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Bats of Nepal
By
Rajesh Rajchal
M. Sc. Forestry Student
Institute of Forestry
Pokhara, Nepal
Submitted to
Course Coordinator
Special Study Course
Institute of Forestry
Office of the Dean, Pokhara
February 2007
Acknowledgements
If a string were tied between all the people who made this study possible, it would make a
mess! But, to me, it would look something more like a perfect snowflake. This study is a
group endeavor and pervading this one was a recognition and thankfulness for all the people
without whom this effort could not have succeeded.
First and foremost I would like to extend my sincere gratitude to Dr. Bimal Keshari Paudyal
(M.Sc. Program Coordinator), Mr. Shree Prasad Dhoubhadel (Associate Professor), Mr. Raj
Kumar Koirala (Lecturer) and Mr. Jeev Nath Pandey (Lecturer), Institute of Forestry,
Pokhara, Nepal for supporting me to choose my ‘special study course’ in bat. Their advices
and encouragements help me to complete this study.
My sincere and cordial thanks go to Sally Walker, Administrative Chair, Chiroptera
Conservation and Information for South Asia and Geeta Shrestha (Founder, NATURE; Nepal
Country Representative for CCINSA, Kathmandu) who encouraged me to step on the study
of bats. Their constant interest and support from the very beginning have helped me to spring
up my knowledge in the field of bat and prepare this report.
Special thank certainly goes to Dr. Luis Ruedas (Assistant Professor, Department of Biology,
Portland State University, US) who taught us about bat and conducted excellent field visits to
capture bats. His advices and information were very helpful to prepare this report.
I would like to convey deep appreciation to my colleagues Mr. Sujas Phuyal, Mr. Pushpa Raj
Acharya, Mr. Narayan Prasad Koju and Ms. Reena Byanju for their cooperation and helps in
every moment during preparation of this report. They encouraged me to be devoted in bat
conservation and help to collect information relevant to bats.
In this moment, I could not forget Mr. Laxman Poudyal (Ranger, Department of National
Parks and Wildlife Conservation) for his inspiration and kind suggestions during literature
review and report preparation.
I cannot stay without thanking my friend Mr. Niroj Man Shrestha who helped me a lot during
preparation of this report. I would also like to thank Mr. Mohan K. C., Hari Adhikari, Rijan
Tamrakar, Mr. Madhav Prasad Baral, Mr. Prabhat Sapkota and Mr. Rabin Kadariya for their
kind cooperation and positive reflection.
II
Preface
Bats are one of the least known, least monitored and the most misunderstood creatures on the
earth. Some evoke revulsion, fear, suspicion and awe, sometimes even hysteria. Conservation
efforts are incomplete when they do not sufficiently focus on the relatively less known groups
of fauna and flora. Since the Hodgson era, in Nepal, there have been significant efforts to
increase the knowledge on taxonomy, biology, ecology and distribution of mammals in the
country, but information about bats is still very difficult to access.
Young bat conservationists are facing many problems including lack of information about
bats. In this regard, the report is intended as a reference for them and hopefully serves
towards a milestone in bat education. This report is based on the literatures mainly browsed
from internet and it includes two sections. First section includes general introduction on bats
which aims to spread information about bats and dispel many superstitions related to bats and
show positive light especially in terms of their importance and contribution to ecosystem
health and human well being. Second section includes list of bats of Nepal, their descriptions
with photographs and their distributions in South Asia including Nepal.
This is my first attempt on bats and I hope that this report will be useful to bat lovers and
students. I also anticipate that many improvements can be made and some errors may need
correction. I would appreciate any comments and suggestions regarding this report.
Rajesh Rajchal
M. Sc. Forestry Student
Institute of Forestry
Office of the Dean, Pokhara, Nepal
Email: [email protected]
Website: www.freewebs.com/rrajchal
Mobile: +977 9803210032
Note:
I apologize for not being able to give credit to the authors of the photographs and diagrams.
All the photographs and diagrams in this report have been downloaded from internet sites,
and I am very grateful to the authors who made them available. In addition, this report is
strictly based on literature review and I tried to include all the sources in the reference
section. Any missing of citation is unintentional and I apologize in advance.
This report is made available for the educational purpose.
III
IV
Table of Contents
ACKNOWLEDGEMENTS ...................................................................................................................................... II
PREFACE ................................................................................................................................................................. III
INTRODUCTION ...................................................................................................................................................... 1
EVOLUTION OF BATS ............................................................................................................................................ 2
GEOGRAPHIC RANGE ........................................................................................................................................... 6
HABITAT .................................................................................................................................................................... 8
SYSTEMATIC AND TAXONOMIC HISTORY..................................................................................................... 9
PHYSICAL DESCRIPTION ................................................................................................................................... 13
SEXING ..................................................................................................................................................................... 21
REPRODUCTION .................................................................................................................................................... 21
LIFESPAN ................................................................................................................................................................. 24
BEHAVIOR ............................................................................................................................................................... 25
COMMUNICATION AND PERCEPTION ........................................................................................................... 27
FOOD HABITS ......................................................................................................................................................... 28
BATS AND THEIR NEIGHBORS ......................................................................................................................... 29
HIBERNATION AND MIGRATION ..................................................................................................................... 33
PREDATORS, ENEMIES AND MEANS OF DEFENSE ..................................................................................... 34
ECOSYSTEM ROLES ............................................................................................................................................. 35
CONSERVATION .................................................................................................................................................... 36
IMPORTANCE OF BATS ....................................................................................................................................... 37
ECONOMIC IMPORTANCE FOR HUMANS: NEGATIVE ................................................................................................ 37
ECONOMIC IMPORTANCE FOR HUMANS: POSITIVE .................................................................................................. 37
THREATS ................................................................................................................................................................. 38
LEGISLATION ........................................................................................................................................................ 39
PREVIOUS STUDIES ON BATS ........................................................................................................................... 40
CHALLENGES AND PROSPECTS ....................................................................................................................... 41
BATS OF NEPAL ..................................................................................................................................................... 43
DISTRIBUTION OF BATS OF NEPAL AND SOUTH ASIA ............................................................................. 62
DICTIONARY OF TECHNICAL TERMS............................................................................................................ 71
INDEX ....................................................................................................................................................................... 73
REFERENCES.......................................................................................................................................................... 75
V
Introduction
Chiroptera (Greek, Cheiros: Hand and Pteron: Wing; spells like ki-rop-ter-ah) are the only
flying (volant) mammals in which forelimbs are modified into wings and are perhaps the
most easily recognizable group of animals. There are mammals, like flying squirrels and
flying lemurs, which glide through the air supported by parachute-like extensions of skin
from their bodies. But such a parachute does little more than prolong the squirrel’s leap and
reduce the impetus of its landing; whereas with bats there is true and sustained flight affected
by an upward and downward beat of wings.
Bats are the second-most speciose group of mammals, after rodents. The most current tally of
mammals (Wilson and Reeder, 2005) recognized 1,116 species of bats worldwide. Those
1,116 bat species represent about a fifth (20%) of the 5,418 known mammal species. Bats are
often divided into two major groups, usually given the rank of suborders, Megachiroptera and
Microchiroptera. Although these groups probably do not represent monophyletic lineages,
there are several relevant ecological differences between them. Megachiroptera includes one
family (Pteropodidae) and about 186 species. All feed primarily on plant material, either
fruit/nectar or pollen. The remaining 16 families (around 930 species) belong to
Microchiroptera (Wilson and Reeder 2005). The majority of species are insectivorous, and
insectivory is widely distributed through all microchiropteran families. However, many
microchiropterans have become specialized to eat other kinds of diets. Some bats are
carnivorous (feeding on rodents, other bats, reptiles, birds, amphibians, and even fish), many
consume fruit, some are specialized for extracting nectar from flowers, and one subfamily
(three species in the subfamily Desmodontinae) feeds on nothing but the blood of other
vertebrates. Megachiropterans and microchiropterans differ in many other ways.
Megachiropterans are found only in the Old World tropics, while microchiropterans are much
more broadly distributed. Microchiropterans use highly sophisticated echolocation for
orientation; megachiropterans orient primarily using their eyes, although members of one
genus, Rousettus, are capable of a simple form of echolocation that is not related to
echolocation in microchiropterans. Megachiropteran species control their body temperature
within a tight range of temperatures and none hibernates; many microchiropterans have labile
body temperatures, and some hibernate.
1
Evolution of Bats
Evolution of bats is poorly understood because of poor fossil records. Fossil remains found in
geological deposits show that even the earliest known bats had wings and were able to fly.
Unfortunately, none of these fossils reveal the stages by which bats attained this power of
flight.
The earliest complete fossil bats (Chiroptera) date from the early Eocene (49–53 million
years ago). The Eocene bat fauna is extremely rich, comprising hundreds of individual
specimens, belonging to 24 genera, including several spectacular forms that are preserved in
their entirety (Simmons and Geisler, 1998). The only possible bat remains from prior to the
Eocene are a few teeth from the late Palaeocene (Gingerich, 1987), but their chiropteran
affinities have been questioned (Hand et al., 1994).
Picture showing starting time of evolution of human and bat
The exceptional preservation of the soft tissues of some specimens of bats from the Messel
deposits (49 million years ago) confirms that the Eocene bats had wing membranes
(Habersetzer and Storch, 1989) and details of articulation of the shoulder and the enlarged
scapulae for attachment of flight muscles (Jepsen, 1970) leave no doubt that they were all
capable of powered flapping flight (Habersetzer and Storch, 1989; Norberg, 1989). A second
important feature of these bats is that they have enlarged cochleae relative to the size of the
skull (Novacek, 1987, 1991; Habersetzer and Storch, 1989, 1992). Modern bats with enlarged
cochleae are all capable of echolocation, while modern bats that do not echolocate, or have
reduced reliance on this form of perception, do not show the same extent of enlargement
(Henson, 1970). Since Eocene bats have enlarged cochleae, and also modified auditory
ossicles comparable with those of extant echolocating bats (Novacek, 1985), they were also
clearly capable of echolocation (Novacek, 1985, 1987, 1991; Habersetzer and Storch, 1989,
1992). By 53 million years ago, therefore, two major behavioral innovations that we currently
2
associate with bats – flight and echolocation – had already evolved. It is remarkable that in
many respects the bats that suddenly appear over the entire globe, in the Eocene are
‘completely developed’ (Habersetzer and Storch, 1989).
The problem presented by the sudden appearance of ‘completely developed’ bats in the fossil
record was recognized by Darwin (1859) in the Origin of Species. Darwin (1859) suggested
that the bats posed a problem for the theory of evolution because a credible scenario for the
evolution of a flying bat from an insectivorous terrestrial mammal, by the process of natural
selection, was too difficult to imagine. This point has been used since as an argument in favor
of special creation, e.g. Morris (1974). Despite Darwin’s suggestion that constructing a
credible scenario was too difficult, there have been many attempts to reconstruct the manner
by which the traits evolved, and their subsequent distributions in extant bats. Extant bats are
all capable of powered flapping flight, but include non-echolocating as well as echolocating
forms, in part mirroring their diverse dietary specializations.
Since there are two key behavioral traits, there are at least three scenarios for their evolution:
echolocation may have evolved first (Hill and Smith, 1984; Fenton, 1984; Fenton et al., 1995;
Arita and Fenton, 1997), flight may have evolved first (Simmons and Geisler, 1998) or the
two may have evolved in tandem (Norberg, 1985a,b, 1989; Rayner, 1991a, 1991b;
Speakman, 1993; Arita and Fenton, 1997). This minimal view assumes that at least one of the
behaviors evolved only once. Multiple origins of both traits (e.g. Pettigrew et al., 1989;
Rayner, 1991a) could lead to much more complex interrelationships.
Here the four major hypotheses concerning the evolution of flight and echolocation in bats
have been summarized. For detail, please see Speakman, 2001.
Hypothesis One: Echolocation First
The echolocation-first hypothesis (Fenton, 1984; Hill and Smith, 1984; Fenton et al., 1994;
Arita and Fenton, 1997), in common with the other hypotheses, suggests that the original
arboreal progenitor of the bats was a small nocturnal insect-eating mammal. This animal
probably used ultrasound as a method of communication (Fenton, 1984), very much like
some modern insectivora and some other small mammals such as rodents (Sales and Pye,
1974). The hypothesis suggests that in addition to gleaning insects from branches and other
surfaces, the animal would reach out from branches and capture insects that were flying past
(Jepsen,1970): called reach hunting (Speakman, 1999). This would involve some sensory
skill in predicting the trajectory of the insects in darkness. The animals may have modified
3
their ultrasound communication calls into broadband biosonar calls to enhance their
capabilities to track incoming insects. The calls would have become frequency modulated to
maximize the ranging information (Fenton, 1984). The digits of the pre-bat would have been
already elongated during the transition to the arboreal habit (Kovtun, 1989). It is
hypothesized that the arms and hands of the animal would have become even further
extended (Jepsen, 1970), allowing it to reach further outwards to capture passing insects.
Over time, the animal would develop a sophisticated echolocation system capable of tracking
incoming targets from a stationary position, and enlarged hands with webbing between the
digits, capable of capturing the passing insects. This hypothesis has not been widely accepted.
Hypothesis Two: Flight First
In the flight-first hypothesis the arboreal
nocturnal insectivorous mammal is suggested
to have first evolved gliding as a progression
from leaping between branches of the trees
(Smith, 1977; Norberg, 1985b, 1986a; Rayner,
1986; Arita and Fenton, 1997; Simmons and
Geisler, 1998). An alternative model in which
the pre-bat moved directly from perching to
hovering flight (Jepsen, 1970; Pirlot, 1977) is
not widely accepted, because hovering flight
is the most energetically expensive mode of
flying (Clark, 1977; Rayner, 1986; Winter,
1998; Winter et al., 1998). In contrast, gliding is energetically much cheaper (Baudinette and
Schmidt, 1974). An initial gliding phase would involve extension of the digits and growth of
membranes between the digits to enhance the lifting surface (Ledenev, 1989 and Smith,
1977). Gliding was gradually replaced by powered flight as this allowed the animals far more
flexibility in their locomotion, and provided energetic and other benefits in terms of increased
foraging ranges, reduced costs of transport (Schmidt, 1972 and Rayner, 1986) and reduced
predation risk (Pomeroy, 1993; Speakman, 1993). By chance, the evolution of flapping flight
may have been facilitated by the positioning of the digits to subdivide the wing membrane
(Hill and Smith, 1984). This positioning would allow differential cambering, tensioning and
folding of the membrane over its surface, which are essential aspects of flapping a wing, but
options that are unavailable in membranes that are not subdivided in this manner, such as
4
those found in some current gliders (e.g. flying squirrels: Thorington, 1984). Once powered
flight had developed there might have been some divergence in the foraging behavior of the
animals, with some of them feeding predominantly on insects taken from the substrate and
colleagues feeding predominantly on fruit.
Hypothesis Three: Tandem Development
An alternative to the flight-first hypothesis (where the leaping animal develops flight in
tandem with a sophisticated visual system, which is then swapped for a sophisticated
echolocation system) is the suggestion that the animals developed an increasingly
sophisticated echolocation system in tandem with flight. This idea was suggested by Norberg
(1985b, 1989) who proposed that the ancestral animals were small, nocturnal and
insectivorous, and used ultrasound to communicate (as in hypothesis number one). These
communication calls may have acted as a rudimentary biosonar allowing the arboreal animals
to make short leaps in darkness between branches. The length of leaps, glides and ultimately
flapping flight would have evolved in this system in parallel with the increasing power and
frequency of echolocation pulse production. This would take full advantage of the energetic
efficiency of coupling biosonar production to wing flapping (Speakman and Racey, 1991;
Rayner, 1991b; Speakman, 1993; Jones, 1993). The loudest and most costly calls would only
be required when the animals were able to produce them most efficiently. Initially, the
animals would be unable to use the echolocation to capture insects (Norberg, 1989), but as
their echolocation and flight capabilities developed, the animals would evolve aerial
hawking.
Hypothesis Four: Chiropteran Diphyly
Smith (1976, 1977) suggested that bats are diphyletic, with the megachiropterans derived
from the primates and the microchiropterans derived from another, possibly insectivoran,
ancestry. The diphyly hypothesis for the origins of flight and echolocation suggests that
microchiropterans evolved from an insectivoran ancestor in the late Cretaceous that had
rudimentary echolocation capability (Pettigrew, 1986b). Flight and echolocation in this group
are presumed to have evolved by one of the models presented above (i.e. flight first,
echolocation first or in tandem). Many millions of years later, probably around the Oligocene,
an early primate group began gliding (ultimately becoming a line which led to the
dermopterans) and some of these evolved powered flights and became the megachiropterans.
Even later than this, some of these bats independently developed novel forms of echolocation
5
(e.g. Gould, 1988). The theory is unequivocal in its suggestion that flight evolved twice and
the wing structures of Mega- and Microchiroptera are convergent.
