Mammalian Biolog~

Zeitschrift fUr Saugetierkunde

ELSEVIER www.elsevier.de/mammbiol

OriginaL investigation
Discovery of extant Natalus major
(Chiroptera: NataLidae) in Cuba
By A. TEJEDOR, G. SILVA-TABOADA, and DIALVYS RODRIGUEZ-HERNANDEZ
Department of Mammalogy, American Museum of Natural History, New York, USA; Museo Nacional de Historia Nat-
ural, La Habana, Cuba; Centra de Investigaciones y Servicios Ambientales, Delegacion Territorial del Ministerio de
Ciencia Tecnologia y Medio Ambiente, Pinar del Rio, Cuba

Receipt of Ms. 09. 12. 2002

Acceptance of Ms. 01. 04. 2003

Abstract
The sole Cuban representative of the bat genus Nato/us, described as a fossil in 1919, was rediscov-
ered in 1992 as an extant colony in a cave in the western tip of the island. Other extant Greater An-
tillean populations of this genus were known before from Jamaica and Hispaniola and, together
with the fossil Nato/us from Cuba, were considered synonyms of the continental species Nato/us
stramineus. Morphometric comparison of the living Cuban Nato/us with other West Indian and
mainland populations of the genus supports the taxonomic separation of the Greater Antillean po-
pulations of Nato/us from Nato/us stramineus. We hypothesize that Nato/us major has a relatively
high extinction potential due to its probable limited capacity for dispersal, and to its specialization
in scarce hot-caves. We suggest that dependence on roosts of unreliable permanence probably in-
troduces a high degree of stochasticity in the process of extinction of obligate cave-dwelling bats.

Key words: Nato/us, Cuba, extinction, caves, rediscovery

Introduction
Extinction has played a determinant role in
shaping the Quaternary land mammal fau-
na of the West Indies (MORGAN and WOODS
1986; MACPHEE and FLEMING 1999). Of a
total of 133 species of land mammals (in-
cluding bats) recorded for the West Indies,
only 60 (45%) have survived until the pre-
sent, with some extinctions taking place just
a few hundred years ago (MACPHEE and
ITURRALDE-VINENT 2000). Although species
loss on Caribbean Islands have been less se-
vere for Chiroptera, most of the extirpation
and range reduction events of mammal spe-
cies in the West Indies have involved bats,
and in particular specialized cave-dwelling
species, i. e., those that roost exclusively in
warm and humid caves. The causes of such
extinctions have been ultimately attributed
to post-Pleistocene climate change with
concomitant changes in cave environments
and distribution (MORGAN 2001). Specula-
tions about the causes of extinction of West
Indian bats, however, have been obviously
limited by the lack of ecological informa-
tion on the extinct taxa.
Among West Indian specialized cave-dwell-
ing bats, species of the family Natalidae,
although spared from global extinctions,

