
1978; Naranjo 1995; Olmos 1997; Tobler 2002). T. bair-
dii is more active during nocturnal hours (Foerster & 
Vaughen 2002). It is known to use latrines or sites of re-
peated defecation (Flesher 1999; Naranjo 2009).
The diet of T. bairdii on BCI has been studied by Ter-
williger (1978) through direct observation of the feed-
ing behavior of individual tapirs. To our knowledge, no 
studies have examined fecal samples on BCI. Estimates 
of the tapir population on BCI have varied greatly over 
time (Enders 1935; Enders 1939; Eisenberg & Thor-
ington 1973; Terwilliger 1978; Eisenberg 1980; Glanz 
1996). More recent estimates of T. bairdii density on 
BCI obtained from diurnal and nocturnal census efforts 
are 0.19 and 0.36 individuals/km2, respectively (Wright 
et al. 1994). The current population size of tapirs on 
BCI is unknown (R. W. Kays, pers. comm.). 
Sampling
We collected fresh fecal samples (dung boluses 
<1 week old as judged by inspection of odor, color and 
integrity of fecal material) at 4 tapir latrine locations 
along the northern shore of BCI, near the terminal por-
tions of the ?Standley? and ?Miller? trails, during May?Aug 
2003. All 4 latrines were located within a straight-line 
distance of approximately 1.5 km (approximately 3 km 
ground distance), with 2 of the latrines in close proxim-
ity (approximately 50 m). Fecal samples were slowly 
dried in a dry closet for 3 days at approximately 35 ?C 
and then spread out on trays in a dark air-conditioned 
laboratory for an additional 7 days until completely dry. 
Each fecal sample was packaged individually in a pa-
per bag and assigned a number for identification, then 
stored in plastic with desiccant until processing, which 
was completed within 8 weeks. We extracted the seeds 
from dry feces manually. This proved more efficient 
than seed extraction using different-sized wire mesh 
screens (as in Naranjo 1995; Fragoso & Huffman 2000), 
because many small seeds passed directly through 5 mm2 
mesh sieves and could not be effectively separated from 
browse material with smaller-mesh sieves. We classi-
fied seeds as small (S < 5 mm), medium (M = 5?20 mm) or 
large (L > 20 mm) and identified seeds to ?morphospe-
cies?. We attempted to identify morphospecies to spe-
cies through comparison with BCI herbarium samples 
and with the assistance of Osvaldo Calder?n and An-
dres Hernandez (BCI botanists). Viability was estimat-
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Small seeds in Tapirus bairdii feces
? 2012 Wiley Publishing Asia Pty Ltd, ISZS and IOZ/CAS
ed by the cut test (Terry et al. 2003); opening each un-
imbibed seed and examining under magnification for 
the presence of at least 1 intact firm white endosperm 
(as in Fragoso & Huffman 2000). Although the cut test 
method is indirect and may vary by species, it has been 
used successfully to determine viability for tapir ingest-
ed seeds (Fragoso & Huffman 2000), for small seeds 
dispersed by frugivorous tortoises (Strong & Fragoso 
2006) and for other small seeds (Piper spp.) where ger-
mination tests failed to distinguish between viability and 
dormancy (Daws et al. 2002) . 
RESULTS
Of 72 fecal samples obtained, 52 from 3 tapir latrine 
locations in the early wet season comprised approxi-
mately 3900 cm3 of fecal matter containing 1084 seeds 
(Table 1). A total of 20 samples from a fourth latrine lo-
cation during the mid-wet season comprised approxi-
mately 1800 cm3 of fecal matter containing 7082 seeds. 
The fecal collections from the 3 early wet season loca-
tions were deposited by tapirs in moist areas that were 
not flooded at the time of collection. Thirteen of the 20 
mid-wet season samples were intact boluses collected 
from a flooded drainage and 7 were collected immedi-
ately adjacent to this drainage on dry soil. 
In total, the feces contained 8166 seeds of at least 
16 plant species. The size range of seeds encountered 
was approximately 1?30 mm in length, with 7864 small 
seeds, 193 medium seeds and 109 large seeds. The ma-
jority of the seeds (96%) and species (50%) were small-
seeded. As much as 83% of all seeds came from the can-
opy tree Laetia procera (Flacourtiaceae); these 2?3 mm 
sized seeds were present in each of the 20 fecal samples 
collected in August. The next most abundant (group of) 
species was Ficus, which made up 11% of the seeds. Of 
all small seeds (<5 mm), the majority (69%) contained 
an intact endosperm. 
DISCUSSION
Tapirs are important dispersers of large-seeded tree 
species (Bodmer 1991; Fragoso 1997; Quiroga-Castro & 
Roldan 2001; Fragoso et al. 2003; Giombini et al. 2009; 
O?Farrill et al. 2012). Previous studies reported that T. 
terrestris feces contain large numbers of small seeds (e.g. 
Fragoso & Huffman 2000; T?foli 2006; Talamoni & As-
sis 2009; Zorzi 2009; Tobler et al. 2010). Our study pro-
vides similar data for T. bairdii and suggests that a large 
proportion of small seeds that pass through the digestive 
tract of T. bairdii are viable. Feces of the Central Ameri-
can tapir on Barro Colorado Island contained large num-
bers of small seeds, which by far outnumbered large 
seeds in the samples. The majority of these appeared vi-
able in cut tests. 