Geographic Range
Bats are found throughout the world in tropical and temperate habitats. They are missing only
from Polar Regions and from some isolated islands. Although bats are relatively common in
temperate regions, they reach their greatest diversity in tropical forests. (Hill and Smith,
1984; Vaughan, Ryan, and Czaplewski, 2000)
Owing to their powers of flight, it is generally assumed that bats are more easily able to cross
barriers like mountains, seas and rivers which so limit the range of terrestrial mammals. Their
powers of flight should give them an advantage in establishing themselves in new territories;
but as a matter of fact there is little reason to believe that it actually does so. The majority of
bats display a tendency to live or remain under particular conditions of environment. Their
range is strictly limited. Flying Foxes, for example, are abundant on islands not 40 miles (65
km) from the African mainland, yet no single species has established itself on the African
Continent. Again, there are many different species of Flying Foxes inhabiting the chain of
islands stretching across the Indian Ocean. Yet each species keeps to its island home. No
inter-migration has been observed.
Fruit bats as a whole are confined entirely to the tropical and subtropical regions. They do not
penetrate much into evergreen rain forests. Their main area of distribution is in the tropical,
semi-evergreen, moist deciduous, and dry deciduous zones of the Peninsula. Some have
colonized the desert or thorn forest zone, where life has been made possible for them as a
result of the planting of fruit trees by human agency.
It is the same with the insectivorous bats. Like fruit bats, their range is largely limited by
physical and other factors in their surroundings. Only one species of bat, the serotine has
succeeded in establishing itself in both the eastern and western hemispheres, in crossing from
the Old World into the New. It is believed that most bats are unable to withstand the cold of
the regions about the Behring Straits, where the passage between the two Hemispheres is the
shortest, and hence have failed to spread themselves as many purely terrestrial mammals, like
the reindeer, the elk, and the glutton have done.
The various genera of insectivorous bats have a distribution more or less circumscribed by
climatic and other factors. The majority of species live in the tropics. The Sheathtails and the
False Vampires are purely tropical. The typical bats (Vespertilionidae) include the Pipistrelles
6
some of which show an adaptability to almost any conditions. Their range reaches from the
confines of the Arctic to below the Equator. Other genera are less adaptable, and are mainly
temperate or tropical in their range. Races of the common European species like the
Longeared Bat (Plecotus auritus), the Serotine (Eptesictu serotinus), and the Barbastelle
(Barbastella barbaslellus) are found in India only in the temperate levels of the Himalayas ;
while species like the Common Yellow Bat (Scotophilus heathii) and Tickell’s Bat
(Hesperoptenus tickelli) keep to the tropical zone of the Peninsula. The Himalayas and the
foothills, where both temperate and tropical conditions prevail, are a meeting ground for both
temperate and tropical species, and provide either a home or temporary shelter for a greater
variety of species than can be found anywhere else in the Indian peninsula.
From what has been written it would be seen that, despite their advantage of wings to carry
them across natural barriers, bats are largely limited in their range by conditions of climate,
temperature, and other physical factors in their surroundings.
Let us consider the bat’s reactions to its physical surroundings. How do bats react to light?
We consider them to be creatures of the dark. But many bats, both fruit-eaters and insecteaters, come out to hunt in the bright light of the evening and like other nocturnal animals,
many hunt by day in cloudy or misty weather, and some even in brilliant sunshine. Again,
some bats spend the day in roosts exposed to intense light. It would appear that bats, in
common with many nocturnal animals, have no inherent objection to light. Temperature, the
movements of their prey and other factors largely influence the time of their emergence and
the direction of their flight.
Conditions of temperature profoundly influence the life and habits of these animals. With the
majority of species, the choice of a diurnal retreat is governed largely by prevailing climatic
factors. There are species like the Flying Foxes and some insectivorous bats which sleep in
open exposed roosts. But these are exceptional, and even to these hardy species such
exposure is not without its risks. It is recorded how Flying Foxes fell to the ground panting
and many quite dead when the moist winds prevailing over Madras suddenly changed to a
hot, fierce blast. A similar incident is recorded from Ghazipur in U.P. Though inured to
exposure, these bats were unable to withstand a sudden climatic change against which there
was no protection in their exposed roosts. Except where their habits have been changed by
living near man, whose works and dwellings offer them shelter, bats may be generally
classified either as tree or as cave dwellers. Some species, particularly in cold regions,
occupy trees in summer but shift to caves or more sheltered dwellings when winter sets in.
7
This seasonal change of the diurnal retreat is less evident in tropical lands where there is less
deviation between winter and summer temperatures. Again, while many tropical species -are
habitual cave dwellers, individuals may be found at times living in caves or in trees. Caves
are however the favored retreat of the majority of our bats. Why? Because trees provide less
protection against the vagaries of climate. The temperature within the hollow of a tree or
under its leaves varies with the temperature of the surrounding air. It is different with caves.
Whatever the conditions outside, the temperature within a cave remains more or less
constant. It varies but little. Caves provide the uniform conditions of temperature that bats
find suitable. In tropical lands, rock-cut dwellings of man, or ancient tombs and temples with
their thick walls and high-domed architecture are much favored by bats, simply because they
reproduce, to some extent, the equable temperature maintained within caves. However high
the temperature of outside air it is cool within these retreats. Caves are in fact essential to the
existence of many species. The presence of caves has made possible their entry and existence
in areas where the climate would otherwise have remained a barrier. The Rousette, or Fulvous
Fruit Bat [Roiuettus leschenmdti (Desmarest)], a habitual cave dweller, has- established itself
at an altitude of 7000 ft. (2150 m.) in the Himalayas, a height at which no other Indian fruit
bat is permanently resident. This bat, like other fruit bats, is a tropical species. It is able to
exist in a temperate clime because it is a habitual cave dweller while other fruit bats live in
trees. In America it has been found that bats living in caves are able to live throughout the
winter in cold northerly latitudes from which other species migrate. Cold can profoundly
influences the habits of bats.
Habitat
Bats can be found in many terrestrial habitats below the Polar Regions. Typical habitats
include temperate and tropical forests, deserts, open fields, agricultural areas and in suburban
and urban environments. Many bats forage near freshwater streams, lakes and ponds, preying
on insects as they emerge from water. Generally, if a terrestrial habitat provides access to
sufficient roost sites and appropriate food, one or more species will be found there. Bats
generally have very specific roosting requirements, which differ among species. They may
roost in caves, crevices, trees, under logs, and even in human dwellings. Bats may also use
different types of roosts at different times. For example, a species that hibernates in a cave
during the winter may use crevices in tree holes as roosts during warmer months.
8
Systematic and Taxonomic History
Traditionally, bats have been considered a monophyletic order (Chiroptera), subdivided into
two suborders – Microchiroptera and Megachiroptera.
Kingdom: Animalia
Phylum: Chordata
Group: Vertebrata
Subphylum: Gnathostomata
Class: Mammalia
Subclass: Theria
Infraclass: Eutheria
Order: Chiroptera
Suborder: Megachiroptera and Microchiroptera
Type: Bat
In the late 1980’s and early 1990’s some researchers argued that Megachiroptera, which
consists solely of the family Pteropodidae (Old World fruit bats), is a sister group to the
primates rather than to Microchiroptera. Most of this evidence was based on similarities
between visual pathways in primates and megachiropterans. Thus, the two groups of bats
were hypothesized to have arrived at the same general body plan through convergent
evolution. This hypothesis found little subsequent support, particularly with the recent
explosion of available molecular evidence. A large body of evidence now supports the
traditional view that all bats evolved from a single, common ancestor. (Teeling et al., 2002;
Teeling et al., 2005; Van den Bussche and Hoofer, 2004; Vaughan, Ryan, and Czaplewski,
2000)
While molecular genetic data helped put one controversy regarding bat systematic to rest, it
has led to another, perhaps equally surprising hypothesis. An increasing number of molecular
studies call into question the monophyly of Microchiroptera (Teeling et al., 2002; Teeling et
al., 2005; van den Bussche and Hoofer, 2004). Instead, results indicate that some
microchiropterans (the families Rhinolophidae, Rhinopomatidae, and Megadermatidae) form
a clade that is most closely related to the family Pteropodidae, containing all megachiropteran
species. This is surprising because of the many morphological and behavioral features that
distinguish megachiropterans and microchiropterans. These results also call into question the
manner in which laryngeal echolocation, a critical mode of sensory perception in all
microchiropterans, and no megachiropterans, has evolved. Did the ancestor of all bats
echolocate, and the ability was subsequently lost in what we now recognize as
9
Megachiroptera? Or did laryngeal echolocation evolve twice in Chiroptera. Both hypotheses
are viable, given the evidence currently available. (Teeling et al., 2002; Teeling et al., 2005;
Van den Bussche and Hoofer, 2004; Vaughan, Ryan, and Czaplewski, 2000)
The clade made up of Pteropodidae and the (traditionally) microchiropteran families
Rhinolophidae, Megadermatidae, and Rhinopomatidae has been called the suborder
Yinpterochiroptera. All remaining microchiropteran bat families make up the suborder
Yangochiroptera. This taxonomic scheme remains controversial, as some molecular and
morphological evidence suggests that Microchiroptera is a monophyletic group, sister to
Megachiroptera. (Teeling et al., 2002; Teeling et al., 2005; Van den Bussche and Hoofer,
2004; Vaughan, Ryan, and Czaplewski, 2000)
Kingdom Animalia: The Animalia is the largest of the 5 kingdoms, comprised of over
1,000,000 species. This kingdom encompasses multicellular, eukaryotic organisms that have
their own means of locomotion. Included in this kingdom are birds, sponges, fish, and people.
Phylum Chordata: The phylum chordata is a group of bilaterally symmetrical animals that
have a notochord during at least one phase of development. Chordates include birds, fish,
reptiles and mammals.
Class Mammalia: Mammals are hairy, warm-blooded, milk-producing animals.
Group Eutheria: This is one of the three groups of living mammals. Eutherian mammals are
also known as placental mammals, because they give birth to live young, which are nourished
before birth in the mother’s placenta. This group includes insectivores, rodents and primates.
Order Chiroptera: These are the only flying mammals and can be described in two suborders – megachiroptera and microchiroptera.
Megachiroptera
Characters
1. It is a larger bat with a wing spread over one and a half meter though the body is only
about 30 cm in length.
2. The body is covered with brown fur.
3. The snout is long and without nose leaf. The face is like that of a fox in appearance.
4. eyes are large
5. Ears are oval and the two edges of the ear are in contact at the base.
6. Tail is absent.
10
7. The thumb and second digits are clawed.
8. Dental formula is I 2/2, c 1/1, pm 2/3, m 2/3. The molars are not tubercular but
marked with a longitudinal groove.
9. Frugivorous, food chiefly consists of figs and guava.
10. Social in habit and move about in droves of considerable size.
11. Geographical Distribution: Pteropus occurs in India, Ceylon, Australia, Africa and
Madagascar.
12. Example: Pteropus, Cynopterus, etc
Microchiroptera
Characters
13. It is small in size and the body is covered with soft fur.
14. Snout is short with or without nose leaf.
15. Pinna are large and often provided with flags, serving as tactile organs and also in
making the power of hearing more acute.
16. Eyes are small and the vision is weak as the visual rods are poorly developed.
17. Tail is included in the inter-femoral membrane provided with a distinct flap.
18. Only the thumb or first digit is clawed in the fore-limbs. Hind-limbs are weak and
have five clawed digits.
19. Molars have cusped crowns with transverse grooves.
20. Nocturnal in habit. During the day they spend their time sleeping in caves or other
dark sheltered places and come out at dusk for feeding.
21. Insectivorous, though occasionally Frugivorous or sanguivorous (blood sucking).
22. Gregarious living in colonies of thousands.
23. They produce ultra-sonic sound waves which after striking on the solid objects are
reflected back and are picked up by their ears; guiding in their flight.
24. Geographical Distribution: Bats are cosmopolitan in distribution.
11
Comparison between Megachiroptera and Microchiroptera
SN
Characters
1. Habitat
2.
Habits
3.
4.
Size
Head
5.
Pinna
6.
Clawed fingers
7.
Tail
8.
Molars
9. Distribution
10. Echolocation
11. Body temperature
12. Hibernation
13. Examples
Suborder 1: Megachiroptera
Suborder 2: Microchiroptera
Deserted area in Old World Rock crevices, hollow trees and
tropics,
abandoned buildings in both the
hemispheres,
Nocturnal, Frugivorous, often Nocturnal, mainly insectivorous,
found in great flocks, hanging gregarious or solitary, hanging
upside down by hind claws upside down with head downward.
with wings folded.
Larger bats
Smaller bats
Head fox-like, Eyes large, Eyes small, Snout short and blunt
Snout elongated without nose- with nose-leaves
leaf
Small, simple, devoid of Usually large bearing leaf-like
appendages
appendages called tragus
First and Second fingers are Only first finger (thumb) is
clawed.
clawed.
Absent or small, Free from Large, Enclosed in a large internarrow inter-femoral membrane femoral membrane
Crowns devoid of sharp cusps, Crowns have sharp cusps,
Longitudinal grooves present
Transverse grooves present
Only in the old world tropics
Much more broadly distributed
Primarily using their eyes, Highly sophisticated echolocation
although members of one
genus, Rousettus, are capable of
a simple form of echolocation
control their body temperature labile body temperatures
within a tight range of
temperatures
none hibernates
some hibernate
Fruit bats or flying foxes: Little brown bat (Myotis),
Pteropus, Cynopterus
Rhinolophus, Vespertilio, Vampire
bat (Desmodus)
12
Physical Description
Bats are unmistakable. No mammals other than bats have true wings and flight. Bat wings are
modified forelimbs, much as are bird wings, except in the case of bats the flight surface is
covered with skin and supported by four fingers, while in birds the flight surface is provided
mostly by feathers and is supported by the wrist and two digits. Like human being, it has five
fingers. The flight membrane usually extends down the sides of the body and attaches to the
hind legs. Bats also often have a tail membrane called a uropatagium. In order to
accommodate powerful flight muscles, the thoracic region of bats is quite robust. In addition
to providing power, a massive chest and shoulders maintains the center of gravity between
the wings, making flight more efficient. The opposite is true of the posterior end of the body,
which is small relative to the chest and back. The hind limbs in particular are generally short
and small, with sharp, curved claws that help bats cling to surfaces in their roost. An
important cranial characteristic for recognizing bat families is the nature of the premaxilla.
(Hill and Smith, 1984; Vaughan, Ryan, and Czaplewski, 2000)
The smallest bat in the world, and perhaps the smallest mammal in the world, is the hognosed bat (or the bumblebee bat or kitti’s bat) (Craseonycteris thonglongyai). It is found in
Thailand and weighs only 2 to 3 grams. It has a 6 inch wingspan. So it can even be fitted into
a match box. This bat is very rare. The Kitti’s hog-nosed bat forages for insects around the
canopy of bamboo and teak trees. The largest bat in the world is a megabat called the
Gigantic Flying Fox (Pteropus vampyrus). This bat has a wingspan of 5-6 feet and weight of
13
about 1.5 kg. The principle food source of the gigantic flying fox is fruit, which it chews up,
swallowing the juice, and spitting out the pulp and seeds. This seed dispersion helps to
reforest vast areas of the tropics. The suborder names, Megachiroptera and Microchiroptera,
imply that megabats are all large and microbats are all small, which is not always the case.
Size varies with each group, however, with the smallest megachiropterans weighing only 13
grams and the largest microchiropterans weighing nearly 200 grams. (Vaughan, Ryan, and
Czaplewski, 2000)
Craseonycteris thonglongyai
Pteropus vampyrus
There are several obvious morphological features that distinguish the two suborders.
Megachiropterans rely on vision to orient in the dark of night, and thus have large, prominent
eyes. All microchiropterans rely heavily on echolocation, and not vision, and generally have
small eyes. Instead most microchiropterans have large, complex pinnae (external ears),
including an enlarged tragus or antitragus. Megabats have claws on the second digits
supporting their wings (with one exception); this is never the case in microbats. Microbats
often have dentition or cheek teeth whose morphology can easily be related to dilambdodont
teeth; megabats have simplified cheek teeth. (Hill and Smith, 1984; Vaughan, Ryan, and
Czaplewski, 2000)
Wings: Their wings, as we have seen, distinguish bats from all other mammals and fit them
completely for life in the air. Let us consider their structure. The name Chiroptera given to
bats is a combination of the Greek words chieros, a hand, and pteron, a wing, describes
exactly the structural plan of a bat’s wing. A bat’s arms and hands are the framework of its
wings. They are built on the usual pattern of the vertebrate forelimb. There is the upper arm
ending at the elbow, the double-boned forearm ending at the wrist, and the hand with a thumb
and four fingers. The thumb is free, the fingers are enormously lengthened and embedded in
the leathery wing membrane to support it. Like the ribs of an umbrella, they open and close
the wing and keep it taut when expanded. The jointed finger-bones give the bat’s wing its
14
special flexibility. The facile movements of the joints adapt the wing to the twists and turns
of flight, and adjust its surface to changing currents of air. A drawing together of the fingers
reduces the wing expanse, ‘takes in sail’ so to speak and instantly checks speed and
momentum. In its flexibility, its power of controlling momentum, the wing of a bat is the
perfect flying organ devised by nature.
