Morphological Variation of Primary
Reproductive Structures in Males of
Five Families of Neotropical Bats

MATEUS R. BEGUELINI,1 CÍNTIA C. I. PUGA,2 FABIANE F. MARTINS,2 ANDRÉ
H. S. BETOLI,1 SEBASTIÃO R. TABOGA,1 AND ELIANA MORIELLE-VERSUTE2*

1Department of Biology, São Paulo State University, UNESP/IBILCE, São Jos�e do Rio
Preto, São Paulo 15054-000, Brazil

2Department of Zoology and Botany, São Paulo State University, UNESP/IBILCE, São Jos�e
do Rio Preto, São Paulo 15054-000, Brazil

ABSTRACT
Bats present unique features among mammals with respect to repro-

duction, and although neotropical bats do not have a hibernation period,
many of their reproductive characteristics vary seasonally and interspecifi-
cally. Thus, this work aimed to examine the reproductive structures of 18
species belonging to five families of Brazilian bats. The gross anatomy of the
testes varied little; however, the epididymis of Emballonuridae and Vesper-
tilionidae showed exceptional structures with a large elongation of the cau-
dal region. We observed a wide variation in the positioning of the testes:
Phyllostomidae and Noctilionidae presented external testes; Emballonuri-
dae and Molossidae presented migratory testes that may be in intra-abdomi-
nal or external positions; and Vespertilionidae displayed scrotal testes. In
the histological evaluation, we observed a different pattern in vespertilionid
species, with testicular regression and sperm retention/storage in the cauda
epididymis in the five species analyzed. Similar testicular regression was
observed in Molossops temminckii; however, sperm retention/storage was
not observed in this species. These data suggest that although the species
analyzed are tropical species that do not present a prolonged period of tor-
por (hibernation), they still maintain a period of seminiferous tubule regres-
sion and sperm storage very similar to that observed in hibernating bats.
Anat Rec, 296:156–167, 2013. VC 2012 Wiley Periodicals, Inc.

Key words: Chiroptera; epididymis; quiescence; testes;
Vespertilionidae

The order Chiroptera is the second largest order of
mammals, with approximately 202 genera and 1116 spe-
cies widely distributed among all tropical and temperate
regions, except for some remote oceanic islands and Ant-
arctica (Reis et al., 2007). It is classically divided into two
suborders: Megachiroptera and Microchiroptera. The first
is found exclusively in the Old World and is composed of
only one family (Pteropodidae), with approximately 42
genera and 186 species; while the second is cosmopolitan,
with 17 families and 930 species (Simmons, 2005).

In Brazil, there are approximately 165 species of Micro-
chiroptera, distributed into nine families: Emballonuridae,
Furipteridae, Molossidae, Mormoopidae, Natalidae, Noctilio-
nidae, Phyllostomidae, Thyropteridae, and Vespertilionidae
(Reis et al., 2006). Of these families, Furipteridae, Mormoo-
pidae, Natalidae, Noctilionidae, Phyllostomidae, and

Thyropteridae are found exclusively in the Americas, while
Emballonuridae, Molossidae, and Vespertilionidae are also
found occurred in the Old World.

Grant sponsor: São Paulo State Research Foundation
(FAPESP); Grant number: 2009/16181-9, and 2009/03470-2.

*Correspondence to: Eliana Morielle-Versute, Rua Crist�ovão
Colombo n� 2265, Jardim Nazareth, 15054-000, São Jos�e do Rio
Preto, São Paulo, Brazil. Tel.: þ55-17-32212369. FAX:
þ 55-17-32212374. E-mail: morielle@ibilce.unesp.br

Received 10 January 2012; Accepted 13 September 2012.

DOI 10.1002/ar.22613
Published online 1 November 2012 in Wiley Online Library
(wileyonlinelibrary.com).

THE ANATOMICAL RECORD 296:156–167 (2013)

VVC 2012 WILEY PERIODICALS, INC.



It has been a great effort by researchers aiming to study
the characteristics of species of temperate and tropical
regions. Although most emphasis has been to the temper-
ate species, which are subject to abrupt changes in
temperature, the results observed for tropical species, has
been very interesting. The most studies in tropical species
generally deal with aspects of female reproduction and
pregnancy (Rasweiler, 1972, 1974, 1978, 1982, 1987, 1993;
Quintero and Rasweiler, 1974; Badwaik et al., 1997; Ras-
weiler and Badwaik, 1997; Badwaik and Rasweiler, 2001;
Komar et al., 2007; Nolte et al., 2009), and only a few
examinations are focused on male patterns, and these are
primarily based in ecological–behavioral perspectives (Wil-
son and Findley, 1971; Fabi�an and Marques, 1989;
Heideman et al., 1992; Heideman and Bronson, 1994;
Zort�ea, 2003; L�eon-Galv�an et al., 2005; Chaverri and
Kunz, 2006; Costa et al., 2007; Ortêncio-Filho et al., 2007).