1616-5047/04/69/03-153 $ 30.00/0. Mamm. biol. 69 (2004) 3 . 153-162

154 A. TEJEDOR et al.
are among the most highly affected by
range reductions. All three natalid genera,
Natalus, Chilonatalus, and Nyctiellus (re-
cently elevated from subgenera of Natalus;
MORGAN and CZAPLEWSKI 2003) have ex-
tinct populations in one or more islands
throughout the region. The genus Natalus
(sensu stricto), reported as extinct from se-
ven islands, has undergone more extirpa-
tion events than any of the two other West
Indian natalid genera (MoRGAN 2001). The
only population of Natalus hitherto consid-
ered extinct on a large island is a form de-
scribed as Natalus primus Anthony, 1919,
first discovered as a fossil in eastern Cuba
and later in 13 other fossil localities
throughout Cuba and Isla de Pinos (SILVA
TABOADA 1979).
Historically, the number of species recog-
nized within the genus Natalus has varied
from one to three depending on the author.
GOODWIN (1959) recognized three species:
Natalus stramineus Gray, 1838 (Mexico to
Brazil and Lesser Antilles); Natalus tumi-
dirostris Miller, 1900 (South America); and
Natalus major Miller, 1902 (Greater Antil-
les); the latter comprising three subspecies:
Natalus major major (Hispaniola); Natalus
major primus (Cuba, fossil); and Natalus
major jamaicensis Goodwin, 1959 (Jamai-
ca). Natalus major was later considered a
junior synonym of Natalus stramineus by
LINARES (1971), and this view was followed
by most authors (e.g., VARONA 1974; SILVA
1979; HALL 1981; KOOPMAN 1993). In con-
trast, MORGAN (1989; 2001) and MORGAN
and CZAPLEWSKI (2003) argued that mea-
surements and discrete characters support
the recognition of N. major as distinct from
N stramineus. Given that the Cuban form
of Natalus was previously known only from
fossils, it was excluded from previous taxo-
nomic analyses or received a merely periph-
eral treatment.
On 1 May 1992, while surveying bats in the
remote Cueva La Barca, at the western tip
of Cuba, AT and DRH discovered the first
living specimens of the same Natalus de-
scribed by ANTHONY in 1919. Here we re-
port this discovery and review the taxo-
nomic status of Natalus major. Also, we
report observations on the natural history
of N major in Cuba and discuss their impli-
cations for the protection of this bat and for
understanding causes of bat extinction in
the West Indies.
Material and methods
Within Peninsula de Guanahacabibes, Cueva La
Barca (= Cueva de La Barca) is located near
Cabo de San Antonio (21 °50'7" N, 84°56'35" W),
the westernmost tip of Cuba (Fig. 1). The vegeta-
tion of the area is largely undisturbed tropical
semideciduous forest growing on exposed Qua-
ternary limestone with scarce open bodies of
water. A weather station at Cabo de San Antonio
registered a mean annual temperature of 25.9 QC,
a mean annual precipitation of 1,423 mm, and
mean relative humidity of 81 % (NUNEZ JIMENEZ
and QUINTANA LARRAZ 1968).
Cueva La Barca has six chambers, with a com-
bined linear extension of about 500 m, and two
permanent bodies of water (ARMAS et al. 1989).
Three of the chambers are more or less well lit
and ventilated and the rest are dark. Given that
one of the dark chambers maintains high, stable
values of temperature and humidity (Tab. 1) Cue-
va La Barca is considered a 'hot cave', which in
the Neotropics are characterized by temperatures
of 26-40 QC and humidity> 90% (SILVA TABOADA
1977; RODRIGUEz-DuRAN 1998). The two other
dark chambers of the cave correspond to the
'temperate phase' of MORGAN (2001) because
their climatic conditions are intermediate be-
tween those of the hot chamber and the exter-
ior.
The bat community that inhabits Cueva de La
Barca includes 4 families and 13 species: Mor-
moopidae (Pteronotus parnelli, P macleayi,
P quadridens, Mormoops hlainvillii); Phyllosto-
midae (Macrotus waterhousei, Brachyphylla nana,
Erophylla sezekorni, Phyllonycteris poeyi, Mono-
phyllus redmani, Artibeus jamaicensis); N atalidae
(Chilonatalus micropus, Natalus major); and Ves-
pertilionidae (Eptesicus fuscus). The cave harbors
(excluding Artiheus, Eptesicus, and Macrotus) 10
of the 11 species of Cuban bats considered to be
"obligate cave-dwellers" by SILVA TABoADA
(1979). Most of the large bat populations that in-
habit Cueva La Barca are concentrated in the
three dark chambers.
We visited Cueva La Barca in May, August and
September of 1992; and in April and July 1993.
Observations on the habits of the bats were re-
corded in all visits, but specimens were collected
 Discovery of extant Nata/us major (Chiroptera: Natalidae) in Cuba 155

Gulf of Mexico

22'

Cabo de
San Anlonio

85' Caribbean Sea 84' 30 '

Fig. 1. Geographic location of Cueva La Barca (large black circle) in Guanahacabibes peninsula and fossil local-
ities of Nato/us major (small black dots in inset map) in Cuba.

Table 1. Air temperature and relative humidity recorded simultaneously inside the hot chamber of Cueva La Bar-
ca and outside the cave.