Source: BCI, 2003
From the wings proper the flying membranes extend to the feet and then spread between the
legs forming here what is called an inter-femoral membrane, which usually encloses the tail,
and is supported also by a spur of bone projecting from each foot. Besides the inter-femoral
membrane, there is an accessory flying membrane, the ante-brachial membrane which, rising
from the region of the neck, connects up with the humerus and forearm. Thus there is, so to
speak, a continuous and uninterrupted parachute of skin around the bat’s body. Motive power
to this parachute is supplied, as we have shown, by the forelimbs, by the arms and the hands,
which have become agents for propelling the bat’s body through the air. As further aids to
flying, the bat has a capacious thorax, which contains a remarkably large heart and lungs, and
offers space for the attachment of the great muscles which sustain the arms in flight. The
large heart and powerful lungs, and the big flight muscles give these animals great staying
power on the wing, many of them flying vast distances in quest of food. But the manner of
flight in bats varies almost as much as it does in birds.
15
Variation in the length and number of the bones of the hand and fingers, so evident in bats,
profoundly influences the shape and size, of the wing, its strength, and power of flight. Bats
with long tapering wings are the swiftest fliers. Such are the wings of our Sheath-tailed Bats
(Emballonuridae). With their long, pointed wings, their speed and grace of movement, they
look almost like swallows in flight. They are among the swiftest bats in the world. Bats with
short rounded wings are slower in movement. The Horseshoe Bats (Rhinolophus) are an
example. They emerge after dark and course slowly round trees and over hedges in search of
insects. They seldom fly high. Intermediate between these extremes are the wings of many of
the typical bats (Vespertilionidae), the Pipistrelle [Pipistrellus coromandra (Gray)] for
example. They hunt at no special level. But how marvellously buoyant is their flight! How
they twist and turn, plunge and rise, in an ever-changing irregular course! Their extreme
mobility in flight admirably displays the superiority of the bat’s wing, whose, perfection is
derived from its skeletal structure, the flexible, many-jointed mammalian hand.
Though converted into wings, the bat’s arms and hands are used, as most mammals use them,
for walking and climbing. They may even be put to such purposes as the holding of food and
the killing of prey. The clawed thumb, as we have seen, is free of the wing. It is used for
securing a hold when walking, climbing or resting. Fruit bats usually have two such claws to
their wings, one at the end of the thumb and one at the end of the first or index finger. The
additional claw must be a help in scrambling and climbing among the twigs and branches of
trees, where these bats get their food. This extra claw is distinctive of fruit bats. It is never
present on the index finger of insect-eating species. The wing is sometimes used like a hand
for holding food. When eating a large fruit, a Flying Fox may hold it in the folds of one wing
or clasp it between its wrists. This habit, suggestive of the use of the wing as a hand, is
adopted also by some insectivorous species. Some of the Horseshoe Bats on capturing an
insect too big to be dealt with in the air, alight and then kill the victim by pounding it against
the tough membrane of the wing, which is thrust against the muzzle by an inward sweep of
the arm. Some bats use their wings to strike at flying insects. The wings are, after all, arms
and hands, and are used as such in special need.
In repose the wings are usually folded along the side of the body. Horseshoes completely
envelop themselves in their wings; so enwrapped, they look almost like great cocoons.
Different again is the way Sheathtailed Bats fold their wings. Their wings are long and
tapering. To pack them away compactly, a special method of folding has been devised. With
most bats the bones of the wings fold inwards, much as our fingers do, i.e. towards the palm
16
of the hand. But with the Sheath tails the second finger folds inwards and then outwards
taking a sort of double bend. This zigzag folding reduces the length of the wing. It appears to
be adopted generally by all long-winged bats.
Legs: Compared with the great development of the fore body and arms, the hindquarters and
legs of bats are weak. The whole structure of the animal is designed to one purpose,
maximum efficiency in flight, but such efficiency has been secured only at a cost to other
forms of progress. With the limbs hobbled to each other, and their free movement
encumbered by enveloping flight membranes, the walking of a bat is reduced to an awkward
hobble. The gait is rendered yet more cumbersome by the peculiar articulation of the kneejoints, which are directed backwards instead of forwards as in other animals. The device
secures maximum spread of the flying membranes but is of little help to other movements.
Many bats are quite helpless on the ground. Others get along with some speed, proceeding
backwards and forwards or crab-wise, moving their fore- and hind limbs much as four-legged
animals do. They climb with ease, using the claw on the thumb and the claws of their toes to
secure a hold. But why the feet of some bats are so large and others so small require
explanation. It has been said that the feet of bats which live habitually in caves are usually
larger enabling them to cling better to a flattened surface. Bats do not use their legs for
holding and catching prey. The Flying Fox sometimes holds a fruit with one of its legs, not
grasping it but striking its claws into it like the prongs of a fork. The claws of the hind legs
are commonly used to clean and comb their fur or even as toothpicks. But walking and
climbing and other uses to which a bat may put its legs are merely incidental. It is a creature
of the air. The main purpose of its legs is as a means of securing a hold after landing. When
alighting a bat usually secures first hold with its hooked thumbs and then, swinging over,
grips its perch with its feet. Some species turn a somersault as they alight and take hold hind
feet first. This is what Horseshoes generally do, but they have noticeably weak thumbs!
Taking hold with its feet, the bat rests and sleeps hanging head downwards. This is the usual
attitude. But some bats cling with their thumbs and feet, the body slung like a hammock or lie
prostrate along a convenient surface. Most of the smaller bats creep into holes and crevices.
The flattened heads of some of them seem especially designed for entry into narrow
openings.
Tail: Tails are a variable quantity in bats. They may be long, short, or so small as to be
scarcely visible. The tails of fruit bats are distinctive. When these bats have a tail it is always
very short and placed beneath the interfemoral membrane, and with this membrane it usually
17
has no connection. In the Horseshoes, the tail is distinct and reaches to the end of the
interfemoral membrane in which it is embedded. The Indian False Vampires
(Megadennatidae) have scarcely a vestige of a tail. In the typical bats (Vespertilionidae) the
tail is long and enclosed in the membrane spread between the legs, but its tip is often free and
is then used as a hook when climbing. In the Sheathtails the tail perforates the interfemoral
membrane towards the middle and appears on its upper surface or reaches considerably
beyond it.
The tail when well developed serves as strut or support for the interfemoral membrane and
controls its movement, bending it inwards or upwards, stretching it full length, adjusting its
surface to movements of flight. In Sheathtailed Bats, the tail slips in and out of the membrane
as from a sheath, and some species are thus able to expand or contract this membrane or, to
use a nautical term, to shake out or take in a reef in the stern sheet. Apart from flight, the tail
and its enveloping membrane may be put to yet other uses. Curved belly wards by the
intending of the tail, the membrane, when sufficiently developed as it is in most of the typical
bats acts as a brake to flight. It may be used as a large and capacious pouch for holding prey.
An insect disabled by a blow of the wing is driven into this pouch, or cleverly ‘netted’ as it
falls in mid-air. Thrusting its head into this pouch, the bat kills its victim. This improvised
Pouch is also used by some of these bats as a cradle for reception of newborn young. When
resting, the tail and connected membrane may be extended, tucked inwards, or folded over
the back, the last is a habit common to many Horseshoes.
Teeth: Insect-eating bats usually, swoop down on smaller insects and seize and eat them on
the wing. Or the prey is eaten after alighting, which accounts for the accumulations of wing
cases and other hard parts of insects which litter the floor below the roosts of these bats. Fruit
bats also carry away fruit to eat at their roosts as a final bonne bouche. But food is not always
taken on the wing. An insect may be snapped up from a tree or bush. Hovering above its
victim the bat settles for a fraction of a second, seizes it and flies upwards again, munching
its capture. Some of the smaller fruit bats also hover over their food, biting off morsels of
fruit and eating them on the wing. The Flying Fox is too big for such agile action. It eats in
the usual resting position, hanging head downwards. All bats are voracious feeders. A small
fruit bat may eat more than its own weight of fruit at a single meal. But there is this about the
eating of fruit bats, all that is taken into the mouth does not find its way into the stomach. All
fruit bats are entirely nectar or fruit juice feeders. They live upon liquid, not upon solid food.
The pulp of the fruit is chewed, but only the juice is swallowed and the substance rejected.
18
The tough fibrous element ground into a mash lodges in the hollow of the palate and is
continually removed and ejected by the tongue. But the substance of glutinous fruits which
liquefy easily, like plantains or ripe guavas, may be swallowed. It is the same with flowers
when large, their nectar is licked up with the finely pointed tongue. Some fruit bats have
particularly long tongues. Minute flowers like those of the mango and cashew nut, favorite
food of some fruit bats, are chewed and crushed to extract their nectar. Such being their food
and way of dealing with it, the cheek teeth of fruit bats are little more than grinding mills.
The oblong molars have quite smooth, or nearly smooth, crowns divided by a deep
longitudinal groove. These grooves are channels by way of which juice crushed out of the
fruit flows into the bat’s gullet. Worked by powerful cheek muscles the fruit bat’s molar teeth
are admirably designed for their special work.
Quite different are the teeth of insect-eating bats. Their molar teeth have sharp cusps to their
crowns. Shaped somewhat like the letter ‘W’, these sharp pointed teeth are well adapted for
holding and piercing the hard-shelled bodies of beetles and other insects. But the hard parts of
an insect’s body are not eaten. The wing cases of beetles and other hard indigestible parts of
the insect’s body are cut away and ejected. The Vampires are bloodsuckers and have their
front teeth especially modified for piercing the skin of animals they feed on. The number of
teeth in different genera of bats is variable and is of great importance in distinguishing
between them.
Senses: Bats feed by night. They fly in the dark with ease and swiftness. Many display an
amazing faculty for avoiding obstacles in the dark. Sight must play its part in guiding some
bats, especially those which come out early in the evening or in the morning. But there are
many species which must depend little on vision to guide them. Bats deprived of sight were
able to fly unhampered in absolute darkness. They found their way between threads stretched
across a room without once touching them. In the same way, these blinded bats evaded
branches and twigs set in their path and came to rest on the walls of the room. Loss of sight
did not hamper their movements. What then is the faculty which enables bats to steer their
way so unerringly in the dark? Recent research has revealed their secret. It was discovered
that bats use a highly developed echo-apparatus, a radar system of their own. Supersonic
sounds emitted by them vibrate through the air and striking upon any object in their path are
deflected back and instantly ‘picked up’ by bats. These ‘warning echoes’ enable bats to locate
and evade obstacles in their course. By using an elaborate sound-detection apparatus, the
experimenters discovered that their bats sent out signal cries at the rate of 10 per second
19
before taking off. The rate increased to 30 per second when they, launched into the air, and
rose to 50 per second as ‘warning echoes’ began to be received. The increased burst of sound
strengthened the intensity and volume of the returning echoes. To emit these high cries bats
have a powerfully developed vocal apparatus. The unusual development of the muscles of the
larynx in these animals was long unexplained. Its purpose is now clearer, the transmission of
supersonic sounds. That re-echoing of sounds so emitted plays an essential part in guiding
these bats was made apparent in the experiment. Bats with their mouths closed tightly to
prevent them from giving out sounds blundered about hopelessly.
How do bats ‘pick up’ these ‘warning echoes’? The faculty of sound perception is probably
not located in any single organ. It arises from a combination of senses acting in unison and
mutually assisting one another. The most important of these is perhaps the sense of hearing. It
is especially acute in insectivorous bats. An insectivorous bat can be told from a fruit bat by
its ears. In fruit bats the margins of each ear meet at the base to form a ringed, funnel-like
opening; in insectivorous bats they do not meet at the base. Besides, many insect-eating bats
have what appears to be an additional hearing aid. This is a process called the tragus. It arises
inside the anterior margin of the ear. There is also a lobe at the base of the outer margin,
known as the anti-tragus, which sometimes attains considerable proportions. These
accessories to the ear are never seen in fruit bats. They have neither a tragus nor an antitragus. Besides, the ears of fruit bats are small; those of insect-eating oats arc frequently
large, very mobile and capable of independent movement. Under stress of excitement, they
are set into tremulous motion. The ears of these bats seem especially attuned to pick up highpitched supersonic sounds, sounds quite inaudible to the human ear. Loud noises like thunder
scarcely disturb them, but the slightest squeak puts them on the alert, so much for the sense of
hearing. Coordinated with hearing is the sense of touch. Few if any animals have so exquisite
a sense of touch as bats. This sense, delicate enough to pick up the slightest pulsation in the
air, is exercised chiefly by the flying membranes. Their wide expanse, abundantly supplied
with nerves and blood vessels, makes them extremely sensitive. Besides these membranes,
many insect-eating bats have yet another organ of perception. This is an expansion of skin
around the nose generally called the nose-leaf. The nose-leaf may be comparatively small or
simple as in the False Vampires, or large and complex as in the Horseshoes (Rhinolophus)
and the Leaf-nosed bats (Hipposideros). Its intricate folds are lined with fine sensitive hairs.
It is evidently an organ of special perception. When alarmed these facial crests, like the ears,
are thrown into tremulous movement, the bat turns its head from side to side seeking the
20
source of danger. The nose-leaf appears to be more specially the equipment of bats which are
more rigidly nocturnal in habit, and which seek their prey among trees and bushes.
Horseshoes come out after dark and hunt much in forests, flying in and out among the
branches. There seems little doubt that the highly developed nose-leaf is an organ especially
designed to aid such intricate movement. This does not mean that bats without nose-leaves
are incapable of such movement. Temporarily blinded and with its ears stopped, a Pipistrelle
not only flew around a room without hesitation, out avoided all obstacles, and skillfully
dodged all attempts to capture it with a net. As stated, it is no single sense or sense organ, but
a combination of senses, which guides these bats in the dark and helps them to find their
prey. Fruit bats are less well-equipped than insect-eating species. They have no extra aids to
hearing or sound perception. The ear of a fruit bat has no tragus and they have no nose-leaf.
Yet the smaller forms emerge well after dark and find their way about easily, though it must
be said they seldom hunt in forests but keep instead to open cultivated country or to the
margin of forests. Flying Foxes on the contrary are early on the wing and sight must largely
guide their flight. They are less successful in avoiding unexpected obstacles in the dark. So,
many of them come to grief at night on telegraph and telephone wires, evidence of
misdirected flight.
Sexing
Without actually getting hold of the bat, sexing can be a remarkably difficult task. If one can
nab the little blighter for long enough to scrutinize it, sexing is reasonably simple; if it has a
penis it’s a male. However, as it is illegal to ‘nab’ bats without a bat handler’s license in this
country, most people will not get close enough. Unfortunately, it is not always true to say
that males are larger than females or vice versa. The variation between different bat species
means that in some species females are larger than males, while in others the reverse is true
and in many species the difference in size between sexes is negligible. During, and directly
prior to, the mating season the testicles of males may descend from the scrotum making
sexing a little easier.
Reproduction
Breeding season and habits naturally vary with climatic and other conditions. In northern
latitudes, the breeding season and time when young are produced are profoundly affected by
cold and other factors. Mating systems vary among bat species. Many temperate bats mate in
the fall as they aggregate near their winter hibernacula. These bats are generally promiscuous.
21
Pteropodids also tend to have promiscuous mating systems. These bats often aggregate in
large groups in one or a few trees and mate with various nearby individuals. In many
Neotropical microchiropterans, one or two males defend small harems of females. Males
secure all matings with their harem females until other males supplant them. While most
species are either polygynous or promiscuous, there are some bats that are monogamous. The
male, female, and their offspring roost together in a family group and males may contribute
to protecting and feeding the young. Examples include Vampyrum spectrum, Lavia frons,
Hipposideros galeritus, H. beatus, Nycteris hispida, N. arge, N. nana, and some Kerivoula
species. One megachiropteran species, Hypsignathus monstrosus, has a lek mating system,
where males gather in a lekking arena to display to females, who then choose the most
desirable of mates. Courtship behavior is complex in some species, while in others, it can be
nearly nonexistent (e.g., males of some species will mate with hibernating females that barely
react to the copulation event). (Hill and Smith, 1984; Vaughan, Ryan, and Czaplewski, 2000)
A large number of bats breed seasonally but the general mating season is at the end of
autumn. Temperate species often breed before they enter hibernation while many tropical
species breed in a cycle that is linked to wet-dry seasonality. All species that are not seasonal
breeders occur in the tropics, where resources may not be as variable as in temperate regions.
The function of seasonal breeding is to coordinate reproduction with the availability of
resources to support newborn young. To this end, many species have also evolved complex
reproductive physiology including delayed ovulation, sperm storage, delayed fertilization,
delayed implantation, and embryonic diapauses. Females generally give birth to one to two
pups per litter. For example the Pipistrelles may have two pups. But in some species in the
genus Lasiurus, litter sizes may reach 3 or 4 individuals (e.g. Lasiurus borealis, L. seminolus,
and L. cinereus). (Hill and Smith, 1984; Nowak, 1991)
The sperms are active in the uterus through the winter, the ovaries remain quiescent till the
spring when hibernation ends and active life is resumed. Then the ovum is fertilized, and
gestation and the actual development of the embryo commences. Thus while mating takes
place in the autumn, the birth of the young is postponed till the spring or early summer.
