American Society of Mammalogists

Foraging and Breeding in Two Sympatric Species of Neotropical Bats, Genus Noctilio
Author(s): Emmet T. Hooper and James H. Brown
Source: Journal of Mammalogy, Vol. 49, No. 2 (May, 1968), pp. 310-312
Published by: American Society of Mammalogists
Stable URL: http://www.jstor.org/stable/1377989 .
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 GENERAL NOTES
FORAGING AND BREEDING IN TWO SYMPATRICSPECIES OF NEOTROPICAL
BATS, GENUS NOCTILIO
It is axiomatic that closely related species must have different requirementsfor limiting
environmental resources in order to avoid competing and thus to persist in the same area.
Despite this belief, there are few quantitative ecological data on sympatric pairs of tropical
mammal species. This paper presents a few data on foraging activities, diets, and re-
production of two sympatric species of Neotropical bats, Noctilio leporinus and N. labialis.
The ranges of the species overlap broadly in tropical America and both forms probably
frequently occur at the same locality. Certainly they do on the Rio Chagres in Panama
(Bloedel, 1955) and on the Rio Higueron, one-half mile east of Finca Jimenez, Guanacaste
Province, Costa Rica, where we obtained both species on the nights of 13-15 February
1966 in a program of mist netting that was part of a field course offered by the Organiza-
tion for Tropical Studies.
Two nets, each 12 m long, were set at different heights (0.8 to 3.0 m and 3.6 to 5.8 m
above the surface of the water) directly across the Rio Higuer6n. The nets were tended
for two successive nights; bats were removed at 2-hr intervals in the periods 7:00 PM to
6:00 AM. At this time of year, the dry season, vertebrates of all kinds probably were con-
centrated along the river where there was more cover and moisture. The riparian forests
comprised a band of green vegetation in the otherwise almost leafless, tropical deciduous
forest.
Capture levels in the nets indicate flight levels of the bats. The data show that the two
species fly at the same heights; all 10 leporinus and 32 of the 34 labialis were captured in
the lower net. This is in harmony with Bloedel's observations (1955) of flight levels of
the species in Panama. Much of the flight time probably is foraging time.
The relative numbers caught in each of the 2-hr periods suggest temporal patterning of
activity (Table 1). A Kolmogorov-Smirnovtest indicates that the flying times of the two
species are significantly different (0.05 > P < 0.01). When and where we observed them,
N. leporinus was active throughout the night, whereas activity of N. labialis was concen-
trated immediately after sundown. Similar peaks in activity soon after sunset have been
observed in temperate vespertilionid bats (Jones, 1965; Cockrum and Cross, 1964). From
the data presented, it appears that the two species of Noctilio forage in the same areas,
but for the most part utilize those areas at different times.
Stomach contents indicate diets of the species. The possible time lapse between capture
and preservationof the bats in formalin varied from approximately15 min to 3 hr. Results
of analyses of the stomachs appear in Table 2, and we gratefully acknowledge the assistance
of R. M. Bailey and T. H. Hubbell in identifying fish and insects in the samples. N. labialis
contained only insect remains. Because these were finely chewed and partly digested, it
is possible that insects with hard, relatively indigestible cuticles (e.g., beetles) constituted
a greater proportion of the reported stomach contents than of the actual diet of the bats.
The identifiable insects ranged from approximately4 to 20 mm in body length. The fullest
stomachs contained representativesof as many as six insect orders, and at least six species
of Coleoptera alone were found in several stomachs. All the insects identified were capable
of flight and could have been captured in the air; there is no good reason to infer that
any were taken from the water. The specimens of leporinus contained remains of both in-
sects and fish, but to judge from this small sample, fish make up a large part of the diet
at this locality. Because leporinus is larger than labialis (an adult of each weighs approxi-
mately 55 g and 30 g, respectively), its food probably also is larger.
These diets resemble those reported for the two species at other localities. Dunn (1934)
and Bloedel (1955) also found nothing but insect remains in the guano and stomachs of
labialis. In stomachs of leporinusthere were both fish and insects (Bloedel, 1955; Goodwin,
310

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May 1968 GENERAL NOTES 311

TABLE 1.-Numbers of Noctilio caught the nights of February 13-14 (N1 below) and
14-15 (N2 below).
N. leporinus N. labialis
Period N1 N2 Totals N1 N2 Totals

Sunset to 7 PM 0 1 1 13 9 22
7-9 PM 2 0 2 3 1 4
9-11 PM 1 1 2 4 1 5
11 PM-1 AM 0 1 1 0 0 0
1-3 AM 2 2 4 0 3 3
3-5 AM 0 0 0 0 0 0