The characteristics exhibited by females are directly
related to the specializations of males and anatomical,
physiological, embryological, and behavioral specializa-
tions observed in females of some species with wide
distribution and representation in the tropics constitute
significant evidence that they have diverse reproductive
patterns (Rasweiler, 1987; Rasweiler and Bonilla, 1992;
Rasweiler and Badwaik, 1997; Rasweiler et al., 2000;
Badwaik and Rasweiler, 2001; Rasweiler et al., 2011).

Despite all these observations, almost no evaluations
of these species from a morphological/histological view-
point have been made (Krutzsch et al., 2002; Krutzsch
and Nellis, 2006; Beguelini et al., 2009; Duarte and
Talamoni, 2010). Thus, this study aimed to analyze the
male primary reproductive structures (testes and epidi-
dymis) of 18 species belonging to five of the great
families of neotropical bats, to investigate the possible
morphological differences.

MATERIALS AND METHODS

Species, Aging, and Licenses

The primary reproductive structures (testes and epi-
didymis) of 18 species of exclusively neotropical bats,

which belong to five different families, Emballonuridae
(two spp.), Molossidae (four spp.), Phyllostomidae (six
spp.), Noctilionidae (one spp.), and Vespertilionidae
(five spp.), were morphologically analyzed in this study.
Most specimens were collected in northwest São Paulo
state, Brazil (São Jos�e do Rio Preto: 49W 220 4500 20S
490 1100); however, some species were obtained from
captures in the central region of Goi�as state, Brazil
(Goiânia: 49W 150 1400 16S 400 4300), and the more diffi-
cult-to-collect species were obtained from the
Chiroptera collection at the São Paulo State University,
UNESP-IBILCE (Table 1). The captures occurred from
June 2008 to June 2010.

Only adult males were used in this study. The bats
were classified as adults based on their body weight,
complete ossification of the metacarpal epiphyses, wear
of the teeth (De Knegt et al., 2005), and the presence of
spermatozoa inside the testes and/or the cauda epididy-
mis. All of the studied animals were sexually mature
adults; however, we were unable to determine whether
the animals were in basal reproductive or peak breeding
conditions or their social status, so our studies indicate
a mixture of these cases, with animals collected during
or outside of their peak breeding periods.

The capture and captivity of all specimens were
authorized by the Brazilian institution responsible for
wild animal care (Instituto Brasileiro do Meio Ambiente,
IBAMA, Process: 21707-1), and a proposal of this study
was assessed and approved by the Ethics Committee for
Animal Experimentation (Document: Protocol. 013/2009
CEEA/IBILCE-UNESP).

The animals were treated according to the recom-
mendations of the Committee on Care and Use of
Laboratory Animals from the Institute of Laboratory
Animal Resources, National Research Council, Guide
for the Care and Use of Laboratory Animals" (Commit-
tee on Care and Use of Laboratory Animals, 1980), and
after being sacrificed they were deposited in the Chi-
roptera collection at the São Paulo State University
(UNESP-IBILCE).

TABLE 1. List of specimens analyzed in the present study

Family Species São Pauloa Goi�asb Collectionc

Emballonuridae Peropteryx macrotis – – 5
Rhynchonycteris naso – 2 3

Molossidae Eumops glaucinus 5 – –
Molossops temminckii 4 4 –
Molossus molossus 25 – –
Molossus rufus 5 – 3

Phyllostomidae Artibeus lituratus 18 – –
Artibeus planirostris 59 – –
Carollia perspicillata 8 – –
Glossophaga soricina 14 – –
Phyllostomus discolor 5 – –
Platyrrhinus lineatus 20 – –

Noctilionidae Noctilio albiventris 5 – 5
Vespertilionidae Eptesicus furinalis 5 – –

Histiotus velatus 5 – 1
Lasiurus blossevillii 1 – 4
Myotis albescens – 3 2
Myotis nigricans 30 1 –

aSpecimens collected at São Paulo state, Brazil (49W 220 4500 20S 490 1100).
bSpecimens collected at Goi�as state, Brazil (49W 150 1400 16S 400 4300).
cSpecimens obtained from the Chiroptera collection at the Sao Paulo State University, UNESP-IBILCE.