Similar conditions may apply to various northern species which have colonized the temperate
levels of the Himalayas. In the Indian peninsula, some of the Pipistrelles apparently produce
their young in any month of the year. The Horseshoes, the Leaf-noses and the False Vampires
(Megaderma) produce their young mainly at the commencement of the hot weather (MarchMay), while the majority of the typical bats (Vespertilionidae) and Sheath-tails are born in
22
the second half of the year. Both for the early and the late breeders the peak periods of
production coincide with peak periods of insect activity. Horseshoes hunt their food mainly
about trees and the young produced between March and May are just in time for our main
flowering season and its corresponding abundance of insect life. For the young of other bats,
which are not especially tree hunters there is an abundance of insect food during and just after
the rains.
Hanging by her thumbs, head up and feet down, she gave birth to her young which,
squeaking lustily, passed into and cradled in her interfemoral membrane. Fully expanded and
tucked upwards the membrane formed a perfect ‘pouch’ for the reception of the offspring. In
the case of the Pipistrelle the mother hung head downwards and received her offspring in her
right wing which she held partially extended for the purpose. The process varies with
different species. To continue the story of the baby Noctule, thrusting her head into the
interfemoral pouch the mother licked her offspring clean, and then resuming her normal head
down position tucked it away under the membrane between her thigh and shoulder where it
had easy access to the nipples on her breast. Bats have two pectoral teats. The young are born
blind and are generally naked. Some newborn Horseshoes and Flying Foxes have a coating of
downy hair. All newborn bats are completely dependent on their mothers for both protection
and nourishment. This is true even in Pteropodidae, where pups are born with fur and open
eyes. Microchiropterans tend to be more altricial at birth.
The baby bat clings tightly to its mother’s body, its feet and claws buried in her fur, its mouth
holding one of her teats in a permanent grip. The great majority of bats are born with a
complete set of milk teeth, all of one pattern. There is no differentiation between incisors,
molars or canines. Their sole function is to fasten on the mother’s teats and so obtain secure
hold of her body during movements or flight. The teats, besides being a channel of
nourishment of the offspring, are a means of gripping on to the parent. Horseshoes and False
Vampires actually have ‘false teats’ for this express purpose, two long nipple-shaped
prominences springing from the region of the pubis. No milk is drawn from them. The baby
bat clings to them with its teeth when not feeding. When hungry it turns round and transfers
its hold to the true nipples on its mother’s breast. The mothers have at first no difficulty in
carrying their young about with them, but they grow rapidly and may become an embarrassment. In some species, the females leave the young at home and return at frequent intervals to
feed them. In others, the young are put together in swarms and feeding is communal, each
female feeding the first that gets at her and sometimes even two at a tune. Certain species,
23
such as Rousettes and Flying Foxes, carry their young continuously till they are able to fly. At
birth, newborn bats weigh between 10 to 30% of their mother’s weight, putting a large
energetic strain on pregnant females.
Aside from the few monogamous bat species, where males contribute to feeding and
protecting young, all parental care in bats is provided by females. Some males defend feeding
territories for their harems, thereby contributing indirectly to the survival of their young after
birth. Bats cannot fly when they are born, so young bats either remain in the roost while their
mothers forage, or cling to their mothers’ during flight. Females of many species form
maternity colonies while they are lactating and rearing young. When the young are left in the
roost as the mother forages, they cluster together to keep warm. Upon their return, mothers
and their respective infants can identify each other by their vocalizations and scent, and thus
can successfully reunite. In some species, females will communally care for young, with
"babysitters" caring for the cluster of young while their roost-mates forage.
Family life is not prolonged. Juveniles grow quickly and can usually fly within 2 to 4 weeks
of birth. As soon as they are able to fend for themselves, which is within a brief span of a few
months or weeks after birth, the young fend for themselves and lead a separate existence.
They are weaned shortly thereafter. Thus, lactation is relatively short, but metabolically
demanding. (Hill and Smith, 1984)
Lifespan
Bats live surprisingly long lives. Typically, mammalian lifespan roughly correlate with their
body size: smaller mammals live short lives, whereas larger mammals live longer lives. Bats
are the only group of mammals that does not conform to this relationship. Despite the fact
that bats are generally small mammals, many bats can live over 30 years in the wild. Where
data on longevity is available, lifespan in the wild are often recorded from 10 to 25 years.
Typically, a given species will live at least 3.5 times longer than other mammals of similar
size. (Wilkinson and South, 2002)
There are several viable hypotheses to explain longevity in bats. Hibernation and daily torpor
may restrict lifetime energy expenditure in individuals, allowing them to live longer. Lack of
predation pressure on adults may also allow bats to live long lives. For their size, bats have
low reproductive rates in a given breeding season. Typically, females give birth to only one
or two young per year, but reproduce many times over a long life. By evolving a reproductive
24
strategy that is more typical of large mammals, perhaps lifespan have evolved to match those
of large mammals as well. (Wilkinson and South, 2002)
The longest-lived bat on record is a little brown bat (Myotis lucifigus). One banded individual
was recaptured 33 years after it was originally tagged. These bats weigh only 7 grams as
adults, roughly 1/3 the size of a house mouse. Myotis lucifugus is one of the most widely
studied species worldwide; thus, it would not be surprising if other, less well-known species
live even longer. (Kurta, 1995)
Behavior
Most people would consider bats to be ‘creatures of the night’ and this is, for the most part,
an accurate surmise. There are some species – such as the Old World Fruit Bats – that venture
out to feed during daylight, but most (if not all) microbats are nocturnal. Although Fruit bats
may be active during the daylight hours, there is a noticeable crepuscular peak in activity (in
other words, fruit bats are most active during the dawn and dusk hours). Some microbats,
such as the Whiskered Bat (Myotis mystacinus), may also be seen during the day, but such
observations are rare. The behavior that unifies all of Chiroptera is flight. Bats are the only
group of mammals to have evolved powered flight (although many species glide), and only
the third vertebrate group to do so. Depending upon the size and shape of their wings relative
to their body mass, different species of bats may have different flight styles. Many species
have large, broad wings and relatively small bodies, which allows them to fly slowly but with
high maneuverability. This flight behavior is useful for chasing evasive insect prey and
maneuvering within a dense forest at night. Some species with large, broad wings can even
hover. This behavior is especially useful for bats that eat nectar or pollen from stationary
flowers. Other species have long, narrow wings, which are useful for achieving high speeds,
but which restricts maneuverability. Many of these species forage in open spaces and may be
able to fly long distances. These two wing morphologies represent the ends of a continuum,
most species have wing morphologies that fall between these extremes. (Hill and Smith,
1984; Nowak, 1991; Vaughan, Ryan, and Czaplewski, 2000)
Many bats live in groups, while some species are solitary. Often, bats roost in colonies for
some portion of the year. Living in a colony can serve many functions. For bats, one of the
main purposes of group living is to collectively conserve heat. Bats are small and have high
metabolic rates, so heat conservation is vital. Many bats hibernate during the winter and
undergo daily torpor to conserve energy. Clustering together while roosting can further
25
reduce heat loss. Some bats that roost together do so in groups of several individuals. Some
groups (e.g. Tadarida) roost in caves in groups of thousands, or even millions. Some bat
species migrate to hibernation sites or to follow a food source (flowering cacti, for example).
Most bat species are not known to defend foraging areas, but this behavior is known from
some tropical species. Territorial defense of roosting sites is also known in some species.
(Hill and Smith, 1984; Vaughan, Ryan, and Czaplewski, 2000)
Why do bats hang upside down?
Scientists from all over the world have speculated
about this. It is now believed that bats adapted to
living in caves as they evolved. In caves, there are no
branches or areas in which to sit upright, so bats were
forced to hang upside down. If bats hung by their
thumbs they would have to let go before spreading
their wings. By hanging upside down they are able to
spread their wings ready for take-off and groom
themselves. They also have a good clear view for both
seeing and hearing before flight. Another advantage to
hanging upside down is that many bats can fit
together in a tight cluster, thus conserving body heat.
A Flying Fox
This is especially beneficial to bats during colder
periods in winter when they hibernate.
Its hind limbs have rotated 180 degrees so that its knees face backwards. This rotation aids in
the bat’s ability to navigate in flight and to hang by its feet. Bats actually have specialized
tendons that hold their toes in place so that they are able to cling to their roosts without
expending any energy. In fact, bats must flex their muscles in order to let go of the roosting
surface. The tendons in their legs and feet are organized in such a way that the weight of the
bat causes the toes and claws to grip the foothold in the roost firmly, even when the animal is
asleep. These adaptations are quite helpful for a flying mammal since bats only need to let go
of the roost in order to drop into flight. Hanging upside down also provides bats with roosting
space away from predators in safe places on the ceilings of caves, in trees, and buildings that
few other animals can use because they have not evolved to hang upside down by their feet.
26
Communication and Perception
Echolocation is another signature life history strategy in bats. All microchiropterans rely
heavily on echolocation to navigate through their environment and to find food. Bats call at
frequencies that are typically higher than humans can hear. These sounds bounce off objects
and produce echoes, which bats can hear and interpret. Bat calls vary in duration and
structure. Some species use short calls (2 to 5 milliseconds) at a high rate of repetition, while
other species use longer (about 20 milliseconds), but less frequent calls. The frequency
(pitch) characteristics also vary within and among species. Differences in characteristics like
frequency and duration affect the ability of an echolocation call to produce echoes from
objects of different sizes, shapes, and at different distances. As a result, echolocation call
structure can reveal quite a bit about the ecology and foraging strategy of a bat species. (Hill
and Smith, 1984; Vaughan, Ryan, and Czaplewski, 2000)
Bats can see very well, probably better than human do at dusk, but even their eyesight needs
some light and they would be unable to find their insect prey in the dark. Human hearing
ranges from approximately 15 to 20 kHz depending on age. By emitting a series of often
quite loud ultrasounds that either sweep from a high to low frequency or vary around a
frequency, bats can distinguish objects and their prey and therefore avoid the object or catch
the insect. The biggest functional difference between vision and echolocation is that vision is
a passive mode of perception, while echolocation is an active mode of perception. Vision
typically relies on external sources of light energy. Echolocation is quite different in that the
energy provided is by the animals themselves. Because bats have tight control over what
kinds of sound they produce, bats can exhibit a high degree of control over what types of
objects they can perceive. Echolocation calls vary among species, within species, and even
within individuals. This variation in echolocation behavior reflects variation in the habitats
bats are using and the food for which they are searching. Bats can also use "passive
echolocation", detecting and locating prey based on prey-generated sounds, such as frogs
calling or the sound of a beetle walking across sand. (Hill and Smith, 1984; Vaughan, Ryan,
and Czaplewski, 2000)
Bats communicate with one another in a variety of ways. Although bats may be able to hear
and interpret the echolocation calls of other bats, there is little evidence that those calls are
used directly in communication. Bats employ a suite of communication calls, most of which
are audible to the human ear. Some species use a diverse repertoire of social calls, which can
27
be useful in intra-specific aggression, mother-infant communication, and mating behavior.
(Behr and von Helversen, 2004; Hill and Smith, 1984; Vaughan, Ryan, and Czaplewski,
2000)
Scent marks and pheromones are also important in bats, as they are in other mammals. Scent
is used to communicate reproductive status and individual or group identity. Many species
have special scent glands near their faces or their wings. One family, the sac winged bats
(Emballonuridae), are so called because of a sac on the leading edge of their wing that may
be a scent gland (Behr and von Helversen, 2004). Bats also communicate with visual
displays, often during courtship. Some species have special markings on their wings or
pelage, and engage in ritualized displays to attract mates (Behr and von Helversen, 2004; Hill
and Smith, 1984).
Food Habits
As a group, bats eat a wide variety of food types. The majority of species eat insects, either
taking them on the wing or picking them off surfaces. Species specialized for eating fruit,
nectar, or pollen are especially abundant and diverse in tropical regions. Some bats eat
vertebrates like frogs, rodents, birds, or other bats. Several species (e.g., Noctilio leporinus
and Myotis vivesi) are specialized to trawl for fish. Three species of bats, the vampire bats
subsist solely on the blood of other vertebrates. Although most stories related to mythical
‘vampires’ originated in the Old World, there are no Old World bat species that feed on
blood. Vampire bats occur only in the neotropics. Vampire bats eat blood by using their sharp
incisors to make incisions in the skin of their prey. An anticoagulant in their saliva keeps
blood flowing while they lap it up. Only one of these three species eats the blood of
mammalian prey, the common vampire bat (Desmodus rotundus). The other two species
(Diaemus youngi and Diphylla ecaudata) are specialized for feeding only on birds. Although
most bats tend to be specialized for a particular diet, most frugivorous bats also include
arthropod prey in their diet when available. At least one extant species, the unusual New
Zealand lesser short-tailed bat (Mystacina tuberculata), is omnivorous (Hill and Smith, 1984;
Nowak, 1991; Vaughan, Ryan, and Czaplewski, 2000).
The different food preferences of bats are widely distributed among families.
Megachiropterans eat only fruit and nectar, but the entire range of diets can be found among
microchiropterans. Insectivory is common in many families, and carnivore on vertebrates is
exhibited by several. The New World leaf-nosed bats (family Phyllostomidae) in particular
28
have undergone an extensive radiation in ecology and food habits. The entire range of diets
exploited by all of Chiroptera can be observed in this single family, which also includes the
only sanguivorous (blood feeding) bats (Vaughan, Ryan, and Czaplewski, 2000).
Bats and their Neighbors
Fruit Bats: Bat-plant interactions are easily observed in the tropical parts of the world. An
animal’s relationship with its neighbors is largely governed by its food. As fruit-eaters, there
is an intimate interrelationship between fruit bats and the various forms of plant life, which
provide them with food. They exert a mutual influence on each other. The distribution and
local movements of these bats, the changes in feeding grounds, and the nature of their food
are largely influenced by the seasonal flowering and fruiting of trees, which explains why
fruit bats may be abundant in a given area at one season and absent in another. The breeding
periods of these bats and the time when the young are born are again coordinated with
flowering and fruiting seasons, in other words with the abundance or scarcity of food
supplies. Except for the Rousette, a habitual cave dweller, trees again usually provide diurnal
retreats for all our fruit bats. The kind of tree selected depends much upon the locality.
Wherever palms are available, they become the favorite shelter of the short-nosed Fruit Bat
(Cynopterus sphinx). Flying Foxes usually select wide-spreading banyan trees, various
species of figs, and feathery tamarinds; while in some localities clumps of bamboo are
chosen. A particular tree or group of trees may be occupied by Flying Foxes year after year,
evidence of their essentially sedentary habit. They are loath to leave a favored locality. These
are some of the ways by which the lives and habits of fruit bats are influenced by the
surrounding plant life. What is the influence they exert on plants? By their destruction of
flowers and fruit, fruit bats become a factor in the control of plant life, but they also function
as agents in its propagation. Their habit of carrying away fruit to their distant roosts makes
them agents in seed dispersal. So great is the quantity of seed so carried that the ground under
which large colonies of Flying Foxes habitually roosted was rented annually for the right of
seed collection. Then, as flowers form an important part of the food of fruit bats, they become
fertilizing agents carrying pollen from one flower or tree to another. When drinking the
nectar, the movements of the bat cause the pollen to be shed on its snout, or head, or other
parts of its body, and so to be carried about. Many of the flowers visited by fruit bats are
species, which open only after dusk, they have a strong scent to guide bats to them and, apart
from this, their shape and structural peculiarities lend themselves especially to fertilization by
29
bats. Bat and plant have co-evolved and co-existed for millions of years in tropical regions
(Heithaus, 1982).
Insectivorous Bats: As predatory animals, the relationship of insectivorous bats with their
animate environment is largely influenced by the creatures upon which they prey. From their
voracity and the myriads of insects they must destroy they are perhaps Nature’s most
important check on nocturnal or crepuscular insect life. Almost any insect that can be caught
is food for these bats. But, because of differences in the levels at which they hunt and
differences in feeding grounds, there is some differentiation in prey. Highflying bats like the
Sheathtails are brought more into contact with highflying insects. The smaller less powerfully
winged species prey largely on moths, flies, and insects, which keep to lower levels. Then
again, some bats, many Horseshoes for example, hunt mainly in hill forests; others keep to
open country, cultivation, or the neighborhood of human dwellings; and others again hunt
habitually over water and become factors in the control of aquatic insect life. As with other
predatory animals, these bats follow the movements of their prey. The seasonal abundance or
scarcity of insects in a given area may lead to a change in hunting grounds. Tanks drying up
in hot weather temporarily become the hunting ground of many insectivorous bats, which
swoop on the swarms of insects hovering round the decaying vegetation. Flowering trees,
attracting insects, also attract bats, which prey upon them. The height at which bats fly and
the time of their emergence are again largely influenced by the movements of the insects
upon which they feed. Insectivorous bats, particularly cave-dwelling species, indirectly
become providers of food and shelter to their neighbours. The accumulations of dung and
insect remains in caves and similar retreats where they roost attract and offer sustenance and
shelter to many insects and small creatures.
Carnivorous Bat: While the majority of bat functions as a potent check on insect life, there
are those specialized species, the Vampires, which have assumed a different role, and have
taken to preying on other animals. The true Vampires are found only in the tropical forests of
the New World. There is no authentic record of these bats ever attacking human beings, a
habit developed by some of their American counterparts.