5 5 10 20 14 34

1928; Goodwin and Greenhall, 1961). Methods employed by leporinus in locating and
capturing fish have been described in detail by Bloedel (1955) and Suthers (1965).
Reproductive tracts provided information on breeding season in the two species. Of 15
female labialis, 13 were pregnant. Each contained a single embryo 3 to 5 mm in length
(12 specimens) or smaller (1 specimen); the size is far short of that at term. Each of
three female leporinus also contained a single embryo; these measured 9, 33, and 43 mm.
The largest of these is obviously near term and the one 33 mm in length is at a more
advanced stage than any of the embryos of labialis.
These data suggest that labialis has a well defined breeding season and that its young
are born later in the year than those of leporinus. Pregnant leporinus have been collected
in Trinidad in February, lactating females were taken in February and March, and a single
parturition was observed in December (Goodwin and Greenhall, 1961). In Panama, N.
labialis mate in late November or December and give birth in late April or early May
(Anderson and Wimsatt, 1963). These authors also found that all embryos from a given
area were at about the same stage of development, and suggested that all the pregnant
females within a population must have bred within 1 or 2 weeks of each other. Wimsatt
(1945) described similar synchronous pregnancies and parturitions in colonies of the tem-
perate vespertilionid, Myotis lucifugus. Synchrony of breeding activities in these popula-
tions prompts one to look for environmental stimuli responsible for initiating reproductive
activity in tropical and temperate bat populations.
Significant differences between closely related sympatric species in the utilization of an
environmental resource suggest, but do not prove, that resource limits the population sizes

TABLE 2.-Stomach contents of samples of Noctilio.

Number of stomachs
Orders
represented N. leporinus N. labialis

Coleoptera 0 17
Lepidoptera 0 7
Hemiptera 1 6
Hymenoptera 0 5
Diptera 0 5
Orthoptera 0 3
Homoptera 0 3
Unidentified 2 25
Teleostei 2 0

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 312 JOURNAL OF MAMMALOGY Vol. 49, No. 2

of those species; the differences in resource utilization represent the evolution of mecha-
nisms reducing competition and permitting coexistence. Lack (1966) argued strongly
that the population sizes of birds which feed on insects or vertebrates are limited by the
availability of food. The several characteristics (activity time, diet, size of food, and re-
productive period) in which these species of Noctilio differ may represent evolutionary
developments which have been effective in reducing competition for food between the
two forms. Assuming that the two species are derived from an insectivorous ancestor, the
piscivorous habit in N. leporinus may have evolved as a result of competition for insect
food and may play an importantrole in the continued coexistence of the two species. Dif-
ferent reproductive periods probably place maximum demands on shared prey items (in-
sects) at different seasons. Differences in activity patterns may simply indicate adaptations
to the different diets, or they may represent a mechanism of further reducing interactions
by temporal segregation of foraging activities.
The National Science Foundation's support of the Organizationfor Tropical Studies and
of the University of Michigan's training program in systematic and evolutionary biology
made this study possible.
LITERATURE CITED
ANDERSON, J. W., AND W. A. WIMSATT. 1963. Placentation and fetal membranes of the
Central American noctilionid bat, Noctilio labialis minor. Amer. J. Anat., 112:
181-201.
BLOEDEL, P. 1955. Hunting methods of fish-eating bats, particularly Noctilio leporinus.
J. Mamm., 36: 390-399.
COCKRUM, E. L., AND S. P. CROSS. 1964. Time of bat activity over water holes. J. Mamm.,
45: 635-636.
DUNN, L. H. 1934. Notes on the little bulldog bat, Dirias albiventor minor (Osgood),
in Panama. J. Mamm., 15: 89-99.
GOODWIN,G. G. 1928. Observationson Noctilio. J. Mamm., 9: 104-113.
GOODWIN,G. G., AND A. M. GREENHALL. 1961. A review of the bats of Trinidad and
Tobago. Bull. Amer. Mus. Nat. Hist., 122: 187-302.
JONES,C. 1965. Ecological distribution and activity periods of bats of the Mogollon
Mountains area of New Mexico and adjacent Arizona. Tulane Studies Zool.,
12: 93-100.
LACK,D. 1966. Population studies of birds. Clarendon Press, Oxford, vi + 341 pp.
SUTHERS,R. A. 1965. Acoustic orientation by fish-catching bats. J. Exp. Zool., 158:
319-348.
WIMSATT, W. A. 1945. Notes on breeding behavior, pregnancy, and parturitionin some
vespertilionid bats of the eastern United States. J. Mamm., 26: 23-33.
EMMET T. HOOPERAND JAMES H. BROWN,Museum of Zoology, University of Michigan,
Ann Arbor,48104, and Department of Zoology, Universityof California,Los Angeles, 90024.
Accepted 8 January 1968.

CHROMOSOMESOF THE GROUND SQUIRREL, SPERMOPHILUS RICHARDSONII

AUREUS (DAVIS)
Controversy exists regarding the taxonomic status of the ground squirrels comprising
Spermophilus richardsonii (Hall and Kelson, 1959). Howell (1938) classified the sub-
species elegans, nevadensis, and richardsoniias members of the same species, S. richardsonii,
because there was gross morphological evidence of intergradation between elegans and
richardsonii in Gallatin County, Montana. In contrast, Davis (1939) did not find inter-
gradation between the ground squirrels of southeast Idaho (aureus) and southwest Mon-
tana (richardsonii). He classified the former population as a new subspecies, aureus, and

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