REPRODUCTIVE CHARACTERISTICS OF BATS 157



Animal Processing and Documentation

The animals were euthanized by cervical dislocation
and placed supinely on the dissection board. An incision
was made in the abdominal region in the caudal direc-
tion, and the skin was moved to the sides. The locations
of the testes and epididymis were documented, and these
organs were then removed for future histological analy-
sis. In animals with testes located inside the abdominal
cavity, the subcutaneous tissue was cut, the area was
exposed with forceps, the region was photographed, and
the testes and epididymis were removed. The gross docu-
mentation was accomplished with a 7.0-mp HP
Photosmart M627 camera (6.0–18.0 mm) with a 3x
optical zoom and a 5x digital zoom.

Corporal and Testicular Weights,
Gonad-Somatic Index, and Statistical Analysis

The animals and the testes were weighed on a balance
with an accuracy of three decimal places. The specimens
from the UNESP-IBILCE collection were weighed and
their weights were increased by 33% to compensate for
the weight lost during the fixation process, as described
by Baker (1958).

The gonad-somatic index was calculated by dividing
the weight of the testes by the corporal weight of the
animal and recording the result to the fourth decimal
place. The values were expressed using the mean value
6 standard deviation.

All data were analyzed by a nonparametric multiple
independent test (Kruskal–Wallis: P � 0.05) using the
Statistica 7.0 software (CopyrightVC Statsoft, Inc.
1984–2004).

Histology

The reproductive status of each specimen was eval-
uated histologically to confirm its sexual maturity. The
presence of spermatozoa inside the testes and in the
cauda epididymis was used as an indication of sexual
maturity and activity. The testicular regression was
characterized by the observation of only the spermatogo-
nia and Sertoli cells in the seminiferous epithelium and
by a relative decrease in epithelium height and increase
of interstitial tissue.

After surgical excision, the testes and epididymis were
immersed in Bouin fixative solution for at least 24 hr,
dehydrated in a graded ethanol series, embedded in
glycol methacrylate (Historesin, Leica Instruments), and
sectioned (1 lm thickness) using a Leica RM 2155 micro-
tome. Tissue sections were stained with hematoxylin-
eosin (Ribeiro and Lima, 2000) and analyzed using an
Olympus BX60 microscope with Image-Pro-Plus for
Windows computer image analysis software.

RESULTS

Position of the Testes–Epididymis Complex

The position and the framework support complex of
the testes and epididymis widely varied in the five fami-
lies analyzed.

The testes–epididymis complex (TEC) of all specimens
of the six species of phyllostomid bats analyzed were
found outside the abdominal cavity, located laterally to

the abdomen (Fig. 1A, arrows), in the proximity of the
inguinal canal (Fig. 1D, arrow). Its dislocation to a more
caudal position, laterally to the base of the penis
(parapenial), was observed in all species (Fig. 1B), with
this movement apparently subject to the control of the
animal and able to be performed within a few minutes.
The TEC was located between the skin and muscles,
with only a thin and transparent tunica vaginalis cover-
ing it (Fig. 1C). The presence of a true scrotum was not
observed in these animals, but only a relaxation in the
skin-muscle connection that allows the movement of the
TEC (Fig. 1A,B).

Similar to phyllostomid bats, the TEC of Noctilio albi-
ventris (Noctilionidae) was also observed outside the
abdominal cavity, laterally to the abdomen; however, it
is enclosed in an unusual pocket-like pouch that is
directly associated with a true scrotum (Fig. 1E–H). The
scrotum presents no hair on its surface and can be
clearly distinguished by having a thin layer of skin with
numerous scattered short barbs and long barbs in its
periphery (Fig. 1F). The pocket-like pouch was consti-
tuted by a thick layer of tunica vaginalis surrounded by
large amounts of adipose tissue (Fig. 1G,H). The disloca-
tion of the TEC of the fold into the scrotum is easily
observed (Fig. 1E,F and 1G,H).

The position of the TEC in Molossidae specimens
varied as follows: (i) both TECs located outside the ab-
dominal cavity, between the skin and muscles, inside a
small scrotum (Fig. 1I) or dislocated to the vicinity of
the inguinal canal (Fig. 1J, arrow); (ii) a TEC located
outside and another inside the abdominal cavity
(Fig. 1K); or (iii) both TECs located within the
abdominal cavity, dorsally arranged below the kidneys
(Fig. 1L–M).