Bats and Man: Man influences the lives and habits of these animals mainly as a provider of
food and shelter. Fruit bats are habitual raiders of our plantations. They feed on almost every
kind of cultivated fruit, and upon the flowers of such trees as the mango and the cashew nut.
Their migrations and seasonal movements are influenced, very considerably by the
abundance or scarcity of food supplies made available by man, and their incursion and
30
establishment in the arid treeless parts of India has been made possible only as a result of
irrigation and the planting of fruit trees by human agency. Indirectly man also becomes a
provider of food for insect-eating bats. Many species take up their abode in or near human
dwelling or in cultivated areas. Human dwellings and cultivation attract their attendant hordes
of insects, and the insects attract their attendant armies of bats.
The roosting habits of bats also have been considerably modified by association with man.
Many species have taken to living in houses and other retreats provided by him. And in the
absence of natural caves, cavernous tombs and temples built, by man offer accommodation to
cave-dwelling species. Such man-made shelters have undoubtedly aided the establishment
and spread of species into areas in which, but for such shelters, they could not have existed.
Insect-eating bats, by their destruction of myriads of crop and other pests, must play a useful
role in human economy; the same cannot perhaps be said of fruit bats. Such good as they may
do in the spread of plant life is largely offset by their enormous damage to human food
resources. Yet bats are rarely molested by man. The flesh of some species, the Flying Fox in
particular, is sometimes eaten. It is said to have a line and delicate flavor. What then are the
enemies of bats? How are their numbers controlled?
Color as a Means of Protection: Generally speaking, most people tend to think of bats as
being either brown or black in color – interestingly, some people I spoke to were even
unaware that bats are covered in fur. The color of this fur varies between species, from the
more typical brown or black -- usually paler underneath -- to a pale orange in the Schneider’s
Leaf-nosed Bat (Hipposideros speoris). Certain bats fall in between this, displaying colored
patches, stripes or frosted tips to their fur. The sombre coloration of most bats, so much in
keeping with the darkness of their diurnal retreats must have its protective value. This applies
even to such brightly colored species as the Painted Bat (Kerivoula picta). Despite its bright
orange fur and startling vermilion and black wings, the bat is most difficult to detect, its color
so matches the deep reds and yellows of fading leaves, a favorite roost. On a different plane
is the coloring of Flying Foxes. Here there is no concealment or camouflage. The bright
golden coloring of the bat’s head and neck glows in contrast with the sombre tones of its
body and wings. Far from concealing, the bold color pattern attracts attention. One may class
these bats with those animals .whose coloration is known as ‘warning coloration’. The
meaning of ‘warning colors’ has been explained. It will be seen that animals which exhibit
warning colors always have some repellent means of defense, e.g. they possess stink-glands
and use nauseous fumes and discharges to divert attack. Also such boldly-colored animals do
31
not fear exposure. These attributes are seen to some extent in Flying Foxes. All the species
have so strong an odour that any of their enemies, man or animal, can easily detect them with
closed eyes! They do not fear exposure, make no attempt at concealment, but sleep in open
roosts, exposed to the gaze of all who pass by.
Interrelationships between Bats
Food is again one of the most important factors governing such interrelationship. The
influence of numbers on these animals, the relationship between population and food
supplies, and the control exerted by this factor on overpopulation arc all matters which
remain to be investigated. Flying Foxes congregate in suitable areas in teeming numbers.
With their capacity for feeding one can well imagine the quantity of food required to support
such large populations.
Among insectivorous bats, equally voracious in habit, some division of food supply is
brought about by differences in hunting habits, and the surroundings or the strata of the air in
which different species customarily hunt their prey. There is also a division of territory
among individuals of many species. Individual bats may have their own ‘beats’ which they
guard against intrusion by wandering members of their own species. Even a paddy field may
be divided into three or four territories each bat keeping more or less strictly within its own
territory. Should a stranger intrude it is at once driven off. Pipistrellus (P. coromandra) also
have their special beats. An individual may be seen keeping to the same round evening after
evening, occasionally varying it by sporting in mid-air with comrades of adjacent beats.
Many Horseshoe Bats also display this habit.
Social Life: Most bats are gregarious. Individuals of a species tend to live together in large or
small colonies. The rule is not invariable. A number of Indian bats are commonly found
living alone or in pairs. But many species congregate in vast colonies. There is no apparent
social organization in these colonies. In the common roost, the individual exhibits some
toleration for the presence of others. This tolerance appears to be the limit of social life.
There is no prolonged association between parents and young, no sustained family life, no
organization or leadership. Each individual fends for itself. Members of a colony of Flying
Foxes leave their roosting trees at about the same time, fly in the same general direction, but
each bat steers its own course. Arrived at the feeding tree each bat does its best to get the
better of its neighbor. There is incessant wrangling. Late comers endeavour to dislodge
earlier arrivals from good feeding places. It is the same when they return to their roosts. They
32
fight for sleeping places, snap viciously at each other, or strike out with the great claws on
their thumbs, shrieking and cackling incessantly. The Rousettes also are highly gregarious.
Thousands roost together in caves and tunnels. The smaller fruit bats (Cynopterus, etc.) do
not assemble in such numbers. The colonies are comparatively small.
It is the same with insect-eating species. The Painted Bat and some of the Horseshoe and
other species live singly or in pairs, Pipistrellus assemble in comparatively, small numbers.
But many species, such as some of the Sheathtails etc, congregate in vast colonies. Different
species occupying the same roost generally roost apart, and in such roosts, individuals keep
their distance. Sometimes different kinds of bats share the same cave or retreat. But even then
there is no intermixing. Each species keeps to its own quarter. The ‘hide-outs’ of Vampires
are usually shunned by smaller bats. Their predatory habits make them unwelcome
neighbours. Apart from sex, the causes, which lead to the congregation or segregation of bats,
remain obscure. As to means of communication bats are rarely silent, and probably have a
keen perception of the voices of their fellows. Young bats call to their mothers as persistently
as the young of any other animal.
Hibernation and Migration
In northern countries, the majority of bats hibernate in winter. With the lowering of all
temperature, all body functions cease and the bats fall into a lethargic sleep. But even in the
depth of winter a warm spell restores them to activity. Their winter sleep is never very
profound. Hibernation enables animals to tide over the scarcity of food, which prevails
wherever winter conditions are severe. In winter, there are few insects about. The numbing
effect of cold on insects is well known. Faced with a shortage of food supply, insect-eating
bats either fall into a lethargic sleep, a condition in which they require no food, or migrate to
warmer climes where food is abundant. Little is known of the hibernation or migration of bats
in India. But such records as are available show that various European species, amongst these
the Barbastelle, the Long-eared Bat, and the Noctule (Nyctalta noctula), which have
established themselves in the higher levels of the Himalayas follow the inherent habits of
their kind and hibernate in winter. Whether any of the European forms migrate from the
Himalayas in winter to warmer southern latitudes is not known. But apparently various
tropical species which visit the temperate levels off the Himalayas in, spring and summer
leave this zone on the approach of winter. The Indian Pipistrelle, common at Simla at other
times of the year, completely disappears in winter. The Mustachioed Bat (Myotis
33
mystacinus), a widely distributed Oriental species, also disappears from the hill-station in
winter and docs not return till the spring. So much for the Himalayas. Bats living in tropical
zones of the Peninsula naturally do not hibernate, but even these animals are reduced to
temporary inactivity under severe and unaccustomed cold. During an unusually cold, Flying
Foxes were found keeping to their roosts through the night. On the other hand extreme heat
has been known to cause the Indian Flying Fox great discomfort. Rain, unless it is heavy,
seems to make little impression on bats in migration and hibernation. Many come out to hunt
on wet evenings and judging by the way they roost in caves and culverts, which are almost
waterlogged, many species appear to be little affected by excessive damp. High winds may
influence the height at which bats fly and, when these prevail, even strong-winged species
like our Sheathtails come down to hunt at more sheltered levels.
Predators, Enemies and Means of Defense
Little is known of the natural enemies of bats. In their exposed roosts, Flying Foxes ought to
become an easy prey to predatory animals, but there is little evidence of their being attacked.
Bats are known to fall victim to a wide variety of predators such as large spiders, snakes,
large lizard, birds of prey (Falconiformes and Strigiformes), mammals and some carnivorous
bats (Hill and Smith 1984, Mickleburgh et al., 1992). Bats are probably most vulnerable to
predators as they roost during the day or emerge in large groups in the early evening.
Predators like snakes or hawks often wait near the entrances of caves at dusk, attacking bats
as they leave the roost. Juvenile bats that cannot yet fly are also at risk of predation if they
fall to the ground. Individual bats flying in the dark of night are probably difficult to catch,
even for owls, which can fly and locate prey well in the dark. Several species of bat have
become specialized for preying on other bats, these include the New World species
Vampyrum spectrum and Chrotopterus auritus, and two Old World species in the genus
Megaderma. (Hill and Smith, 1984). Bats generally avoid predation by staying in protected
roosts during the day and through agile flight at night. Most bats are also cryptically colored.
The sombre coloration of most bats, so much in keeping with the darkness of their diurnal
retreats must have its protective value. This applies even to such brightly colored species as
the Painted Bat (Kerivoula picta). Despite its bright orange fur and startling vermilion and
black wings the bat is most difficult to detect, its color so matches the deep reds and yellows
of fading leaves, a favorite roost. On a different plane is the coloring of Flying Foxes. Here
there is no concealment or camouflage. The bright golden coloring of the bat’s head and neck
glows in contrast with the sombre tones of its body and wings. Far from concealing, the bold
34
color pattern attracts attention. One may class these bats with those animals whose coloration
is known as warning coloration. It will be seen that animals which exhibit warning colors
always have some repellent means of defense, e.g. they possess stink glands and use
nauseous fumes and discharges to divert attack. Also such boldly colored animals do not fear
exposure. These attributes are seen to some extent in Flying Foxes. All the species have so
strong an odour that any of their enemies, man or animal, can easily detect them with closed
eyes. They do not fear exposure, make no attempt at concealment, but sleep in open roosts,
exposed to the gaze of all who pass by. The real enemies of bats are the various parasitic
flies, and to a lesser extent fleas and mites which feed on the fur, membranes, and blood of
these animals. Various species of bats appear to have their particular insect parasites. To rid
themselves of these irritating pests bats constantly comb their fur, using the claws of their
feet, and even their teeth where possible.
Ecosystem Roles
Bats are not the most popular animals in the world! Many people associate bats with bad
luck, with disease, with witches, evil, vampires, rabies or at least crop raiding of fruiting
trees. In fact bats do some damage to crops by eating the fruits but they probably make up
for it by dispersing seed. Insect eating bats consume an enormous number of insects that are
harmful to crops. When speaking of bats, then, their contribution to mankind’s well being is
both positive as well as negative.
Because of their high metabolic needs and diverse diets, bats can impact the communities in
which they live in a variety of important ways. They are important pollinators and seed
dispersers, particularly in tropical communities. Also, carnivorous and insectivorous bats may
significantly limit their prey populations. Bats may be keystone species in many
communities, particularly in the tropics where they are most abundant and diverse. (Hill and
Smith, 1984; Jones, Purvis, and Gittleman, 2003; Nowak, 1991; Vaughan, Ryan, and
Czaplewski, 2000)
Bats are associated with many kinds of internal and external parasites. They are known to
harbor several protozoans that cause malaria (e.g., Plasmodium, Hepatocystis, Nycteria and
Polychromophilus) although none of the malarial parasites found in bats cause malaria in
humans. Trypanosome protozoans, that may cause a variety of diseases, such as sleeping
sickness, are also found in a number of bat species. Many flatworms (Cestoda and
Trematoda) and roundworms (Nematoda) spend at least part of their life cycle within the
35
tissues of bat hosts. Bats commonly harbor external, arthropod parasites. Ticks, mites and
insects such as true bugs and fleas are known to live and feed on bats. An entire family of
flies, Streblidae, has co-evolved with bats. These flies have secondarily lost the ability to fly,
living only in the fur of bats. Species that parasitize bats exhibit a range of host-specificity:
some are found on one or a few bats, others occur on a wider variety of bat species, and still
others can parasitize bats as well as other taxonomic groups. (Hill and Smith, 1984)
Conservation
Approximately 25% of all species within Chiroptera (nearly 240 species) are considered
threatened by the International Union for the Conservation of Nature (IUCN). At least twelve
species have gone extinct in recent times. Megachiropterans tend to be more at risk than
microchiropterans (34% and 22% of species, respectively), but both groups are facing
substantial threats from habitat loss and fragmentation. Destruction of, or disturbances to,
roost sites is particularly problematic for bats. Pesticide use also indirectly harms bats that eat
insects or plant products that have been chemically treated. Species with relatively small
geographic ranges and/or that are ecologically specialized tend to be at greatest risk. (Jones,
Purvis, and Gittleman, 2003)
In recent years, the general public has become increasingly aware of the beneficial roles that
bats play in ecosystems and their unique and amazing life histories. A wealth of research now
demonstrates that bats are a vital component of many ecosystems and an important resource
for humans. Efforts to protect bats have increased. For example, many caves that serve as
large hibernacula are fixed with gates that allow access by bats, but not by humans. Rather
than trying to eradicate bats from homes and neighborhoods, many people are placing bat
houses in their yards to give bats appropriate roosting habitat. In the United Kingdom, all bats
and bat roosts are protected by law. Several large roost emergences, including evening
emergences from a roost under the Congress Avenue Bridge in Austin, Texas, draw millions
of tourists each year. Conservation organizations like Bat Conservation International
(www.batcon.org) have growing memberships among the general public and run many
successful bat conservation projects, including projects in the developing world designed to
increase awareness and appreciation. (Bat Conservation International, 2004; Fenton, 1997)
36
Importance of Bats
Economic Importance for Humans: Negative
Although bats are often perceived as much more of a threat to human interests than they
actually are, bats may negatively impact humans in at least two ways. Some species roost in
human dwellings and can become a nuisance. This is particularly true if a large colony takes
up residence in a home, producing a great deal of guano and an unpleasant odor. Bats also
carry and transmit rabies. In general, bats rarely transmit rabies to other species, including
humans and domestic animals. Vampire bats, on the other hand, regularly transmit the disease
to domestic cattle, representing a large financial burden for the cattle industry in the New
World tropics. Rabies is transmitted through saliva and other body fluids and vampire bats
exhibit several behaviors, which make them especially effective vectors of the disease (e.g.,
social grooming and food sharing). Their feeding habits result in their saliva contacting the
blood of other animals, which is an ideal situation for rabies transmission (Hill and Smith,
1984; Vaughan, Ryan, and Czaplewski, 2000). Although bats are extremely beneficial and
help maintain balance in our delicate ecosystem, there are times when bats become a problem
or nuisance to home or business owners. Bat colonies living in human structures can be noisy
and sometimes create unpleasant droppings or odors. Bats living in buildings do not cause
structural damage nor do they chew on wires or wood. Chemical toxicants should never be
used to solve bat problems. They are unnecessary and may create far worse problems since
the chemicals may be dangerous to humans and their use may cause poisoned bats to fall to
the ground where they die slowly and are more likely to come into contact with children or
pets.
Economic Importance for Humans: Positive
Although many people consider bats to be harmful pests, bats play pivotal roles in ecological
communities and benefit humans in numerous ways. Many species of insectivorous bats prey
heavily on insects that transmit diseases or are crop pests. In addition, bat guano (feces) is
often used to fertilize crops. Many tons of guano are mined each year from caves where bats
aggregate in large numbers. In other words, some species eat crop pests and excrete crop
fertilizer! Evidence continues to accumulate in support of the immense economic benefit of
insectivorous bats for the agricultural industries worldwide. Frugivorous bats are important
seed dispersers, helping promote the diversity of fruiting trees in the tropics. Bats that eat
pollen and nectar are important pollinators, and some plants they pollinate are economically
37
important to humans, such as Agave and bananas (Musa). Larger bats, such as pteropodids
are sometimes eaten by humans. (Hill and Smith, 1984; Nowak, 1991)
Recently, common vampire bats have become an important focus of medical research.
Vampire bats are generally considered a significant threat to human interests because they
regularly transmit rabies to cattle (and sometimes to people). However, the anticoagulant
protein in their saliva (Desmoteplase) is being studied in an effort to help prevent blood clots
in humans, such as those being treated for stroke (Reddrop et al., 2005). The increasing
popularity of bats has led to a booming ecotourism industry, often surrounding large roost
emergences, such as those of Mexican free-tailed bats. (BCl, 2004)
Threats
Scientists say almost half the world’s bat species face some degree of threat to their survival.