Date Time Temperature QC Relative humidity % Reference

In the hot Outside the In the hot Outside the
chamber cave chamber cave
28 Feb. 1987 01:00 29.2 26.1 98 89 ARMAS et al. (1989)
17:00 30.0 96
22 Jul. 1993 16:00 29.0 32.0 This study
22:30 28.5 26.5
8:00 28.1 25.0

only on the first and last visits. The bats were cap-
tured inside the cave using butterfly nets and
were held inside cloth bags for up to two hours
or sacrificed upon capture. Stomachs for diet
identification were extracted from bats captured
on the morning of 23 July 1993. Flight behavior
was incidentally observed on some animals re-
leased outside the cave during daylight hours,
10-15 m away from its entrance. Specimens were
preserved in the standard fashion (e. g., HANDLEY
1988) as skin/skull preparations or in fluid. Mor-
phological measurements follow SILVA TABOADA
(1979): forearm, tibia, greater skull length, condy-
lobasal length, zygomatic breadth, postorbital
width, braincase width, width across molars, and
mental length. Standard t-tests (two tailed,
n = 0.05, ZAR 1999) were used to compare mea-
surement means between sexes and between
neontological and Pleistocene material. Measure-
ments were taken from specimens housed in the
following institutions: American Museum of Nat-
ural History (AMNH); Florida Museum of Natur-
al History (FLMNH); Field Museum of Natural
History (FMNH); Museo Nacional de Historia
Natural, Havana (MNHN); National Museum of
Natural History, Smithsonian (USNM); or were
obtained from GOODWIN (1959) and LINARES
(1971).
Specimens examined: Natalus major Cuba (51):
AMNH 41009, MNHN 1-50 (field numbers), Ja-
maica (11) AMNH 181999, 182000, 214131,
246137, 246138; FLMNH 13688-13692; FMNH
93784, Dominican Republic (8) AMNH 97589,
97590, 238148, 2338149; USNM 101395 a,
156 A. TEJEDOR et al.

101395b, 221022; FLMNH 5517, Natalus strami-

neus Belize (5) FLMNH 9844, 9845, 9847; FMNH
121072, 121073, Guatemala (2) FMNH 64809,
64810, Costa Rica (1) USNM 566451, Dominica
(4) FMNH 18215, 44209-44211, Anguilla (I)
USNM 544834, Saba (1) FMNH 47701, Montser-
rat (1) FMNH 47702.

Results and discussion

MorphoLogy and taxonomy
The mandible of the living Natalus from
Cueva La Barca agrees with the holotype
of Natalus primus (AMNH 41009, Fig. 2) in
general morphology and shows the charac-
ter that diagnoses the taxon: long and slen-
der dentary with heavy teeth (molars taller
than 2/3 the width of the dentary). Like-
wise, the mental length (12.3 mm) of the
holotype falls within the range of variation
in mental length of La Barca specimens
(11.6-12.6 mm).
3mm
Fig. 2. Comparison between the holotype (A, compo-
site) of Nato/us primus and a specimen of Nato/us ma-
jor from Cueva La Barca (B). Note relationship between
height of molars and width of dentary.
The living Natalus from Cueva La Barca
show slight secondary sexual dimorphism
in size, with males having longer tibias and
larger skulls than females (P < 0.05, Tab. 2).
Comparison of recent specimens of Natalus

Table 2. External and cranial measurements of adult specimens of Nato/us major (Cueva La Barca, Guanahaca-
bibes). n = sample size, x = mean, sd = standard deviation. Level of significance of the differences between
means: * = P < 0.05.

Males Females t-value

n n
Variable x ± sd (min.-max.) x ± sd (min.-max.)

Forearm 26 24 1.0241
48.8 ± 1.24 (46.1-51.2) 48.5 ± 0.80 (47.0-50.1)
Tibia 26 24 4.3771 *
27.2 ± 0.71 (26.0-28.6) 26.4 ± 0.58 (25.4-27.1)
Greatest skull length 18 17 4.2240*
19.2 ± 0.37 (18.5-19.9) 18.7 ± 0.33 (18.1-19.5)
Condylobasallength 16 12 5.2907*
17.9 ± 0.29 (17.5-18.4) 17.4 ± 0.21 (17.0-17.7)
Zygomatic breadth 13 11 3.9598*
9.6 ± 0.24 (9.2-10.0) 9.3 ± 0.12 (9.1-9.5)
Postorbital width 18 18 3.1192*
3.4 ± 0.08 (3.2-3.5) 3.3 ± 0.11 (3.0-3.4)
Braincase width 14 11 1.4325
8.5 ± 0.11 (8.3-8.7) 8.4 ± 0.21 (8.1-8.7)
Width across molars 12 8 1.4037
6.7 ± 0.15 (6.5-6.9) 6.8 ± 0.16 (6.6-7.0)
Mental length' 18 17 3.2267*
12.1 ± 0.22 (11.9-12.6) 11.9 ± 0.14 (11.6-12.3)
 Discovery of extant Nata/us major (Chiroptera: Natalidae) in Cuba 157

