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Home , Emballonuroidea, Furipteridae, Lesser false vampire bat, Lesser yellow bat, Megaderma, Megadermatidae

Prof. Wipula Yapa started studies on Sri Lankan over 30 years ago. After graduating from the University of Kelaniya in 1985, he carried out an ecological study on cave dwelling A Field Guide to the bats in . He proceeded to Germany in 1990 for his doctoral studies and received his PhD from the Ludwig Maximillians University of Munich. He returned to Sri Lanka in 1995 and joined the Department of Zoology, University of Colombo. In 1995, he initiated an ecological survey on the distribution of bats in Sri Lanka, which was one of the detailed surveys carried out in Sri Lanka in recent times. His research expands to many areas such as breeding habits, roosting ecology, migration, social behavior and pathogens of bats. He is Bats of Sri Lanka currently serving as a Professor in Zoology and is the Head of the Department of Zoology & Environment Sciences of the University of Colombo.

Dilmah Conservation was initiated in 2007 by Dilmah to incorporate environmental conservation efforts into the MJF Charitable Foundation, which focuses on social justice. Dilmah Conservation works towards the sustainable use of the environment in partnership with various governmental and non-governmental organisations. The pledge made by Dilmah founder Merrill J. Fernando to make business a matter of human service is deeply ingrained in the work carried out by Dilmah Conservation. For additional information visit our website at:

www.dilmahconservation.org 77671 00L

Wipula Bandara YapaWipula A Field Guide to the

BATS of Sri Lanka Wipula Bandara Yapa A Field Guide to the Bats of Sri Lanka

Wipula Yapa Contents

Foreword 08 Preface 09 Acknowledgements 10 1. Introduction and Evolution 15 1.1. Bats 15 1.2. Evolution 16 1.3. Classification 18 Classical 18 Modern 19 2. Diversity Habits and Habitats 21 2.1. Morphological Diversity 21 2.2. Reproduction and Social life 24 2.3. Roosting Habits and Habitats 25 2.4. Activity Patterns 30 2.5. Diet 31 2.6. Echolocation 34 3. Conservation and Importance of Bats 37 3.1. Humans and Bats 37 3.2. Economic Importance 38 3.3. Bats and Diseases 40 3.4. Conservation Issues 41 3.5. How Can We Help? 43 4. Bats of Sri Lanka 44 4.1 Country 44 4.2 How to use this book 46 4.3 Family and Species profiles 48 Pteropodidae 48 Rhinolophidae 60 66 78 84 Embalonuridae 110 Molossidae 118 5. Key 124 6. Glossary 128 7. Bibliography and Resources for further study 136 8. Index 141 8.1. Scientific Names 141 8.2. Common Names 141

A Field Guide to the Bats of Sri Lanka 13 Lesser false vampire spasma (GR)

52 A Field Guide to the Bats of Sri Lanka 1. Introduction

1.1. Bats - (Order: Chiroptera)

Bats are ; and just like humans they give birth to live young, nurse them with milk and sometimes even carry them when they go in search of food. They are also a fascinating group of . In fact almost everything about bats is so extraordinary and unique, setting them well apart from all other mammals. For example, they are the only group of mammals capable of sustaining powered flight. For this, their forelimbs are modified to form a wing, which is so different from that of birds. The bat wing is a membrane which covers the hand and also sides of the body (Figure 1). Bats have also developed an effective orientation system which involves the use of sound signals for perceiving the environment, the process which is known as echolocation. Although some other groups of vertebrates such as oil birds, whales and certain species of shrews are known to echolocate, echolocation in bats is far more superior and advanced than the rest. These two specialisations, namely the evolution of flight and echolocation in bats, are most probably the main parameters in the diversification of feeding and roosting habits, reproductive strategies and social behaviour of bats. Thus, bats have been able to exploit many ecological niches that were inaccessible to other group of mammals, making them a very successful group of mammals. So far, over 1200 species of bats have been recorded, amounting to roughly ¼th of the total number of mammals recorded at present. Thereby, they are the second largest mammalian order (next to rodents) in number of species and probably exceed all other such groups in overall abundance. They have successfully colonised almost every continental region on the earth (except Antarctica), as well as many oceanic islands and archipelagos. Their widespread distribution and the unique specialisation in morphology, physiology and behaviours have led to a variety of feeding habits which include feeding on fruits, nectar, leaves, flowers, pollen, insects, fish, other smaller vertebrates and even blood.