 The most abundant seed in the fecal samples we col-
lected was L. procera, an uncommon species in the old-
growth forest of BCI, which produces 15?20 mm large 
berrylike capsules containing 10?20 seeds, in Aug?Sep 
(Croat 1978). These berrylike fruits are rich in crude 
fat and energy content (Castellanos & Chanin 1996) 
and are known to be eaten by a variety of birds and pri-
mates that collect the fruits directly from the trees (Snow 
1971; Van Roosmalen et al. 1988; Castellanos & Chanin 
1996; Wehncke et al. 2003). The tapirs producing the 
feces of this study must have fed on L. procera fruits 
that had fallen to the forest floor and deposited the seeds 
at latrines 79% intact. 
As our study covered just 4 different latrines and pos-
sibly few individual tapirs, the abundances of various 
plant species we recorded in feces might not be repre-
sentative of an unbiased sample. For example, L. pro-
cera appears to be overrepresented. This study did not 
address the relationship between abundance of seeds 
found in feces and the availability of the same species at 
the time of fecal sample collection. Year-round sampling 
in combination with large-scale inventory and monitor-
ing of tree fruiting phenology would allow us to assess 
whether tapirs feed selectively, or rather on whatever is 
available.
Our results suggest that tapirs might act as dispers-
ers not only of large-seeded plant species, as document-
ed by previous studies (Bodmer 1991; Rodrigues et al. 
1993; Fragoso 1997; Fragoso & Huffman 2000; Quiro-
ga-Castro & Roldan 2001; Giombini et al. 2009; To-
bler et al. 2010; O?Farrill et al. 2012), but also of plant 
species with relatively small seeds that are not present-
ly considered ?tapir-dispersed?. To assess this role with 
more certainty, it would be worthwhile investigating the 
occurrence and viability rates of small seeds (<5 mm) 
in tapir dung at other locations and in all seasons. In ad-
dition, although cut tests provide acceptable estimates 
of viability (Terry et al. 2003; as in Fragoso & Huffman 
2000, Daws et al. 2002 and Strong & Fragoso 2006), 
germination tests are recommended to determine viabil-
ity with greater certainty.
Ultimately, the effectiveness of tapirs as dispers-
ers of small seeds will also depend on the fate of the vi-
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T
ab
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 1
 S
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it
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(A
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(p
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M
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.7
 (
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? S
ee
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s 
ar
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ap
pr
ox
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at
e.
 S
iz
e 
cl
as
s 
(l
en
gt
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: 
S
 <
 5
 m
m
, M
 =
 5
?2
0 
m
m
, L
 >
 2
0 
m
m
. 
? F
re
sh
 m
as
s 
of
 L
ae
ti
a 
pr
oc
er
a 
ob
ta
in
ed
 f
ro
m
 G
ua
ri
gu
at
a 
(2
00
0)
; 
al
l 
ot
he
r 
fr
es
h 
m
as
s 
re
co
rd
s 
pr
ov
id
ed
 b
y 
J.
 W
ri
gh
t (
pe
rs
. c
om
m
.)
. ?
F
re
qu
en
cy
 =
 p
er
ce
nt
ag
e 
of
 to
ta
l f
ec
al
 s
am
pl
es
 th
at
 c
on
ta
in
ed
 th
e 
sp
ec
ie
s.
 ? M
ea
n 
es
ti
m
at
ed
 v
ia
bi
li
ty
 (
%
) 
ac
ro
ss
 s
pe
ci
es
. 
N
A
, n
ot
 a
va
il
ab
le
.
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? 2012 Wiley Publishing Asia Pty Ltd, ISZS and IOZ/CAS
able seeds in the feces in relation to the habitat. Ta-
pirs are known to deposit feces in both dry and flooded 
sites (Fragoso & Huffman 2000; Tobler et al. 2010). 
For flooded sites, tapir-mediated dispersal might be es-
pecially effective for plant species associated with wet 
habitats (Salas & Fuller 1996), to which tapirs may pro-
vide directed dispersal (Wenny 2001). Dispersal by ta-
pirs might also be effective for other small-seeded plant 
species when combined with secondary seed dispersal 
by dung beetles that move and bury dung in the soil (e.g. 
seeds in howler monkey feces [Estrada & Coates-Estra-
da 1991]). 
ACKNOWLEDGMENTS 
We thank the Smithsonian Tropical Research Institute 
for hosting this study; Jim Dalling for use of his labora-
tory; Osvaldo Calder?n and Andres Hernandez for seed 
identification; Eloisa Lasso De Paulis for assistance in 
the field; Ricardo Moreno for assistance in the field and 
help with herbarium seed comparison; Jos? M. V. Frago-
so and Andrew Taylor for advice; Joe Wright for seed 
mass data; Roland Kays for current BCI tapir population 
information; and Jos? M. V. Fragoso, Aaron Shiels, Lau-
ra Capece, Michael Szperka for comments on previous 
versions of the manuscript. Patrick A. Jansen acknowl-
edges financial support from the Netherlands Founda-
tion for Scientific Research. 
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