from the Greater Antilles with those of the
Lesser Antilles and the continental Neotro-
pics shows that Greater Antillean Natalus
have the largest body size within the genus.
The mean forearm length of Greater Antil-
lean Natalus (47.4 mm, combining speci-
mens from Cuba, Jamaica, and Hispaniola),
is 4.4 standard deviations larger than that of
pooled specimens from the Lesser Antilles
and mainland populations (37.8 mm,
Fig. 3). Therefore, we concur with MORGAN
(1989,2001) and MORGAN and CZAPLEWSKI
(2003) in recognizing all Greater Antillean
populations of Natalus as representatives
of a distinct species: Natalus major. A study
of genic variation in Caribbean Natalus by
ARROyo-CABRALEs et al. (1999) also points
in this direction by showing that Jamaican
N. major may belong to a lineage basal to a
Central America/Lesser Antilles Natalus
stramineus clade.
Pleistocene fossils of Natalus major from
Masones-Jagtiey cave system in Central
Cuba are significantly larger than the living
population from Cueva La Barca in four
cranial dimensions (Tab. 3). This pattern
opposes that found in the Masones-Jagtiey
deposit for nine other Cuban bat species,
which increased in body size from older to
younger layers (SILVA TABOADA 1974). The

Mexico
Central America n = 81 n =71
Lesser Antilles Greater Antilles
South America v = 0.9 v =5.2

34 36 38 40 42 44 46 48 50 52
forearm length (mm)
Fig. 3. Variation in forearm length of adult specimens (sexes combined) of insular and continental populations
of the genus Nata/us. n = sample size, v = variance, range observed (horizontal line), mean (vertical line), one
standard deviation on each side of the mean (rectangle). Note non-overlapping ranges between mainland/Lesser
Antilles populations (Nata/us stramineus) and Greater Antilles populations (Nata/us major).

TabLe 3. Differences in skull dimensions (mm) between fossil and neontological adult specimens of Nata/us ma-
jor from Cuba (sexes combined). n = sample size, x = mean, sd = standard deviation Level of significance of the
differences between means: ** = P < 0.01; *** = P < 0.001.

variable Fossil deposit Living population t-value

Masones-JagUey Cueva La Barca
n n
x ± sd (min.-max.) x ± sd (min.-max.)