The TEC position in the emballonurid bats was
unique, with each cauda epididymis always externally
located in the lateral of the base of the penis (parape-
nial), within a subcutaneous pouch (Fig. 1O), whereas
the testes varied in position from the parapenial (Fig.
1O) to an intra-abdominal position (Figs. 1P–Q). When
the testes were externally located, the epididymis made
a loop around them (U-shaped), with the caput and
cauda epididymis close to each other (Fig. 1O). When
within the abdominal cavity, the testes were located
behind the prostate complex, arranged caudally to the
kidneys (Fig. 1P–Q), and the caput epididymis was
joined with the testes. Additionally, in these cases, the
corpus epididymis elongates a great deal (Fig. 1Q), and
the cauda epididymis remained enclosed in the pouch.
The maintenance of the cauda epididymis position is due
to the presence of suspensor ligaments (connective
tissues) that attach it directly to the subcutaneous
pouch. Despite the presence of an external pouch, we do
not observe a true scrotum in these species (Fig. 1N).

We observed that the testes of all five vespertilionid
species studied were external and permanently scrotal
(Fig. 1R–U). The maintenance of their position is
achieved by suspensor ligaments (connective tissues)
that connect the prolonged cauda epididymis to the
caudal skeleton and the juxtaposition of the cauda epidi-
dymis between the two interfemural membranes
(uropatagium). The length of the cauda epididymis
seems to vary seasonally and, at the apex of its elonga-
tion, it may be observed in the living animal (Fig. 1R,
arrow).

158 BEGUELINI ET AL.



Figure 1 (Legend, overleaf.)

REPRODUCTIVE CHARACTERISTICS OF BATS 159



Morphology of the Testes and Epididymis

The gross anatomy of the testes varied little, being
more rounded in the Phyllostomidae species (except
P. discolor, which had bean-shaped testes) (Fig. 2A–F),
oval in the Emballonuridae (Fig. 2M–O), and more elon-
gated in the Molossidae (Fig. 2I–L), Noctilionidae
(Fig. 2G–H), and Vespertilionidae species (Fig. 2P–T).

The epididymis in all species analyzed was formed by
a long and highly convoluted tubule that can be divided
into three main regions: (i) caput, the initial portion that
is juxtaposed with the confluence of the rete testes net-
work; (ii) corpus, the medial part of the epididymis; and
(iii) cauda, the end portion of the epididymis that
communicates directly with the deferens ducts (Fig. 2).
In contrast, the size and positioning, especially of the
cauda epididymis, varied considerably.

The epididymides of Phyllostomidae, Molossidae, and
Noctilionidae were connected laterally to the testes for
their entire extension. Phyllostomidae had a low degree of
adhesion to the testis and a highly developed caudal region,
which remained fully attached to the testis in all species
(Fig. 2A–F), except in G. soricina, where the caudal region
appeared to be free (Fig. 2D), and in P. discolor, where it
formed a lateral elevation (Fig. 2E). Molossidae presented
the highest degree of adhesion to the testis and the small-
est size of the caudal region (Fig. 2I–L). Unlike the other
families, Noctilionidae presented a further development of
the caput than the cauda epididymis (Fig. 2G–H).

In Emballonuridae and in the four genera of Vesperti-
lionidae analyzed, the epididymis showed a large
elongation of the caudal region that was not attached to
the testes. The size of this elongation varied individually
and in some specimens was larger than the testis itself.
In all of the emballonurid specimens analyzed, the cauda
epididymis remained enclosed inside the scrotum, form-
ing a loop around the testes (Fig. 2M–N), when it was
external or had an elongated shape (Fig. 2O) when it
was internal. However, in Vespertilionidae, it extended
parallel to the caudal skeleton, with its position being
maintained by filaments that connected it to the caudal
skeleton and by juxtapositions between the anterior and
posterior interfemural membranes (uropatagium). It was
not directly attached to the testes, but, in most speci-
mens, remained near them because both were covered
with a highly pigmented fascia (black) (Fig. 2P–T).

Corporal and Testicular Weights, Gonad-So-
matic Index, and Statistical Analysis

The corporal and testicular weights and the gonad-so-
matic index of all analyzed species and the statistical
analysis are shown in Figure 3. Because of an inability
to acquire data from M. albescens and R. naso, they are
absent from the figure. The Phyllostomidae family pre-
sented the highest values and the largest variation in
the corporal weights, presenting significant differences
among all species (Fig. 3A). The Molossidae family also
presented a large variation; however, their corporal
weights were intermediate between those of the Phyllos-
tomidae and Vespertilionidae families. In contrast, the
Vespertilionidae species presented the lowest corporal
weights values and the smallest variation between spe-
cies. The corporal weight of N. albiventris closely
approximates that of the medium-sized Phyllostomidae
species and that of the large Molossidae species, whereas
the corporal weight of P. macrotis approximates that of
the Vespertilionidae species (Fig. 3A).