They say bat populations in many countries are experiencing alarming declines. Threats are
being caused mainly by humans and the greatest threat to bat populations is habitat loss. Bats
produce only one pup a year so their number
grows slowly. Loss of food supply is another
threat especially to the fruit bats. Natural disasters
(flood, fires) as well as man made development
like electric wires, pollution, pesticides, chemical
toxicants, industries and vehicles are threatening
them. People also hunt and trade for food and
medicines. Chepang (A tribal group) of Chitwan,
Nepal is known to eat meat of bat as a very
special food. Some people vandalize their roosts
or needlessly kill bats out of ignorance or fear and
disturb the roosting bats in trees, caves, buildings
Threat to Bats by Electrification
and the hibernating bats. Visiting caves for fun
with light is also disturbs the bats and decline of bat species in the caves has been observed in
several cases. Some people sealing off caves so that bats cannot enter inside. Moreover,
several animals, including owls, hawks, raccoons, skunks, and snakes prey on bats; yet,
relatively few animals consume bats as a regular part of their diet.
38
Legislation
Bats are protected in many parts of the world due to their demonstrated ecological value.
Ironically, temperate countries of the western world (Europe, U.K., and USA) are far more
organized in this aspect than the tropics, where the diversity of Chiropteran species is
enormous. In the tropics, Australian legislation includes bats in strong protective legislation.
Mexico has strong legislation including full protection of caves partly because of their role as
bat habitats. (Hutson et al, 2001).
Anthony Hutson, Co-Chair of the IUCN SSC Chiroptera Specialist Group comments that
many countries are currently updating wildlife legislation because of the Rio Convention. In
this regard is not sensible to sign the Convention of Biological Diversity and continue to treat
pollinating and seed disbursing animals as ‘noxious’. CITES regulations regarding Pteropus
and Acerodon also have influenced some states to introduce protective legislation for bats
(Hutson, 2001).
In South East Asia, as mentioned previously, the Malaysia Parliament in Borneo has
protected all bats under a Wild Life Protection Ordinance (1998) which requires a license for
domestic possession of bats or any part or derivative. A more recent Malaysian law requires a
license for the sale and use of all mist nets with severe penalties for their sale and use. In fact,
peninsular Malaysia has included fruit bats in legislation for control of hunting since 1972
and there is similar legislation for other states in Malaysia. Other South East Asian states are
not so specific.
Legislation for Chiroptera is a vexed subject in the region of South Asia, particularly in India
where fruit bats are caught and eaten as food by some local people, and used for medicines to
cure headache and female ailments by others. They can be trapped for zoos or laboratory
work in any numbers with impunity. Fruit bats are killed in great numbers from time to time
due to what is considered their nuisance value to farmers when they damage fruit orchards.
Participants in the C.A.M.P. workshop recorded 11 species of Microchiroptera 8 species of
Megachiroptera were hunted for food or medicinal use in India, Nepal, Sri Lanka and
Myanmar.
Species
Microchiroptera
Megachiroptera
Bats hunted for
Food
Medicine
5
6
6
2
39
No South Asian country protects bats in principle. Sri Lankan legislation gives full protection
to one species, Rousettus leschenaulti seminudus. Other countries, such as Pakistan go to the
other extreme of exempting bats from wildlife legislation. Bats are exempted from the
regulation of international trade in Pakistan and the Punjab excludes Pteropus giganteus from
protection.
In India, fruit bats are listed as a group on Schedule V of the Indian Wildlife (Protection) Act,
which is the only Schedule that carries no penalty or restriction at all for the killing or capture
of bats, crows, mice, and rats. Several appeals to the government have been made over the
years to remove fruit bats from Schedule V considering their ecological utility, including a
recommendation in a study done from 1985-1989 under the auspices of the Wildlife Institute
of India, the premier wildlife institution in the country, which is attached to the Ministry of
Environment. These recommendations have not been taken up, but this year for the first time
the Ministry for Environment amended the Wildlife (Protection) Act to include two Critically
endangered bats, one of them a fruit bat, on Schedule I, providing a high level of protection.
This will, perhaps, create a window for delisting of fruit bats from Schedule V and upgrading
to a schedule, which will provide some protection.
Insectivorous bats are not listed in any schedule and are similarly persecuted if they prove to
be a nuisance to human beings. They settle in temples and in the eaves of houses and in
deserted structures where they are driven out by various means without regard for their value
in nature. As insectivorous bats are not listed anywhere in the Wildlife (Protection) Act,
1972, the only circumstance under which anyone can be charged and prosecuted for harming
them is within a Protected Area, where every living creature comes under the protection of
the Chief Wildlife Warden of the state.
Legislation cannot be implemented if officials whose duty is to uphold the law are not aware
of its implications. Many foresters not only do not know that bats are so essential to healthy
ecosystems; they are unaware of the number and variety of species. Forest officers were
shocked to hear that a workshop would be conducted for 130 species of bats as they were
familiar with only one or two.
Previous Studies on Bats
In Nepal, very little work has been done on bats and information regarding this species is
very limited. Some occasional studies can be found by students of Tribhuvan University and
some data found are based on the few opportunistic collection and museum specimens. B. H.
40
Hodgson, who collected 373 mammal species, belonging to 70 genera and 114 species, was
the first collector who described the fauna in this country. Hence, he contributed some to the
understanding of Nepalese bats. Abe (1982) has recorded about 570 terrestrial small
mammals, consisting of 28 species from central Nepal and contributed in identification of
several bat species (Phuyal, 2005). First legal paper – Biodiversity Profile Project – was
published in 1995 in which 37 bat species was considered to occur in Nepal. Bates and
Harrison (1997) indicated that 47 species have been recorded from Nepal whereas Hutson et
al. (2001) reported 51, of which 39 species recently remained in various categories of threat
(IUCN 2006), including in descending order of seriousness of threat category: 2 vulnerable; 6
near threatened; 29 least concern; and 2 data deficient, including one species possibly
endangered to Nepal.
Malla (2000) studied the diet analysis of Hipposideros armiger and Rhinolophus pusillus
(Microchiroptera) of Nagarjuna Cave using stomach content analysis to find out the food
items. Phuyal (2005) surveyed the bats of Pokhara valley and documented a total of twelve
bat roosting sites and eleven species. Acharya (2006) studied the distribution of roosting and
survival threats of bat in Pokhara valley with reference to species and population survey at
Bat Cave, Pokhara and found that the cave is roosting site for more than 3,000 (13.75 per sq
m at winter and 0.21 per sq m at summer) and he identified two bat species viz. Rhinolophus
pussilus and Hipposideros armiger in the cave. Shrestha (1997) has described 35 bat species
in Nepal and more extensive and detail information has been described by Molur et al. (2002)
(available in www.southasiantaxa.org) where information of all the species and their
distribution map have been given.
Challenges and Prospects
Not surprisingly, ‘additional studies are needed’ is the type of phrase often heard in
connection with bat fatalities. Decline of bat species have been reported in almost every
report but the issue of bats has not been well explored. Bat is not so charismatic to see and
people have many miss-mythical concepts towards bats. Nationally it is not getting protection
initiatives till now and most of the conservationists of Nepal are worried towards larger
endangered mammals. The study about bats found in Nepal seems like occasional studies and
documentation is very poor. Paucity of information regarding status, distribution, systematic
taxonomy and ecology is still troubleshooting the young bat conservationists. The
documentation of bats in Nepal is poorly documented. Proper technical guidance and finance
41
support to study are also the main constrains in the conservation of bats in Nepal. BPP (1995)
noted 37 species in Nepal but descriptions of all the species have not been given. Most of the
Nepali names given were the same (Chamero) which bring confusion in the identification.
Bates and Harrison (1997) mentioned that 21 more species may probably found in Nepal but
the study has not been yet started. Malur et al. reported 51 species but the status of one
species is still in question mark and their detail descriptions, geographic distribution and
identification techniques have not been given. These confounding and unclear data need to be
solved. Moreover, bats are vanishing at an alarming rate and village people are killing bat
intentionally or accidentally around Nepal. Roosts are being destroyed or disturbed with
regularity, foraging habitats are being lost, large numbers of bats are being consumed for
food and many have died from pesticide poisoning. Many species may vanish before their
decline is even recognized.
Optimistically, these problems commence prospects in the field of bat conservation. As
limited work has been done in Nepal, it opens the door for new bat conservationists to live in
all balanced and harmonious relationship with bat and all its elements with a new theme in a
new journey towards new horizons, new hopes and new success. We all literate people can
become involved in trying to save and embark conservation education to people at large.
Action by both governmental and non-governmental organizations of Nepal is vital for
promoting the protection of bats in this country. This is particularly true with bats because of
the fact that so many people still harbor misconceptions about them. Scientists, planners and
developers of Nepal can directly influence bat conservation efforts by becoming active
members of environmental protection clubs/organizations, which cannot survive without an
influx of new members.
42
Bats of Nepal
According to BPP (1995), 37 species were documented in Nepal. Bates and Harrison (1997)
recorded 47 species in Nepal whereas Malur et al. (2002) reported 51 including threat
categories: 5 vulnerable, 2 critically endangered, 5 data deficient, 1 endangered, 17 least
concern and 20 near threatened. To the date, 51 species of bats are known to live and breed in
Nepal, that comprise 41% of the South Asian and 5% of the global bat fauna. According to
Shrestha (1997), greatest diversity of the bats are found in central and western Nepal –
Pokhara valley, Rupse Chahara (Baglung), Dumja, Bardia and Baitadi. Here, names and
status of 51 species of bats have been listed and their English and Nepali names have been
given. The descriptions of the species and photographs have been taken from different
sources
especially
www.zmmu.msu.ru,
www.mammalogy.org,
www.ngensis.com,
www.fieldmuseum.org,
www.animaldiversity.ummz.umich.edu,
www.batthai.com,
www.nara-edu.ac.jp,
www.news.bbc.co.uk,
www.parchilagomaggoire.it,
www.univ-
lehavre.fr, www.dinets.travel.ru. The distribution map of the species has been taken from
Molur et al. 2002 from the site www.southasiantaxa.org.
CR, 2, 4%
VU, 5, 10%
DD, 5, 10%
CR Critically Endangered
EN, 1, 2%
DD Data Deficient
EN Endangered
LC Least Concern
NT Near Threatened
LC, 17, 34%
VU Vulnerable
NT, 20, 40%
Figure showing the status of bats of Nepal
43
Check list of Bats found in Nepal
SN
Scientific Name
1. Areilulus circumdatus
Temminck, 1840
2. Barbastella leucomelas
Cretschmar, 1830/31
3. Cynopterus sphinx Valh, 1797
4. Eptesicus gobiensis
Bobrinski,1926
5. Eptesicus serotinus Schreber,
1774
6. Hesperoptenus tickelli Blyth
1851
7. Hipposideros pomona Andersen,
1918
8. Hipposideros armiger Hodgson,
1835
9. Hipposideros cineraceus Blyth,
1853
10. Hipposideros fulvus Gray, 1838
11. Kerivoula picta Pallas, 1767
12. la io Thomas, 1902
13. Megaderma lyra E. Geoffroy,
1810
14. Miniopterus pusillus Dobson,
1876
15. Miniopterus schreibersi Kuhl,
1819
16. Murina aurata Milne-Edwards,
1872
17. Murina huttonii Peters, 1872
18. Murina leucogaster MilneEdwards,1872
19. Myotis blythii Tomes, 1857
20. Myotis csorbai Topal, 1997
21.
22.
23.
24.
25.
26.
Myotis formosus Hodgson, 1835
Myotis longipes Dobson, 1873
Myotis muricola Gray, 1846
Myotis mystacinus Kuhl, 1819
Myotis sicarius Thomas, 1915
Myotis siligorensis Horsefield,
1855
27. Nyctalus montanus Barrett-
English Name
Black-gilded
Pipistrelle
Barbastel or wide
eared bat
Short nosed fruit bat
Northern bat
Nepali Name
Status
NT
Chamero
NT
Chamero
Chamero
LC
DD
Common serotines
Chamero
NT
Tickell’s Bat
DD
Andersen’s Leafnosed Bat
Large Himalayan leafnosed bat
Least leaf-nosed bat
NT
Fulvous Leaf-nosed
Bat
Painted bat
Great Evening Bat
False vampire bat
Nake chamero
LC
NT
LC
Chirbire
chamero
Chamero
LC
CR
LC
Long-winged bat
CR
Jerdon smokey bat
LC
Little tube-nosed bat
NT
Hutton’s Tube-nosed
Bat
Greater Tube-nosed
Bat
Lesser Mouse-eared
Bat
Csorba’s Mouse-eared
Bat
Hodgson’s bat
Kashmir Cave Bat
Vespestilled bat
Whiskered bat
Myotis bat
Small horseshoe bat
DD
Himalayan noctule
VU
VU
DD
Chamero
NT
NT
Bokso chamero LC
VU
VU
NT
Chamero
NT
44
28.
29.
30.
31.
32.
33.
34.
Hamilton, 1906
Nyctalus noctula Schreber,
1774
Philetor brachupterus
Temminck, 1840
Pipistrellus affinis Dobson,
1871
Pipistrellus coromondra Gray,
1838
Pipistrellus javanicus Gray,
1838
Pipistrellus tenuis Temminck,
1840
Plecotus auritus Linnaeus, 1758
35. Plecotus austriacus Fischer,
1829
36. Pteropus gyganteus Brunnich,
1782
37. Rhinlophus subbadius Blyth,
1844
38. Rhinolophus affinis Horsfield,
1823
39. Rhinolophus lepidus Blyth, 1844
40. Rhinolophus luctus Temminck,
1834
41. Rhinolophus macrotis Blyth,
1944
42. Rhinolophus pearsonii
Horsfield, 1851
43. Rhinolophus pusillus
Temminck, 1834
44. Rhinolophus rouxii Temminck,
1835
45. Rhinolophus sinicus Andersen,
1905
46. Rousettus leschenaulti
Desmarest, 1820
47. Scotomanes ornatus Blyth, 1851
48. Scotophilus heathii Horsfield,
1831
49. Scotophilus kuhlii Leach, 1821
50. Sphaerias blanfordi Thomas,
1891
51. Taphozous longimanus
Hardwicke, 1825
Noctule
LC
Rohu’s Bat
VU
Chocolate bat
?
Indian pipistrelle bat
LC
Javan pipistrelle
LC
Indian Pygmy bat
LC
Long eared bat
Lamkane
chamero
Brown long-eared Bat
Flying fox
NT
NT
Phayre
chamero
LC
Chestnut Horseshoe
Bat
Intermediate Horseshoe Bat
Horseshoe bat
Large leaf bat
EN
Big eared horseshoe
bat
Pearson’s leaf bat
NT
Least Horseshoe Bat
LC
Rufous bat
NT
Andersen’s Rufous
Horseshoe Bat
Fulvous fruit bat
LC
Harlequin Bat
Commong yellow bat
LC
NT
NT
LC
Chamero
Pahelo
chamero
NT
NT
LC
Lesser yellow bat
Blanford’s Fruit Bat
NT
NT
Long-winged Tomb
Bat
DD
45
Flying Fox (Pteropus gyganteus)
It is a common fox bat of midland Nepal. Its
head and body measure about 23cm and
wingspan is 112cm. It weighs well over
630gm. The head of the fox bat is dark brown,
sometimes blackish. The shoulder and rear
part of the neck is brown. The abdomen is
yellowish brown. The chin, neck, and flanks
of the bat are darker. The wings are black due
to exposure to heat.
Pteropus gyganteus
Large Horseshoe Bat (Hipposideros armiger)
It is a light dark brown horseshoe bat. Its distribution is in the Himalayan foot-hills. The leaf
nosed are distinguished from horseshoe bats
Rhinolophus by absence of a notch separating
the antitragus from the outer margins of the ear
and in all toes having two joints only. It is found
in
Dharan
and
Barahachettra.
A
good
concentration of horseshoe bat occurs in
Biratnagar. This bat is the largest one of the
genus. The head and body of the measure 8.5cm
and tail measures about 5cm. This bat has a
narrow leaf nose sinuate slightly elevated at the
centre. Some warlike granular patches and
vertical ridges are prominent in nose leaf. The
eyes are well developed and form large thick
Hipposideros armiger
elevations on each side. The color of the bat
variable according to the locality. It is often regarded as the most interesting species having
wider distribution. It is found in the entire range of the Himalayas such as Nepal. Sikkim,
Khasia hills and Ceylon.
46
Great Eastern Leaf Bat or Pearson’s Leaf Bat (Rhinolophus pearsonii)
It is a large sized bat. Its ear is very large. The
head is broad and acutely pointed; nasal
apparatus is very complicated. The lower nose
leaf is large, concealing upper lip like a door
knocker. The upper leg resembles a graduate
spine. The ears are transversely straight with a
large semicircular lobe at the base of the ear.
The body of the animal is beset with long dense
fur. The head and body measure 7cm, tail
2.2mc, wing expanse is about 3.8cm. The wing
membrane is densely pigmented. This bat has
been recorded from Darjeeling and Khasia hills.
In central Nepal, this bat occurs in Godawari.
Rhinolophus pearsonii
Rufous Leaf Bat (Rhinolophus rouxii)
It is a bluish brown bat with fuscous pale pelage
and light drab venter. The head and body of the bat
measure 9.3cm, tail 2.2cm. Its fore arms about 6.6
cm, and the hind foot 9.5 cm. nose leaf is
characteristic in the species. This bat weighs about
9.5cm, it is the largest species of the genus. The
body fur of this bat is wooly jet black in color. The
hairs are tipped with ashy brown tips. In this bat
and leaf nose is large, projecting over lip and is
deeply incised at the middle. The lower lip
possesses a median groove. Tail is encased in the
large inter femoral membrane. The bat lives in
caves and old building. Generally they roost alone
or in pairs. This bat noiselessly flies over house and
Rhinolophus rouxii
trees. Prey upon active insects. They hibernate in
47
the foot-hills of Himalayas during winter. Two races, R. perniger (Hodgson) and R.
beddonori (Anderson), have been reported from India.