Greatest skull length 16 35 6.60***

20.0 ± 0.21 (19.7-20.4) 19.0 ± 0.41 (18.1-19.9)
Zygomatic breadth 11 24 4.29***
9.6 ± 0.10 (9.6-9.9) 9.3 ± 0.23 (9.1-10.0)
Postorbital width 21 36 1.33**
3.5 ± 0.09 (3.4-3.7) 3.3 ± 0.10 (3.0-3.5)
Braincase width 19 25 11.10***
8.9 ± 0.13 (8.7-9.1) 8.4 ± 0.16 (8.1-8.7)
158 A. TEJEDOR et al.
lack of neon to logical specimens of N. major
from Masones-Jagiiey, however, prevents
concluding whether the difference is due to
geographical or chronological variation.
NaturaL history
Roosting: On July 1993, visual estimates in-
dicated that a few thousand Natalus major
inhabited Cueva La Barca. Observations
made as recently as August 2001 suggest
that this bat remains abundant in the cave
(F. BALSEIRO pers. comm.). Natalus major
occupies the two temperate chambers of
Cueva la Barca, roosting almost exclusively
along the east walls of the chambers, which
are more sinuous than the west walls and
are farther from entrances. Roosting groups
contain a few dozen to a few hundred bats.
Individual bats hang from one or both feet
without ventral contact with the substrate
and regularly spaced from each other, keep-
ing a distance between themselves usually
greater than 10 cm. The roosting groups
scatter on the lower parts of walls, at about
1 m high, and occasionally on the low roofs
of wall niches. Specific roosting spots were
regularly occupied by similarly sized groups
of N. major in all visits to the cave, indicat-
ing that seasonal migrations out of Cueva
La Barca, as mentioned by LINARES and Lo-
BIG-A (1973) for other Natalidae, are unli-
kely.
Six other bats (Pteronotus parnelli, Mor-
moops blainvillii, Brachyphylla nana, Ero-
phylla sezekorni, Artibeus jamaicensis, and
Chilonatalus micropus) roost together with
Natalus major in Cueva La Barca, yet none
of the species were ever observed in mixed
groups. Groups of N major sometimes roost
adjacent to groups of M. blainvillii (which
also appear to favor the cave's walls as roost-
ing areas). A close spatial association be-
tween N major and M. blainvillii has also
been observed in Hispaniola (K. CHRISTEN-
SON pers. com.), suggesting that both species
have similar preferences for roosting sites
inside a cave. Captive Natalus major are ag-
gressive and frequently attack bats of other
species, even of larger size, if confined to-
gether in a small enclosure.
In general, individuals of Natalus major re-
main active while roosting during the day,
taking flight at the slightest sign of distur-
bance (human steps, distant glare from a
flash light), but usually some individuals al-
low the extreme close proximity of the ob-
server for a few seconds. If disturbance
(e. g. artificial illumination) persists for
some minutes, all bats move to alternative
roosting areas of the same chambers, join-
ing the groups that may already be present
there. When moving to other areas inside
the cave, the bats invariably fly extremely
close to the walls and about 1 m from the
floor, forming a highly distinctive, continu-
ous stream of bats along the wall. No
N. major was ever captured with a butterfly
net more than 2 m away from any cave wall.
A similar in-cave flight behavior of N ma-
jor was observed in St. Clair Cave, Jamaica,
from which GOODWIN (1970) reported that
the bats "fluttered about in a relatively
tight flock like a mass of large moths".
Foraging and reproduction: The flight of
Natalus major is extremely slow and highly
maneuverable. Several individuals released
during the day in the forest outside the cave
entrance showed a greater tendency and
ability to fly through highly cluttered un-
derstory vegetation than most other species
(except Chilonatalus micropus) present in
Cueva La Barca. As hypothesized for spe-
cies of the Old World vespertilionid genus
Kerivoula (KINGSTON et al. 1999) such a
flight style probably enables N major to fly
and exploit food resources in very narrow
spaces. The echolocation calls of the Cuban
Natalus are unknown but it produces weak
and high-pitched audible sounds when held
in the hand.
Analysis of stomach contents obtained at
dawn on 23 July 1993 revealed that the bats
had consumed insects belonging to eight or-
ders. The most commonly represented
groups were: Lepidoptera, Orthoptera
(Gryllidae), and Coleoptera. The remaining
orders (represented by single cases) were
Hymenoptera (Formicidae), Neuroptera,
Diptera, Homoptera, and Hemiptera.
Given the slow flight of this bat and its high
rate of dehydration outside the cave (as
 Discovery of extant Nata/us major (Chiroptera: Natalidae) in Cuba
judged by the rapid increase in brittleness
of patagia of handled animals) it is probable
that its foraging range is relatively small.
On two occasions (April and July 1993) the
species was not seen inside the cave be-
tween 22:00 and 24:00 hours and animals
collected early in the morning on 23 July
1993 had full stomachs indicating that fora-
ging may extend until daybreak. The open
bodies of water present in caves where
N. major roosts (Cueva La Barca, Saint
Clair Cave in Jamaica, and at least three
caves in Hispaniola; FINCHAM 1997;
K. CHRISTENSON pers. corn.), might serve as
sources of drinking water for this bat.
The reproductive activity of Natalus major
is largely unknown. At midday, on 17 April
1993, most N. major groups were observed
to have abandoned their usual roosting
areas in Cueva La Barca and moved to the
entrance of the hot chamber. Most of the
N. major groups observed in this unusual
location were composed of copulating pairs,
which were much more easily approached
by humans than isolated bats. On the same
occasion, other bat species, most notably
Monophyllus redmani, were also observed
copulating in large numbers in the entrance
to the hot chamber. Three females of
N. major collected on 1 May 1992 were
pregnant, each holding a single embryo.
The extant Cuban Natalus and West Indian
bat extinctions
During the late Quaternary, 27 bat species
have undergone either global or local ex-
tinctions in the West Indies, with local ex-
tinctions often encompassing a few islands
and rarely the entire region. As many as
75% of the extinction events have involved
obligate cave-dwelling bats, including mor-
moopids, phyllonycterines, and natalids. Ex-
tinct populations of natalids have been
documented from nine West Indian islands:
the Bahamas (Abaco, Andros, New Provi-
dence, Cat, Exuma, Grand Caicos), Grand
Cayman, Isla de Pin os, and Cuba (MoRGAN
2001).
Before the discovery of the extant popula-
tion of Natalus major in Guanahacabibes,