The Phyllostomidae family also presented the highest
testicular weight values (Fig. 3B) and high gonad-so-
matic index values (Fig. 3C). Phyllostomus discolor,
although not the largest species, presented the largest
gonads and also the highest index. The smallest species
of Phyllostomidae, C. perspicillata and G. soricina, also
had the smallest gonads (Fig. 3B); however, their
indexes were greater than those of larger species, such
as A. lituratus (Fig. 3C).

The Molossidae family presented intermediate gonad
weights, with the exception of M. temminckii (Fig. 3B).
Again, we observed that smaller species have higher
indexes (M. temminckii and M. molossus; Fig. 3C). Ves-
pertilionidae presented the lowest gonad weights (Fig.
3B). Lasiurus blossevillii presented the lowest index,
whereas E. furinalis and M. nigricans presented high
indexes. Noctilio albiventris presented a testicular
weight and gonad-somatic index near those of the molos-
sids, and P. macrotis (Emballonuridae) presented values
near those of the vespertilionids (Fig. 3A–B).

Histological Evaluation

The histological evaluation of all the analyzed species
of Phyllostomidae (Fig. 4A–L), Noctilionidae (Fig. 4S–T),

Fig. 1. General arrangement of the testes–epididymis complex
(TEC) of the analyzed bats. A–D. Phyllostomidae pattern, Artibeus pla-
nirostris. Note the dislocation of the TEC from the lateral of the abdo-
men (A, arrows) to the base of the penis (B, arrow), the thin and
transparent tunica vaginalis that covers it (C) and the passage of the
duct deferens (DD) and spermatic artery (Sa) through the inguinal
canal (D, arrow). E–H. Noctilionidae pattern, Noctilio albiventris. Note
the presence of a true scrotum (E, TEC outside the scrotum; F, TEC
inside the scrotum) that presented numerous scattered short barbs (F,
arrow-head) and long barbs at its periphery (F, arrow); also, note that
the TEC may dislocated from the scrotum to an unusual pocket-like
pouch (E-F and G-H) and that this pocket-like consisted of a thick
layer of tunica vaginalis surrounded by large amounts of adipose tis-
sue (G-H). I–M. Molossidae pattern, Eumops glaucinus (I), Molossops
temminckii (J and L-M) and Molossus molossus (K). Note the presence
of a true scrotum (I) and the migratory pattern of the TEC: (i) both
external TECs (J); (ii) a TEC outside and another inside the abdominal

cavity (K, arrows); and (iii) both TECs located within the abdominal
cavity (L-M). When both are outside, they may stay in the scrotum (I)
or be dislocated to the vicinity of the inguinal canal (J, arrow). N–Q.
Emballonuridae pattern, Rhynchonycteris naso (N and P-Q) and Per-
opteryx macrotis (O). Note the absence of a scrotum (N); the loop of
the epididymis around the testis when it was located externally, inside
the subcutaneous pouch (O); and the elongation of the corpus epidi-
dymis (Q) when the testes were located within the abdominal cavity,
behind the prostate complex, arranged caudally to the kidneys (P). R–
U. Vespertilionidae pattern, Myotis nigricans (R-S), Histiotus velatus
(T), and M. albescens (U). Note the considerable elongation of the
cauda epididymis and the external, permanently scrotal testes of all
species, which may be observed in the living animal (R, arrow). (A,
anus; At, adipose tissue; Cd, cauda epididymis; Co, corpus epididy-
mis; Cp, caput epididymis; DD, ductus deferens; P, penis; PP, pocket-
like pouch; Pr, prostate complex; S, scrotum; Sa; Spermatic artery; T,
testis; TV, tunica vaginalis). Scale bars ¼ 0.5 cm.

160 BEGUELINI ET AL.



Fig. 2. Basic testicular and epididymal morphology of the 18 species
analyzed. Phyllostomidae: A. Artibeus lituratus. B. Artibeus planirostris.
C. Carollia perspicillata. D. Glossophaga soricina. E. Phyllostomus dis-
color. F. Platyrrhinus lineatus. Noctilionidae: G-H. Noctilio albiventris.
Molossidae: I. Eumops glaucinus. J. Molossops temminckii. K. Molos-

sus molossus. L. Molossus rufus. Emballonuridae: M. Ventral view of
the gonad in Peropteryx macrotis. N. Dorsal view of the gonad in Perop-
teryx macrotis. O. Rhynchonycteris naso. Vespertilionidae: P. Eptesi-
cus furinalis. Q. Histiotus velatus. R. Lasiurus blossevillii. S. Myotis
albescens. T. Myotis nigricans. Scale bars ¼ 0.5 cm.