Large Leaf Bat (Rbinolophus luctus)
It is a large-sized bat. Its ear is very large. The head is broad
and acutely pointed; nasal apparatus is very complicated.
The lower leaf is large, concealing an upper lip like a door
knocker. The upper leaf resembles a graduated spine. The
ears are transversely straight with a large semi circular lobe
at the base of the ear. The body of the animal is beset with
long dense fur. The head and body measure 9.25cm; tail
4.25cms; wing expanse is about 42cms. This bat had been
recorded from India, Darjeeling and Khasia Hills, western
ghat and Ceylon. This bat roosts alone or in pairs in corners
Rhinolophus luctus
of old deserted caves. It emerges to hunt in the evening, flying mostly around gardens above
15 to 25 feet. The breeding habit of the bat is not clearly known. Females are been breeding
their young during April and May.
Lesser Leaf Nosed Bat (Rhinolophus affinis)
It is one of the uncommon bats of Nepal. It is large-sized bat
with head and body 6.3cms, tail 2.2cms and wing span 15cms.
It weight about 15gms. The ear of this bat is large than head;
obtusely pointed and avoid; nasal appendages quadrate; with a
transverse bar nearly surrounding it. The upper leaf is
triangular with marinated sides. Dorsal part is brown and
ventral part is wood brown. The head and face is pale brown.
It has been recorded from Godawari of central Nepal by Ade
Rhinolophus affinis
(1971). This bat also occurs in Shrilanka and Burma.
Common Yellow Bat (Scotophilus heathii)
This is an interesting bat of midland Nepal. The head and body of the bat measure 75mm tail
50mm and arm about 54mm. Its dorsal aspect is yellowish brown and under part is canary
yellow. It occurs all over India and South East Asia. This is a one of the gregarious bats; it
roosts in small colonies and hibernates in crevices or crack. Generally, the bat engage noisy
quarrel while hunting and resting. The bat is active in evening. It feed on flying ants.
48
Big-Eared Horseshoe Bat (Rhinolophus macrotis)
The big eared bat is often found in dense forests
and caves. Its ears are very large, broad, oval,
with pointed recurved tip and a large obtuse
targus. The anterior central crest of nose leaf
produced in front over the top of the flat
transverse front edge, hinder leaf lanceolate
triangular, above sooty brown or light earthy
olive brown, paler below, some with a rufous or
Rhinolophus macrotis
isabelline tint and no pubic eats. It was reported
by Blyth (1844) from Nepal. Its distributional range is Indo-Malayan region.
Bay-Leaf Bat (Rhinolophus subbadius)
Its ears are smaller than the head, obtusely pointed and ovoid, nasal appendage quadrate with
a transverse bar nearly surmounting the upper leaf triangular with slightly emarginated sides.
It is clear brown above, paler below and on head and face. It was reported by Blyth (1844).
Its distributional range is Indo-Malayan region.
Javan Pipistrelle Bat (Pipistrellus javanicus)
It is a medium-sized blister back and with
pale-tinged fur on the venter. The head
and body of the measure 4.6mc, tail
3.4cm, forearm is about 3.1cm and hind
foot measures about 6.5cm. The bat
weighs about 7.2gm. It is known to occur
in Pokhara valley and Biratnagar. It is a
dark brown bat with black muzzle the
crown of the head and forehead are
Pipistrellus javanicus
densely furred. The ears are sub-triangular, rounded at the extremity. It is a most common bat
frequently encountered on roofs of houses and buildings. This bat emerges to hunt in the earl
evening. Generally, the bat preys upon flies and insects. It flight is fast and erratic. This bat
also undergoes hibernation during winter. Its abundance increases in May. Females contain
embryos at this time.
49
Nepalese Whiskered Bat (Myotis muricola)
This is an elegant bat of midland Nepal. It has long dense bicolor fur on head and back. The
back of the bat is colored and under parts are smoky grey. The anterior part of the body is
covered by sparse golden hairs and remaining posterior and under part consist of pure grey
wooly hairs. The size of the bat is moderate. Its head and body measure 50cm and tail 32cms.
The bat weighs about 4gms. Tale (1941) distinguishes Myotis mystacinus pepalonsis from
other bats by its white ventral fur.
Asiatic Lesser Yellow House Bat (Scotophilus kuhlii)
This bat has relatively short, close and sleek fur. The color of the coat varies from reddish to
olive-brown dorsally and a scattering of dark reddish tipped hairs in the mid-belly region.
The ears are relatively small and rounded and the targus is half-moon shaped being markedly
convex on its posterior border and concave interiorly the tip is slender and forward pointing
and extends half way up the ear. The upper part of the muzzle behind the eyes often has
glandular swellings and the canines appear powerful it goes out at dusk for hunting. It shows
relatively strong direct type of flight, presumably covering a fairly extensive territory in its
hunting. In this species twins are found to be commonly produced as shown by study in
Mysore, South India (Gopalakrishna, 1949). This bat has head and body length generally up
to 7.5cm and tail 4.5-5cm.
Short-Nosed Fruit Bat (Cynopterus sphinx)
It is a small-sized fruit bat. They occur in small
colonies. Wherever palms are available, they
become favorite roosting places for this bat. The
head and body of this bat measure about 9.4 to
10.6cm and tail 0.75 to 0.95cm. It weighs about
50gm. The forearm measures 6.3 to 7cm and
hind foot 1.4cm. This bat is easily distinguished
by white margined naked ears and divergent
nostrils. Generally, males are reddish brown.
This bat occurs in garden with palms, plum
trees, oranges, and bottlebrush trees Kathmandu
Cynoperus sphinx
50
city. It has also been recorded from Chisapani (near Kathmandu) by Johnson in the year
1980. This bat also occurs in Makalu Barun National Park, Bardiya National Park and Rara
National Park. The Indian race Cynopterus sphinx (Vahl) occurs in South India.
Jerdon’s Smoky Bat (Miniopterus schreibersi)
This is one of the commonest bats of
central Nepal. The size of the bat is
medium. Its head and body measure
6.3cm tail 4.3cms. This bat weighs
about 10.5gms. The wing span measures
about 30cms. The feet are very small
and are often included in the wing
Miniopterus schreibersi
membrane nearly to the end of the toes. The ears are acutely pointed, shorter than the head:
muzzle grooved, wholly scaly brown. This bat has been reported from central Nepal by Sir
Brian Hodgon. It has been recorded from Pokhara Valley by Abe (1971). Its allied species
also are known to occur in Japan, Java and China. Originally this bat was known by the name
Scotophilus fulligenous. Two other species, S. heathii and S. belangeri, also have been
reported from the Indian frontiers. It is a gregarious bat often found roosting in colonies in
crevices or creeks, on roofs, rafter and in holes in ceiling. The yellow bat is quite quarrelsome
and often including in noisy squabbles among themselves in their diurnal hunts. It is known
to feed primarily on ants. It is known to hibernate in the Himalayan foothills.
Fulvous Fruit Bat (Rousettus leschenaualti)
It is medium sized bat, uniformly light brown but occasionally yellowish in color. The older
male is grey flanks. During spring and summer moults completely hairless individuals may
be seen. The bats have an odor like that of fermented fruit. It measures 12.5cm and tail 2cm.
Fulvous fruit bat roosts in noisy colonies of 10 to 2000 in caves and old ruins. Most members
of a colony leave at dusk, flying heavily with slow wing beats in search fruit bearing trees. It
has very good sense of smell and memory of fruit trees in the area. It may travel fairly long
distance. It only swallows fruit juices. The young ones are pink and naked at birth and carried
by the mother for two month during her nocturnal flight. The juveniles after being
independent live in exclusive colonies of their own. This bat occurs in Annapurna
Conservation Area, Makalu Barun National Park, Chitwan National Park and Rara National
Park. Extralimitally it occurs in Indo-Malayan region. Pakistan, Indian, Sri Lanka, Burma, S.
51
China. It roosts in caves, man-made structures such as tunnels, rocks, caves, wells and rooms
in old ruins. It is found up to 2150m.
Great False Vampire Bat (Megaderma lyra)
This is a large bat with a rather ugly
appearance due to its big had with
prominent muzzle, huge upright seminaked ears bluntly rounded at tips and
peculiar nose leaf. The body fur is
noticeably greyer than other bats and
blue-grey dorsally and consisting of long
silky hair. The abdominal fur is a paler,
more yellowish grey. The wings are
Megaderma lyra
rather broad due to the last of fifth digit
being relatively long. Such a design probably results in a slower but more controlled type of
flight which is adapted to their method of hanging. There is no sign of tail in this species but
the interfemoral membrane is well developed, stretching from heel to heel and being
supported by long but weakly developed calcars. There are two conspicuous dark vein-like
lines stretching diagonally from the base of where the tail would normally be, to the heels.
Usually hind feet are comparatively large in this species sand further peculiar feature is in the
development of the first digit which consists of only two joints (phalanges) whilst the
remaining digits have three joints. Its eye is quite large and conspicuous and elongated nose
leaf extends up between the eyes. The skin of the nose leaf is pink and naked and roughly
lozenge shaped with a narrow vertical ridge running down its centre. The upper lip tends to
be rather sparsely haired with a fleshy furrow dividing the middle of the lower lip. There are
no incisors in the upper jaw. Females have two pectoral mammae with two more false teats in
the pubic region.
Blanford’s Fruit Bat or Mountain Fruit Bat (Sphaerias blanfordi)
This bat was reported by Legagul and McNeely (1977) from eastern Nepal. Extralimitally it
is distributed in Indo-Malayan: North India, Bhutan, Burma, Northwest Thailand, South
Tibet, South China. It is distributed in altitudinal range is 1000m.
52
Brown Long-Eared Bat (Plecotus auritus)
This bat is reported from Makalu Barun National Park and Rara National Park. It ranges
Palaearctic and Nearctic. In Indo-Malayan: North India and Pakistan. Palaearctic from West
Europe to Japan.
Least Leaf-Nosed Bat (Hipposideros cineraceus)
It
is
recorded
from
Annapurna
Conservation Area. Extralimitally it is
distributed in Indo-Malayan: north East
Pakistan, north India, Burma, Thailand,
Vietnam, Malaya. It roosts in caves, rock
crevices and hollow trees up to 1500m
altitude.
Hipposideros cineraceus
Hodgson’s Bat (Myotis formosus)
It is found in Annapurna Conservation Area. Extralimitally is
distributed in Indo-Malayan region: North India, South China,
Taiwan, Afghanistan, Tibet, N. China, Korea and Japan. A
wide variety of roosters is utilized from caves and rock
shelters to man-made structures such as well or mine shafts.
Some roost in hollow trees, among foliage, under loose bark,
in the internodal spaces of bamboos, even in unusual bird
nests or in flower. Some colony is found in ruined attics of
houses or other ruined buildings.
Myotis formosus
Whiskered Bat (Myotis mustacinus)
The whiskered bat is a small bat with soft fine body fur of blackish-brown with a few of the
longer dorsal hairs tipped reddish brown. The belly fur is slightly paler and blackish grey.
The tail is enclosed up to the extreme tip in the intermemoral membrane and there is a
longish flexible calcar with a sub-calcarial lobe. The ears are longer and naked with blackish
color. The head and body length is about 4.5cm and tail 3.5cm. It is a crevice rooster in its
diurnal biotope. This species hibernate in winter. It is reported from Annapurna Conservation
and Makalu Barun NP. Cosmopolitan in Indo-Malayan region North Pakistan, North east
53
India, South China. Also in the Palaearctic from West Europe and North Africa to Korea and
Japan. It inhabits a wide variety of roosts from caves and rock shelters to man-made
structures such as well or mine shafts. Some roost in the hollow trees, among foliage, under
loose bark, in the internodal spaces of bamboos, even in disused bird nests or in flowers.
Some colony is found in ruined attics of house or other ruined buildings.
Himalayan Whiskered Bat (Myotis siligorensis)
It is reported from ACAP and Makalu
Barun National Park. It ranges IndoMalayan: North and Northeast India,
Southern China, Vietnam. It inhabits a
wide variety of roosts from caves and
rock shelter to man-made structures
such as or mine shafts. Some roost in the
hollow trees, among foliage, under loose
Myotis siligorensis
bark, in the internodal spaces of
bamboos, even in unused bird nests or in flowers. Some colony is found in ruined attics of
house or other ruined buildings.
Indian pipistrella (Pipistrella coromandra)
It has dorsal fun and blackishbrown and the tip of the hairs
slightly rufescent. The belly fur
is slightly paler brown. There is
a small lobe below the calcar
and tragus is forward curving
and blunting round. The head
and body length of animal is 44.5cm and tail 3-3.5cm. It is
reported from ACAP, Makalu
Pipistrellus coromandra
Barun National Park, Bardia
National Park, Chitwan National Park. It inhabits caves, crags, large enclaves, abandoned
ruins and temporary sheds. It is also reported from Ilam district. Extralimitally it is distributed
54
in Indo-Malayan: North-west Pakistan, India, Sri Lanka, Burma, North Thailand, Afghanistan
and Tibet.
Eastern Barbastelle (Barbastelle leucomelas)
This is rather a small and delicately built bat with dorsal fur long and silky blackish grey
color basally, with the extreme tip of the hairs a pale golden brown. The belly fur is paler
grayish-brown. The ears are large and conspicuous being rather squarish in outline and
forward slanting. It measures about 5cm. This bat is reported from ACAP, Makalu Barun
National Park and Rara National Park. It ranges Paleaearctic and adjacent part of IndoMalayan, Afghanistan, North Pakistan, North India, South China.
Northern Bat (Eptesicus gobiensis, syn, E. nilssoni)
The Northern bat is reported from the Makalu Barun National Park. Exralimitally
cosmopolitan in distribution, Europe to East Siberia, possibly Japan.
Common Noctule (Nyctalus noctula)
Common noctule is a large, robust-looking bat with
very glossy fur usually quite bronzey red when
viewed dorsally. The wings are long and narrow and
the under surface is thickly furred along the flank up
to the elbow as well as in a narrow band extending
below the forearm. It has board and flat head, ears
oval and broad. The outer margin of the ears convex,
reflected backwards and forming a thick lobe
Nyctalus noctula
terminating close to the angle of the mouth. Tragus
short and curved inwards, muzzle devoid of hair, fur dark reddish brown. It is a noisy bat
both in its diurnal roosts in its hunting at night time. They are particularly quarrelsome and
noisy towards evening before they emerge to hunt. It is gregarious in its diurnal roost and the
females form separate maternal colonies. It was reported by Hodgson (1835) as V. labiata. It
occurs in Palaearctic and Indo-Malayan regions.
Himalayan Noctule (Nyctalus montanus)
Himalayan noctule has darker color than common Noctule. The dorsal fur is blackish brown
lacking any rufescence and belly fur is paler brown. The wings are long and narrow and
extended to the ankle of the hind foot. The low rounded ear is blackish and naked. Its head
55
and body measures 6-6.5cm and tail 0.5-0.8cm. It is distributed in ACAP and Makalu Barun
National Park. Exralimitally it is distributed in Indo-Malayan region: East Afghanistan, North
India.
Painted Bat (Kerivoula picta)
Painted bat is a beautiful small
bat. Its body is covered with fine
woolly fur. The dorsal surface is
yellowish-red or golden rufous,
beneath is less brilliant and more
yellow. Its wing membrane is
inky black with rich orange
stripes
along
the
fingers
extending in indentation into the
Kerivoula picta
membrane. Its head and body measures 3.5cm and tail 2.5cm. This animal is reported from
Chitwan National Park and Bardia National Park. Extralimitally it range Indo-Malayan:
India, Sri Lanka, Burma, south China, Vietnam, Thailand, Malaya in altitude below 1000m.
Common Serotine (Eptesicus serotines)
It is large of Serotines. The body fur is
very long and silky and dorsal fur is dark
brown with a scattering of silvery-buff
tipped hairs especially over the region of
shoulder. The belly fur is shorter and more
grey. The naked ears are rather long and
narrow and blackish brown in color. Its
Rousettus leschenaulti
head and body measures 6-8cm and tail
5.5-5.8cm and weight 5.5gms. The common serotine bat is reported from Makalu Barun
National Park. Extralimitally cosmopolitan. In Indo-Malayan realm: Pakistan, North-west
India, America, Palaearctic and Africa, predominantly Palaearctic in the old world, extending
marginally into the northern part of the Indo-Malayan Region.
Mandelli’s Mouse-eared Bat (Myotis sicarius)
It is reported from Annapurna CA and MB NP. Extralimitally is distributed in Indo-Malayan:
Northeast India, Sikkim. It inhabits a wide variety of roosts from caves and rock shelter to
56
man-made structures such as or mine shafts. Some roost in the hollow trees, among foliage,
under loose bark, in the internodal spaces of bamboos, even in disused bird nests or in
flowers. Some colony is found in ruined attics of house or other ruined buildings.