159
Cuba seemed anomalous as the only large
island of the Antilles where a species of Na-
talidae had gone extinct. The present study
dispels the idea that N. major is extinct in
Cuba and brings the island's extant bat
count to 27 species. With three species of
Natalidae (Nyctiellus lepidus, Chilonatalus
micropus, and Natalus major), Cuba has
the highest diversity (in sympatry) of this
family in the entire Neotropics. Two of
these species occur syntopically in Cueva
La Barca, (and Nyctiellus lepidus has been
reported from a locality 20 km W of Cueva
La Barca; SILVA TABOADA 1979), implying
that coexistence is possible among N. major
and other Cuban Natalidae.
Unfortunately, since Cueva La Barca is
likely to be the sole locality of extant Nata-
Ius major on the entire island, the only Cu-
ban form of the genus Natalus may be in
high risk of extinction. Although no extant
colonies of N. major were ever found in
Cuba until 1992, fossil localities of this bat
are widespread throughout Cuba and Isla
de Pinos (SILVA TABOADA 1979). Considering
(1) that natalids are exclusive cave dwellers,
usually congregating in large, conspicuous
colonies and (2) that Cuban caves have
been surveyed extensively with regard to
bat fauna, the probability that living indivi-
duals of this taxon have remained unde-
tected in other caves all these years is very
low. Efforts to protect this rare bat must
take into consideration the causes of its ap-
parent restriction to Guanahacabibes.
Cueva La Barca is located within one of the
largest remaining tracts of lowland forest in
Cuba (DINERSTEIN et al. 1995). The appar-
ent relictual distribution of extant Natalus
major in Cuba, therefore, may seem to be
related to historical changes in vegetation
cover on the island, both anthropogenic or
climatic in origin (PREGILL and OLSON
1981). In Hispaniola, however, N. major
persists in areas partially cleared for agri-
culture (TEJEDOR own observ.), and is found
throughout the island from the dry south
(arid scrub) to the moist northeast (rain for-
est), spanning a 700-2400 mm precipitation
gradient (BOLAY 1997), suggesting that
neither vegetation type nor moderate forest
160 A. TEJEDOR et al.
disturbance are the most important factors
limiting this bat.
Specialized roosting requirements, in con-
trast, may be a limiting factor for Natalus
major. Natalids generally roost in associa-
tion with large, multi-species communities
of obligate cave-dwelling bats that select
warm, humid caves - mostly hot caves - as
permanent roost sites. Although caves are
common geomorphologic features in the
West Indies, a small proportion of these
(8% in Puerto Rico, RODlUGUEZ-DuRAN
1998) are hot and house diverse bat com-
munities. Only caves with small entrances
and a topography that promotes minimal
ventilation can become hot by trapping the
heat and water generated by large numbers
of bats (SILVA TABOADA 1979; RODRfGUEZ-
DURAN 1998). An increase in air circulation
caused by cave erosion could disrupt the
hot cave environment.
About 80% of the fossil localities of Nata-
Ius major in Cuba (ANTHONY 1919; SILVA
TABOADA 1979) and other Caribbean islands
(M ORGAN 1989, 1994) correspond to 'fossil
hot caves' that are currently well ventilated
yet contain fossils of typical hot-cave bats.
This suggests that the extinction of N major
throughout most of Cuba did not result
from elimination of a few sensitive species
from supersaturated hot-cave communities
as has been speculated (MoRGAN 2001).
Rather, extirpation events may have coin-
cided with the accidental (stochastic) col-
lapse of sections of caves due to erosion,
leading to the disappearance of entire hot-
cave communities.
That Natalus major disappeared from large
areas of Cuba following the obliteration of
hot caves while most other obligate cave-
dwelling bats did not can be explained by
the apparent low dispersal ability of nata-
lids. Due to their slow flight and extreme
propensity for dehydration (SILVA TABOADA
1979; HOYT and BAKER 1980; this study), na-
talids may be more adversely affected by
the sudden loss of a roost site than perhaps
longer-ranging hot-cave bats (e. g. most
mormoopids), which may be better suited
for moving into new habitat. Because of its
larger size relative to other natalids, N ma-
jor may also need larger spaces for main-
taining viable colony sizes, further limiting
its choices of roost sites. The demographic
dynamics of N major may be thus more
strongly bound to stochastic losses of habi-
table cave environments than that of other
hot-cave bats.
As known thus far, the survival of Natalus
major in Cuba depends on a single perma-
nent roost, Cueva La Barca. Although
anthropogenic disturbance may not have
played a determinant role in the extirpation
of N major throughout most of Cuba, hu-
mans currently represent the greatest po-
tential threat to this bat. Even in the ab-
sence of active harassment, large bat
colonies formerly present in other Cuban
caves have nearly disappeared with increas-
ing numbers of visitors to these sites (SILVA
TABOADA 1979). In Cueva La Barca, this
kind of decline in bat populations would
likely result in a disruption of the hot-cave
environment. Minimization of human incur-
sions should thus be a top priority for the
protection of Cueva La Barca.
AcknowLedgements
We are grateful to EMILIO ALFARO, JORGE ISAAc,
YAZMIN PERAZA, ESTHER PEREz, DIANELIS RODRi-
GUEZ, TIRSO RODRIGUEZ, and ARMANDO URQUIOLA
Jr., for field assistance. We also thank Grupo Espe-
leol6gico Guaniguanico for loan of equipment and
MARINO FELIZOLA, LUIS R. HERNANDEZ (who also
identified insect remains), and DAVID S. SMITH for
logistic support. Access to collection material was
possible thanks to the late KARL F. KOOPMAN
(AMNH), ALFRED L. GARDNER (USNM), the late
PHILLIP HERSHKOVITZ (FMNH), and JOSE A. OT-
TENWALDER (FLMNH). OMAR LINARES shared with
us his archives on Natalidae while FERNANDO BAL-
SEIRO, and KEITH CHRISTENSON provided unpub-
lished observations. We thank ANNIKA KEELEY for
translations into German. Critical reviews by Ro-
BERT P. ANDERSON, PATRICIA CAULFIELD, LILIANA
M. DAVALOS, NANCY B. SIMMONS, ERIC STINER, VA-
LERIA DA C. TAvAREs, and MARCELO WEKSLER re-
suIted in a significant improvement of the manu-
script version. The visits to the mentioned
institutions were financed by the RARE Center
for Tropical Conservation (Philadelphia). Field
work in Cuba was supported by Museo Nacional
de Historia Natural, Havana.
 Discovery of extant Nata/us major (Chiroptera: Natalidae) in Cuba 161