REPRODUCTIVE CHARACTERISTICS OF BATS 161



and Emballonuridae (Fig. 4U–Z) and of the molossid spe-
cies E. glaucinus (Fig. 4M–N), M. molossus (Fig. 4O–P),
and M. rufus (Fig. 4Q–R) corroborated the sexual matu-
rity of their specimens and indicated that no cases of
reproductive latency was presented in these species.

The analysis of Vespertilionidae showed a different
pattern, with testicular regression and sperm retention
in the cauda epididymis in the five species analyzed. In
E. furinalis, we observed normal spermatogenesis in
three specimens, regressed testes in two specimens
(Fig. 5A) and sperm retention in the cauda epididymis
in all specimens (Fig. 5B). In H. velatus, we observed
normal spermatogenesis in two specimens, regressed
testes in four specimens (Fig. 5C) and sperm retention
in the cauda epididymis in all specimens (Fig. 5D). The

testicular epithelium was regressed in all specimens of
L. blossevillii (Fig. 5E) and M. albescens (Fig. 5G), with
sperm retention in their cauda epididymis (Fig. 5F,H,
respectively). In M. nigricans, we observed normal sper-
matogenesis in the majority of specimens, regressed
testes in five specimens (Fig. 5I) and sperm retention in
the cauda epididymis in all specimens (Fig. 5J).

Molossops temminckii also showed signs of testicular
regression; we observed normal spermatogenesis in four
specimens (Fig. 5K), testes in regression in two speci-
mens (Fig. 5M) and regressed testes in two specimens
(Fig. 5O). This pattern was similar to that observed in
the vespertilionid species; however, it was different in
that the sperm storage in the cauda epididymis was
observed only in the specimens that were active (Fig.
5L) or in regression (Fig. 5N), with regressed individuals
having only a few spermatozoa in their epididymides
(Fig. 5P).

The period in which the specimens with regressed
testes were captured varied from species to species: for
E. furinalis, the captures were in December; for H. vela-
tus, the captures occurred in July–August; for L.
blossevillii, they occurred in November; for M. albescens,
they occurred in June; for M. nigricans, they occurred in
June and November; and for M. temminckii, they
occurred in November–December.

DISCUSSION

Many species of Chiroptera share with other mamma-
lian species, some reproductive characteristics that are
observed commonly in the temperate species, such as
enter into hibernation and present a regression in the
seminiferous in which only spermatogonia and Sertoli
cells can be observed (Racey, 1974; Fuentes et al., 1991;
Lee et al., 2001; Kurohmaru et al., 2002; Lee, 2003; Lee
and Mori, 2004). Possibly to adapt to the hibernation pe-
riod, the mammals species have developed unique
characteristics, such as prolonged sperm storage in the
cauda epididymis in males and in the oviducts and/or
uterine cornua in females, asynchrony between sperma-
togenesis and the mating period and late ovulation,
fertilization and implantation in the female reproductive
tract (Anand-Kumar, 1965; Racey, 1979; Rasweiler, 1993;
Crichton and Krutzsch, 2000; Lee et al., 2001; Encarna-
ção et al., 2004; Sharifi et al., 2004, Beguelini et al.,
2009, 2011).

Although neotropical bats do not have a hibernation
period, many of their reproductive characteristics vary
interspecifically. According to Krutzsch (2000), the loca-
tion of the TEC varies interfamiliarly and
interspecifically and may also vary seasonally or daily.
According to the authors, the position can be classified
into four classes: (i) permanently abdominal; (ii) perma-
nently inguinal or scrotal; (iii) migratory, in which they
migrate (seasonally or daily) from the abdomen into the
scrotum via the inguinal canal; and (iv) external, where
they remain outside the abdominal cavity.

In the present study, we observed variation in the
positioning of the TEC among families. We observed that
Phyllostomidae and Noctilionidae have external TECs,
with their position varying from the base of the penis to
the vicinity of the inguinal canal in phyllostomids and
from the vicinity of the inguinal canal to the scrotum in
noctilionids. Emballonuridae and Molossidae presented

Fig. 3. Corporal (A) and testicular (B) weights and the gonad-so-
matic index (C) of all analyzed species. The different letters indicate
significant differences (P � 0.05).

162 BEGUELINI ET AL.



Fig. 4. General arrangement of the testicular and cauda epididy-
mis tissues stained with hematoxylin eosin. Phyllostomidae: Artibeus
lituratus (A-B); Artibeus planirostris (C-D); Carollia perspicillata (E-F);
Glossophaga soricina (G-H); Phyllostomus discolor (I-J); Platyrrhinus
lineatus (K-L). Molossidae: Eumops glaucinus (M-N); Molossus

molossus (O-P); Molossus rufus (Q-R). Noctilionidae: Noctilio albi-
ventris (S-T). Emballonuridae: Peropteryx macrotis (U-V); Rhyncho-
nycteris naso (X-Z). Note the active pattern in the testes and the
sperm storage in the cauda epididymis of all species. Scale bars ¼
20 lm.