A Field Guide to the Bats of Sri Lanka 15 Knee

Wrist Tail Elbow Uropatagium Propatagium

Calcar Ear Tragus

Foot with 5 toes

Upper arm (humerus)

Forearm (radius) Plagiopatagium

Thumb Fifth finger

Second finger

Dactylopatagium

Fourth finger

Third finger

Figure 1 General body plan of a bat

1.2. Evolution

Bats are believed to have evolved before early , 50 million years ago, as a complete skeleton of a bat, Icaronycteris index was found from an early Eocene site. This is the earliest known fossil record of a bat. The skeleton was assigned to the suborder Microchiroptera based on the structure of the forelimb and the morphology of the upper molars. The first true Megachiropteran with dentition adapted to a fruit eating diet was found much later, in the early/middle epoch about 20 million years ago.

16 A Field Guide to the Bats of Sri Lanka Fossil of a bat

Until the 1980s most workers agreed that the common ancestor of all extant bats was a flying , a hypothesis which implies that bats are monophyletic and powered flight evolved only once in mammals (Fig 2). This classical view that Megachiroptera and Microchiroptera are the closest relatives of each other has much supporting evidence from biochemical and molecular studies and from some morphological characteristics such as the vascular system of the skull, foetal membranes and the nervous system.

However, there is also evidence to support the view that bats are diphyletic, which means that flapping flight evolved twice in mammals. The first event is believed to have taken place in the period when an insectivorous mammal lineage evolved from leaping to full flight in pursuit of flying insects, thus giving rise to Microchiroptera. The second event would have probably occurred in the Tertiary period when an early primate began to glide in search of fruits. Some of these gliders are believed to have given rise to Megachiroptera. This flying primate hypothesis arose from the unexpected finding that Megachiroptera shares a number of advanced visual pathway characteristics with primates that were not found in any other mammalian order, nor significantly in Microchiroptera. This hypothesis has found increasing support from other sources of evidence such

A Field Guide to the Bats of Sri Lanka 17 as brain features, immunological studies of serum proteins with monoclonal antibodies and analysis of restriction sites and protein sequences, globin and a-crystallin. These alternatives and different hypotheses concerning the relative phylogenetic positions of non-volant and gliding taxa relative to bats – have significantly different implications in relation to the origin of powered flight in mammals.

Flying Primates Bat Monophyly

Figure 2 Phylogenatic tree

1.3. Classification

1.3.1. Classical Classification

Bats belong to the Order Chiroptera and they are the sole members of this Order. According to classical classification, it is divided into two suborders; Microchiroptera and Megachiroptera. The Megachiroptera are represented by only one family, the Pteropodidae, which are restricted to the Old World tropics of and Asia. There are over 170 species of Megachiropterans. The modern Microchiroptera are divided into four super families, , , Phyllostomoidea and , which indicate evolutionary affinities among 17 families. There are three families in the super family Emballnuroidea: Rhinopomatidae (Mouse-tailed bats), (Sheath-tailed bats) and Craseonycteridae (Bumble bee bat). Rhinolophoidea includes four families: Nycteridae (Hollow-faced or Slit-faced bats), Megadermatidae (False Vampire bats), Rhinolophidae (Horseshoe bats) and Hipposideridae (Leaf-nosed bats). Superfamily Phyllostomoidea includes: Noctilionidae (Fisherman bats), (Moustached bats) and Phyllostomidae (New World Leaf-nosed bats). The remaining superfamily Vespertilionoidea includes families: (Funnel-eared bats), (Smoky bats), Thyropteridae (Disc-winged bats), Myzopodidae (Old world Sucker-footed bats), Vespertilionidae (Evening bats), (New zealand Short-tailed bats) and Molossidae (Free-tailed bats). Finally there is a fifth Superfamily, the Palaeochiropterygoidea, which includes three families of fossil

18 A Field Guide to the Bats of Sri Lanka Microchiropterans (all extinct): the Palaechiropterygidae, Archaeonycteridae and Icaronycteridae. Order Chiroptera

Suborder Megachiroptera Family Pteropodidae (Fruit bats)