Zusammenfassung
Zur Entdeckung Lebender Natalus major (Chiroptera: NataLidae) auf Kuba
Der einzige kubanische Vertreter der Fledermausgattung Nata/us wurde im Jahr 1919 als Fossil be-
schrieben und im Jahr 1992 als eine lebende Kolonie in einer Hohle auf der westlichen Spitze der
Insel wiedergefunden. Auf den GroBen Antillen waren andere Populationen dieser Gattung schon
aus Jamaica und Hispaniola bekannt. Zusammen mit dem fossilen Nata/us-Exemplar aus Kuba wur-
den sie als synonym mit der kontinentalen Art Nata/us stramineus betrachtet. Morphometrische Ver-
gleiche der lebenden kubanischen Nata/us-Exemplare mit anderen westindischen Populationen und
Populationen der Gattung vom Festland unterstUtzen jedoch die taxonomische Trennung der Nata-
/us-Populationen von den GroBen Antillen von Nata/us stramineus. Wir stellen die Hypothese auf,
daB Nata/us major aufgrund seiner wahrscheinlich begrenzten Ausbreitungsfahigkeit und seiner
Spezialisation auf seltene warme Hohlen ein relativ hohes Aussterbepotential hat. Wir nehmen an,
daB die Abhangigkeit von permanenten aber erforderlichen Quartieren wahrscheinlich einen hohen
Grad von Stochastizitat in dem AussterbeprozeB von obligaten Hohlen bewohnenden Fledermausen
bedingt.

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Authors' addresses:
ADRIAN TEJEDOR, Department of Mammalogy,
American Museum of Natural History, Central
Park West at 79 Street, New York, NY 10024,
USA (e-mail: tejedor@amnh.org); GILBERTO SILVA
TABOADA, Museo Nacional de Historia Natural,
Obispo 61, Plaza de Armas, La Habana Vieja,
Ciudad de La Habana, Cuba; DIALVYS RODRf-
GUEz-HERNANDEZ, Centro de Investigaciones y
Servicios Ambientales ECOVIDA, Delegaci6n
Territorial del CITMA. Calle Col6n no. 61 entre
Maceo y Virtudes, Pinar del Rio, Cuba