Blyth’s Horseshoe Bat (Rhinolophus lepidus)
Its habit is solitary as well as colonial and
its main habitat is forests. Its niches
include caves, ruins, dungeons, tunnels,
subterranean soils, old houses, ruined
temples. It is found up to 2388m. It has
been recorded from Ilam and Sindhuli. Its
main threats are human interference,
tourism. The influence on the population
well understood, not reversible and have
Rhinolophus lepidus
not ceased to be a threat.
Least Horseshoe Bat (Rhinolophus pusillus)
Its habit are colonial and occurs in cave
dweller,
humid
uplands,
hills
and
valleys, coastal areas in about 2000m. It
is
found
Nagerjunban,
in
Kathmandu
Bimalnagar,
Valley,
Pokhara,
Sundarijal. Its main threats is human
interference. The influence on the
population
well
understood,
not
reversible and have ceased to be a threat.
Rhinolophus pusillus
Brown long-eared Bat (Plecotus austriacus)
Its habit is insectivorous, colonial and habitat
is montane moist mixed conferous and
deciduous forest. Its niche includes crevices of
tunnels, forts, dilapidated buildings, caves and
occurs in 1450-3600m. Specimens from
Indian subcontinent and Afghanistan are
Plecotus austriacus
57
provisionally referred to as Plecotus austriacus wardi (Bates and Harrison, 1997). No
population information is known. On conservative estimate of 40 individuals in 17 locations
the population could be 680. Wide distribution from Afghan to Jammu and Kashmir. Status
based on habitat loss in its distributional range.
Serotine Thick-eared Bat (Eptesicus serotinus)
Its habit is solitary or in small numbers,
insectivorous and its habitat includes caves and
cracks in rocks, rocky riverines, montane, tree
hollows, caves and cracks in rocks and occurs in
462-2338m. Its main threat is habitat destruction
and exploitation as medicine. It is found in
Makulu Barun National Park.
Eptesicus sp.
Fulvous Leaf-nosed Bat (Hipposideros fulvus)
Its habits are colonial (single and mixed roosts), and is
insectivorous. Its habitat is habitat subterranean caves,
wells, ruins of houses, thorn scrub, caves. It is found in
Kathmandu
Valley.
Its
main
threats
are
human
interference, recreation / tourism, stone-quarrying, sealing
and fumigation of caves. Threat due to stone quarrying is
irreversible whereas threat in caves is reversible.
Fulvous Fruit Bat (Rousettus leschenaultia)
Hipposideros fulvus
It occurs in colonies of as much as 10,000. It hunts for meat at Garo hills in Meghalaya and it
has been found in Baitari and Kathmandu.
Nicobar Long-fingered Bat (Miniopterus pusillus)
Its habit is cave dweller, insectivorous, lives in colonies
of up to 700 individuals. Its habitats are limestone caves,
rock clefts, culverts, crevices of trees and occurs up to
1200m. It is found in Pokhara. Its main threats are human
interference, ecological imbalance, habitat loss and
disturbance of roosts by man. It is difficult to calculate
extent of occurrence with certainty since the localities are
Miniopterus pusillus
58
so fragmented. In Australia similar taxa is known to have a small home range of <300 sq. km
for each location. The known localities are fragmented and far apart. In Pokhara, Nepal
earlier studies have reported the species however presently there is a decline because of
habitat destruction.
Lesser Mouse-eared Bat (Myotis blythii)
Its habit is colonial/clusters, insectivorous and its habitat is
scrub forest and low foothills with low rainfall, tropical
semi evergreen forests and also occurs in caves, buildings
(unused), trees, crevices in 170-1754m. It is found in
Tumlingter. The main threats are habitat loss and
deforestation. The influence on the population is well
Myotis blythii
understood, not reversible and has not ceased to be a threat.
Schreiber’s Long-fingered Bat (Miniopterus schreibersi)
Its habit is large colonies and habitats are hilly
and forested country-side and also occurs in
caves, caverns and crevices in rocks up to 3381230m. It is found in Kathmandu Valley, Kakani,
Bimalnagar and Syangja.
Miniopterus schreibersi
Andersen’s Rufous Horseshoe Bat (Rhinolophus sinicus)
Its habit is insectivorous, forest dweller with rainfall, colonial
(1-100 individuals) and its habitat is Montane forests. It also
occurs in humid caves, tunnels, wells, hollow trees, temples,
houses in 500-2769m. It has been found in Godavari, Num,
Parchung, Pulchowki, Shivapuri and Thankot. The main threats
are habitat loss and habitat alterations. Threats influencing the
status of the taxa are not understood, not reversible and have not
ceased to be a threat. Rhinolophus sinicus was upgraded to
species level from Rhinolophus rouxii sinicus by Nikki Thomas
basing on mtDNA analysis (Srinivasulu; Bates).
Rhinolophus sinicus
59
Tickell’s Bat (Hesperoptenus tickelli)
Its habit is solitary in foliage; high
flying and its habitat is common in
low lands and paddy fields, dry and
wet zones and occurs in large foliage
up to 1000m. The main treats are
habitat
loss,
deforestation,
exploitation, hunting, hunting for
medicine and human interference.
It’s a known high flyer, hence rare in
recent
collections,
no
recent
Hesperoptenus tickelli
information.
Andersen’s Leaf-nosed Bat (Hipposideros ponoma)
Its habit is insectivorous, associated with
other Hipposideros sp. Its habitat includes
caves, subterranean habitats and also
occurs in caves, crevices of rocks and
buildings up to 2000m. Controversy
regarding the systematics of the species
and it was proposed that it should be
synonymized with H. hypophyllus. The 2
species are discrete.
Hipposideros ponoma
Little Tube-nosed Bat (Murina aurata)
Its habit is insectivorous and habitat is
subtropical forest and also occurs in near
steep mountain slopes with thick forest in
2000-4154 m. The specimen was collected
last from Shivpuri in Nepal, where threats
are minimal to the species.
Murina aurata
60
Greater Tube-nosed Bat (Murina leucogaster)
Its habit is low flying insectivore, solitary or
groups up to six and its habitat includes
grasslands, plantation and mixed woodland
and also occurs in vegetation or caves up to
2000m. It has been recorded in Kathmandu
valley but very few records and no recent
information has been found.
Murina leucogaster
Csorba’s Mouse-eared Bat (Myotis csorbai)
It is found in caves around 1300m. Though exact location
has not been identified, it is said that it is found in central
Nepal.
Myotis csorbai
61
Distribution of Bats of Nepal and South Asia
62
63
64
65
66
67
68
69
70
Dictionary of Technical Terms
Altricial: Young are born in a relatively
Eocene: Presence of modern mammals,
underdeveloped state; they are unable to
from about 58 million to 40 million years
feed or care for themselves or locomote
ago
independently for a period of time after
Evolution: The gradual development of
birth/hatching. In birds, naked and helpless
something into a more complex or better form;
after hatching
the historical development of a related
Cochleae: The snail-shaped tube (in the
group of organisms
inner ear coiled around the modiolus)
Hibernation: The state that some animals
where sound vibrations are converted into
enter during winter in which normal
nerve impulses by the organ of Corti
physiological processes are significantly
Colonial: Used loosely to describe any
reduced, thus lowering the animal’s energy
group of organisms living together or in
requirements. The act or condition of
close proximity to each other - for example
passing winter in a torpid or resting state,
nesting shorebirds that live in large
typically involving the abandonment of
colonies. More specifically refers to a
homoiothermy in mammals
group of organisms in which members act
Keystone species: A species whose
as specialized subunits (a continuous,
presence or absence strongly affects
modular society) - as in clonal organisms
populations of other species in that area
Crepuscular: Active at dawn and dusk
such that the extirpation of the keystone
Cretaceous: From 135 million to 63
million years ago; end of the age of
reptiles; appearance of modern insects and
flowering plants
Cryptic: having markings, coloration,
shapes, or other features that cause an
animal to be camouflaged in its natural
environment; being difficult to see or
otherwise detect
species in an area will result in the
ultimate extirpation of many more species
in that area
Lek
Mating
System:
Hypsignathus
monstrosus, has a lek mating system,
where males gather in a lekking arena to
display to females, who then choose the
most desirable of mates
Migratory: Makes seasonal movements
between breeding and wintering grounds
71
Monogynous: A mating system, having
Synapomorphies: Powered flight and
one mate at a time
many morphological adaptations for flight,
Motile: Having the capacity to move from
including forelimbs modified into wings.
one place to another
Territorial: Defends an area within the
Nocturnal: active during the night
home range, occupied by a single animals
or group of animals of the same species
Oligocene: Presence of modern mammals,
from 40 million to 25 million years ago
and held through overt defense, display, or
advertisement
Ossicles: A small bone; especially one in
Torpor: A dormant, inactive state
the middle ear
Vampire bats: The bat species which
Pheromones: Chemicals released into air
or water that are detected by and
solely depend on the blood of other
vertebrates and found in the Neotropics
responded to by other animals of the same
species
Visual: Uses sight to communicate
Polygynous: A mating system, having
Viviparous:
more than one female as a mate at one
fertilization and development take place
time
within the female body and the developing
Reproduction
in
which
embryo derives nourishment from the
Promiscuous: A mating system, also
called
polygynandrous;
the
kind
female
of
polygamy in which a female pairs with
Warning
coloration:
several males, each of which also pairs
through body color
Federal
signal
with several different females. See also
polygynous
Scent
marks:
Communicates
by
producing scents from special gland(s) and
placing them on a surface whether others
can smell or taste them
Sedentary: Remains in the same area
Social: Associates with others of its
species; forms social groups
Solitary: Lives alone
72
Index
Altricial, 71
Andersen's leaf-nosed Bat,
44, 60
Animalia, 9, 10
Areilulus circumdatus, 44
Asiatic lesser yellow house
bat, 50
Barbastel bat, 44
Barbastella leucomelas,
44, 55
Bats and man, 30
Bats of Nepal, 43, 62
Bay-Leaf Bat, 49
Behavior, 25
Big eared horseshoe bat,
45
Black-gilded pipistrelle, 44
Blanford's fruit bat, 45, 52
Brown long-eared bBat,
45, 53, 57
Carnivorous bat, 30
Challenges, 41
Characters, 10, 11, 12
Check list, 44
Chestnut horseshoe bat, 45
Chiroptera, 2, 9, 10, 14,
25, 29, 36, 39, 77, 78,
79, 81, 82
Chocolate bat, 45
Chordata, 9, 10
Cochleae, 71
Colonial, 71
Colour, 31
Common serotines, 44, 56
Common yellow bat, 7, 48
Commong yellow bat, 45
Communication, 3, 5, 27,
33, 80
Comparison, 12
Conservation, 36, 51, 53,
76, 77, 78
Craseonycteris
thonglongyai, 13
Crepuscular, 71
Cretaceous, 5, 71
Cryptic, 71
Csorba’s mouse-eared bat,
44, 61
Cynopterus sphinx, 29, 44,
50, 51
Darwin, 3, 77
Defense, 34
Dictionary, 71
Distribution of bats, 62
Eastern barbastelle, 55
Echolocation, 1, 2, 3, 5, 9,
12, 14, 27, 76, 77, 78,
79, 80, 81
Economic importance, 37
Ecosystem, 35
Enemies, 34
English Name, 44
Eocene, 2, 71, 77, 79
Eptesicus gobiensis, 44, 55
Eptesicus serotines, 44, 56,
58
Eutheria, 9, 10
Evolution, 2, 71, 76, 77,
78, 79, 80
False vampire bat, 44
Flying fox, 45, 46
Food, 28, 32, 39
Food habits, 28
Fruit bats, 6, 8, 9, 12, 16,
17, 18, 21, 25, 29, 30,
31, 33, 39, 40
Fulvous fruit bat, 25, 29,
45, 51
Fulvous fruit bat, 8, 51, 58
Fulvous leaf-nosed bat, 44,
58
Geographic range, 6
Great evening bat, 44
Great false vampire bat, 52
Greater tube-nosed bat, 44,
61
Habit, 4, 11, 16, 18, 21, 29,
30, 32, 48, 57, 58, 59,
60, 61
Habitat, 8, 12, 76
Harlequin bat, 45
Heperoptenus tickelli, 7,
44, 60
Hibernation, 12, 24, 33, 71
Himalayan noctule, 44, 55
Hipposireros armiger, 44
Hipposideros cineraceus,
44, 53
Hipposideros fulvus, 44,
58
Hipposideros pomona, 44,
60
Hodgson's bat, 44, 53
Horseshoe, 16, 32, 33, 45,
46, 49, 57, 59
Hutton's tube-nosed bat, 44
Hypothesis, 3, 4, 5
Importance, 37
Indian pipistrella, 45, 54
Indian pygmy bat, 45
Insectivorous, 11, 30, 40
Intermediate horse-shoe
Bat, 45
Interrelationships, 32
Javan pipistrelle, 45, 49
Jerdon smokey bat, 44, 51
Kashmir cave bat, 44
Kerivoula picta, 31, 34,
44, 56
Keystone species, 71
la io, 44
Large himalayan leafnosed bat, 44
Large horseshoe bat, 46
Large leaf bat, 45, 48
Least leaf-nosed bat, 44,
53
Legislation, 39, 40
Legs, 17
Lek mating, 71
Lesser leaf nosed bat, 48
Lesser mouse-eared bat,
44, 59
Lesser yellow bat, 45
Lifespan, 24
Little tube-nosed bat, 44,
60
73
Long eared bat, 45
Longevity, 24
Long-winged bat, 44
Long-winged tomb bat, 45
Mammalia, 9, 10, 77
Mandelli's mouse-eared
bat, 56
Medicine, 39
Megachiroptera, 1, 9, 10,
12, 14, 39
Megaderma lyra, 44, 52
Microchiroptera, 1, 6, 9,
10, 11, 12, 14, 39, 77,
79
Migration, 33
Migratory, 71
Miniopterus pusillus, 44,
58
Miniopterus schreibersi,
44, 51, 59
Monogynous, 72
Motile, 72
Murina aurata, 44, 60
Murina huttonii, 44
Murina leucogaster, 44, 61
Myotis bat, 44
Myotis blythii, 44, 59
Myotis csorbai, 44, 61
Myotis formosus, 44, 53
Myotis longipes, 44
Myotis muricola, 44, 50
Myotis mystacinus, 25, 34,
44, 50
Myotis sicarius, 44, 56
Myotis siligorensis, 44, 54
Neighbors, 29
Nepal, 39, 41, 43, 44, 46,
47, 48, 49, 50, 51, 52,
59, 60, 61, 76
Nepalese whiskered bat,
50
Nepali Name, 44
Nicobar Long-fingered
Bat, 58
Noctule, 23, 33, 45, 55
Nocturnal, 11, 12, 72
Northern bat, 44, 55
Nyctalus montanus, 44, 55
Nyctalus noctula, 45, 55
Oligocene, 5, 72
Ossicles, 72
Painted bat, 44, 56
Pearson's leaf bat, 45
Perception, 27
Pheromones, 72
Philetor brachupterus, 45
Physical description, 13
Pipistrellus affinis, 45
Pipistrella coromandra,
45, 54
Pipistellus javanicus,45,
49
Pipistrellus tenuis, 45
Plecotus auritus, 7, 45, 53
Plecotus austriacus, 45,
57, 58
Polygynous, 72
Predators, 34
Promiscuous, 72
Prospects, 41
Protection, 31, 39, 40
Pteropus gyganteus, 45, 46
Pteropus vampyrus, 13
Rbinolophus luctus, 48
References, 76
Reproduction, 21, 72
Rhinolophus affinis, 45, 48
Rhinolophus lepidus, 45,
57
Rhinolophus luctus, 45
Rhinolophus macrotis, 45,
49
Rhinolophus pearsonii, 45
Rhinolophus pusillus, 45,
57
Rhinolophus rouxii, 45, 59
Rhinolophus sinicus, 45,
59
Rhinlophus subbadius, 45,
49
Rohu's bat, 45
Roles, 35
Rousettus leschenaulti, 40,
51, 45
Rufous bat, 45
Scent marks, 28, 72
Schreiber's long-fingered
bat, 59
Scotomanes ornatus, 45
Scotophilus heathii, 7, 45,
48
Scotophilus kuhlii, 45, 50
Sedentary, 72
Senses, 19
Sexing, 21
Short nosed fruit bat, 44,
50
Small horseshoe bat, 44
Social, 11, 32, 72
Social life, 32
Solitary, 72
sperms, 22
Sphaerias blanfordi, 45, 52
Status, 44, 58, 77, 78, 81
Synapomorphies, 72
Tail, 10, 11, 12, 47
Taphozous longimanus, 45
Taxonomic history, 9
Technical terms, 71
Teeth, 18
Territorial, 26, 72
Threats, 38, 59, 77
Tickell's bat, 7, 44, 60
Torpor, 72
Upside down, 12, 26
Vampire bats, 28, 37, 38,
72
Vespestilled bat, 44
Visual, 72
Viviparous, 72
Volant, 1
Warning coloration, 72
Whiskered bat, 25, 53, 44,
54
Wings, 14
74
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