REPRODUCTIVE CHARACTERISTICS OF BATS 163



migratory TEC, and Vespertilionidae presented perma-
nently scrotal TECs, with the scrotum enclosing only the
testes and the caput and corpus epididymis.

These data from Phyllostomidae are similar to those
described by Crichton and Krutzsch (2000) in Macro-
tus californicus and by Krutzsch and Nellis (2006) in
Brachyphylla cavernarum, but differ somewhat from
the findings of Orsi et al. (1990) in Desmodus rotun-
dus, in which the TEC can migrate within the
abdomen at will. Thus, despite the small number of
species analyzed, we propose that the preferential
localization of the Phyllostomidae TEC is external,
outside the abdomen, near the crest of the pubis, lat-
eral to the base of the penis (parapenial), or near the
vicinity of the inguinal canal, with this dislocation
subject to the control of the animal. Although the
intra-abdominal condition was not observed in this
study, it cannot be neglected in relation to other spe-

cies of this family. Similarly, the observation of the
dislocation of the TEC of N. albiventris from
the pocket-like pouch to the scrotum agrees with the
observations of Dunn (1934).

Migratory TECs in Molossidae were also observed in
Mormopterus planiceps (Krutzsch and Crichton, 1987),
Tadarida brasiliensis mexicana (Krutzsch et al., 2002),
T. condylurus, T. pumila (Happold and Happold, 1989),
and T. hindei (Crichton and Krutzsch, 2000). In all these
species and in all species studied herein, we observed
that the TEC might migrate seasonally from an intra-ab-
dominal to an external position. We also note that, as
observed in T. condylurus (Mutere, 1973; Happold and
Happold, 1989) and in the species studied herein, the
testicular position (intra-abdominal or external) did not
provoke a significant variation in sperm production, dif-
fering from T. hindei (Marshall and Corbet, 1959), which
intra-abdominal testes tend to be regressed.

Fig. 5. General arrangement of the testicular and cauda epididy-
mis tissues stained with hematoxylin eosin. Vespertilionidae: Eptesi-
cus furinalis (A-B); Histiotus velatus (C-D); Lasiurus blossevillii (E-F);
Myotis albescens (G-H); Myotis nigricans (I-J). Molossidae: Molos-
sops temminckii (K-P). Note the testicular epithelium regressed in

E. furinalis (A), H. velatus (C), L. blossevillii (E), M. albescens (G) and
M. nigricans (I) with sperm storage in the cauda epididymis (B, D, F,
H and J, respectively), and the active (K), in regression (M) and
regressed (O) testicular epithelium of M. temminckii. Scale bars ¼
20 lm.

164 BEGUELINI ET AL.



Elongation of the cauda epididymis of vespertilionid
bats was also observed in Pipistrellus kuhlii (Sharifi
et al., 2004), P. pipistrellus (Racey and Tam, 1974),
P. subflavus (Krutzsch and Crichton, 1986), Myotis dau-
bentonii (Encarnação et al., 2004), Corynorhinus
mexicanus (Le�on-Galv�an et al., 2005), and Neoromicia
nanus (Van der Merwe and Stirnemann, 2007), all of
which are hibernating vespertilionids. Thus, we conclude
that most vespertilionid bat genera already studied,
including the four nonhibernating genera analyzed
herein, present this elongation. A similar elongation of
the cauda epididymis of P. macrotis was also observed in
other species of emballonurid bats such as Taphozous
georgianus (Jolly and Blackshaw, 1988).

The variation in the positioning of the testes and
epididymis of eutherian mammals seems to be linked to
a balance between the production, capacitation, and stor-
age of spermatozoa (Bedford, 1978, 2004, 2008). The
exposure of the testes to deep body temperatures (intra-
abdominal condition) seems to directly influence the
sperm production; however, it is not incompatible with
normal spermatogenesis. Although body temperature
does not suppress sperm maturation in the epididymis,
it dramatically influences the storage capacity and the
support of sperm viability and maturation in the cauda
epididymis (Bedford, 2008).

It is known that the sperm production is directly cor-
related with the size of the testis (Short, 1997; Bedford,
2008) and to the number of germ cells nursed by each
Sertoli cell in the seminiferous epithelium (Russell and
Griswold, 1993; Hermo et al., 2010a,b). According to sev-
eral authors, the lower epididymal temperatures
facilitate sperm maturation and storage (Foldesy and
Bedford, 1982; Djakiew and Cardullo, 1986; Jolly and
Blackshaw, 1988), thereby clarifying that the positioning
of the testes and epididymis directly influences the
reproductive success of each species, with their position-
ing representing a balance between sperm production
and storage. Optimizing sperm storage may enable lower
sperm production by the testis without compromising
male fecundity (Bedford, 2008), while saving energy in
small animals that show great metabolic stress. In the
same way, it seems that the sperm storage serves to
ensure that adequate amounts of sperm are available at
the peaks of reproductive activity, mainly in species that
form harems.

The elongation of the cauda epididymis in Emballo-
nuridae and Vespertilionidae was associated with the
small size of their testes and was contrasted by the ab-
sence of elongation in others species (Molossidae,
Phyllostomidae and Noctilionidae families). The fact that
these species have larger testes seems to corroborate the
trade-off between sperm production and storage.

Our data show similarities between the phyllostomid
and noctilionid bats. This fact is not surprising because
they belong to the same superfamily and present many
other very similar features, such as characteristics of
the female reproductive system and function and of
early developmental biology (Bleier, 1975; Rasweiler,
1972, 1978; Rasweiler et al., 2011).

Male Reproductive Activity

The absence of cases of reproductive latency
(regressed testes) in the six species of Phyllostomidae

corroborated the observations of others authors: Hollis
(2005) described the presence of scrotal males and preg-
nant females of A. planirostris in many months and
predicted that this species may be able to produce young
at any time throughout the year; Costa et al. (2007)
described a similar pattern in P. lineatus; and Cloutier
and Thomas (1992), Alvarez et al. (1991), and Kwiecin-
ski (2006) observed a bimodal polyestry with great
reproductive activity during the year for the species C.
perspicillata, G. soricina, and P. discolor, respectively.
However, in contrast to the postulation by Oliveira et al.
(2009), the presence of a period of gonadal regression in
A. lituratus was not observed.

Similarly, other studies also indicated the absence of
regressed testes in E. glaucinus (Best et al., 1997) and
M. molossus (Fabi�an and Marques, 1989), but no clear
literature is available for M. rufus and N. albiventris.

Studies of the reproductive cycle of emballonurid bats
are scarce for neotropical species. Plumpton and Jones
Jr. (1992) postulated that the number of pregnancies per
R. naso female of ranges from 0 to 2 per year, and Yee
(2000) described that P. macrotis breeds during almost
the entire year. These data along with our own observa-
tions suggest that both species may have active males
during almost the entire year.

The Case of Vespertilionidae

Our data indicate the presence of a reproductive
period of quiescence (regression of the seminiferous
tubules) and sperm retention in the five species of neo-
tropical Vespertilionidae analyzed that is very similar to
that observed in hibernating bats. Other studies per-
formed at latitudes, such as those by Van der Merwe
and Rautenbach (1987) in Nycticeius schlieffenii at a
latitude of 22�S, Van der Merwe and Rautenbach (1990)
in Pipistrellus rusticus at a latitude of 24�S, and Van
der Merwe and Stirnemann (2007) in Neorimicina nanus
at a latitude of 25�S, also demonstrated a period of tes-
ticular regression in these vespertilionid bats.

These data suggest that, although the species ana-
lyzed are tropical species that do not present a
prolonged period of torpor (hibernation period), they still
maintain a period of seminiferous tubule regression and
sperm retention/storage, a trait probably retained from
the common temperate vespertilionid ancestor. The
maintenance of these traits in a tropical environment is
interesting and is worthy of further study.

The Case of M. temminckii

Although no clear data were observed for the repro-
ductive patterns of M. temminckii, Krutzsch and
Crichton (1987) and Krutzsch et al. (2002) observed a
period of testicular regression in the molossid bats Mor-
mopterus planiceps and Tadarida brasiliensis mexicana.
The former study occurred in Southeast Australia (36�S)
and the latter was in Texas (USA � 30�N). Thus, these
data indicated that the occurrence of testicular regres-
sion in molossid bats might occur in both hemispheres.
However, the fact that only one species of the four
Brazilian molossid bats analyzed presented this pattern
was intriguing and deserves future investigation.

REPRODUCTIVE CHARACTERISTICS OF BATS 165



ACKNOWLEDGEMENT

The authors are grateful to Luiz Roberto Falleiros Jun-
ior for technical assistance. The scholarships awarded to
Mateus Rodrigues Beguelini by the Brazilian Research
Foundation (CAPES) and to Cintia Cristina Isicawa
Puga and Fabiane Ferreira Martins by the São Paulo
State Research Foundation (FAPESP) are also gratefully
acknowledged.

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