Suborder Microchiroptera

Superfamily Emballonuroidea Family Emballonuridae (Sheath tailed bats) Family Craseonycteridae (Bumble bee bat) Family Rhinopomatidae (Mouse-tailed bats) Superfamily Rhinolophoidea Family Rhinolophidae (Horseshoe bats) Family Hipposideridae (Old world Leaf nosed bats) Family Megadermatidae (False Vampire bats) Family Nycteridae (Slit faced bats) Superfamily Phyllostomoidea Family Phyllostomidae (New world Leaf-nosed bats) Family Mormoopidae (Moustached bats) Family Noctilionidae (Fisherman bats) Superfamily Vespertilionoidea Family Thyropteridae (Disc-winged bats) Family Furipteridae (Smoky bats) Family Mystacinidae (New Zealand Short-tailed bats) Family Myzopodidae (Old world sucker-footed bats) Family Vespertilionidae (Evening bats) Family Molossidae (Free tailed bats) Family Natalidae (Funnel-eared bats)

Table 1. 1. Classical classification of bats

1.3.2. Modern Classification

Recently, in studies based on molecular phylogeny, some researchers (for details refer Teeling et al., 2002) made a drastic change in the classical classification of Order Chiroptera by introducing two new suborders; Suborder - and Sub order - . All the Megachiropterans (fruit bats) and the members of the superfamily Rhinolophoidea (which were Microchiropterans in the traditional classification), were grouped together in the Suborder Yinpterochiroptera while the remaining families of Microchiropterans were included in the suborder Yangochiroptera. Thus, the traditional division of the two Chiropteran suborders, (the Megachiroptera which included non-

A Field Guide to the Bats of Sri Lanka 19 echolocating bats and Microchiroptera (echolocating bats) have been changed. The suborder Yinpterochiroptera comprises five families: Pteropodidae (all fruit bats or megachiropterans), Rhinopomatidae (mouse-tailed bats), Megadermatidae (false vampire bats), Craseonycteridae (hog-nosed bat or bumble bee bat) and Rhinolophidae. The family Rhinolophidae consists of two sub families: Rhinolophinae and Hipposiderinae. In the classical classification, Family Rhinolophidae consisted of only horse-shoe bats, containing a single , Rhinolophus, and Family Hipposideridae with old world leaf-nosed bats. However, in the new classification horseshoe bats or rhinolophids were included in subfamily, Rhinolophinae and, all old world leaf-nosed bats were included in the subfamily Hipposiderinae. Family Craseonycteridae contains a single species Craseonycteris thonglongyai. Suborder Yangochiroptera comprises 12 families; Nycteridae, Emballonuridae, Phyllostomidae, Mormoopidae, Noctilionidae, Furipteridae, Thyropteridae, Mystacinidae, Myzopodidae, Vespertilionidae, Natalidae and Molossidae.

This sister-taxon relationship between the rhinolophoid and the megachiropterans suggest that laryngeal echolocation within bats evolved more than once. Or alternatively, if echolocation evolved only once, then this ability was lost subsequently in Megachiropterans. There remains considerable uncertainty in both subordinal and superfamilial classifications within bats, where both morphological and molecular data conflict and different molecular data sets provide varying support. Thus, phylogeny of Chiroptera is not yet fully resolved and further studies may one day clear the taxonomic ambiguity of bats.

Order Chiroptera

Suborder Yinpterochiroptera Superfamily Pteropodoidea Family Pteropodidae (Fruit bats) Superfamily Rhinolophoidea Family Rhinopomatidae (Mouse-tailed bats) Family Megadermatidae (False Vampire bats) Family Craseonycteridae (Hog Nosed bat) Family Rhinolophidae Subfamily Rhinolophinae (Horseshoe bats) Subfamily Hipposiderinae (Old world Leaf nosed bats)

Suborder Yangochiroptera Superfamily Emballonuroidea Family Emballonuridae (Sheath tailed bats) Family Nycteridae (Slit faced bats) Superfamily Family Phyllostomidae (American Leaf nosed bats) Family Mormoopidae (Moustached bats)

20 A Field Guide to the Bats of Sri Lanka Family Noctilionidae (Fisherman bats) Family Furipteridae (Smoky bats) Family Thyropteridae (Disk-winged bats) Family Mystacinidae (New Zealand Short-tailed bats) Family Myzopodidae (Old world sucker-footed bats) Superfamily Vespertilionoidea Family Vespertilionidae (Evening Bats) Family Molossidae (Free tailed bats) Family Natalidae (Funnel-eared bats)

Table 1. 2. Modern classification of bats 2. Diversity, Habits and Habitats

2.1. Morphological Diversity

In general Megachiropterans are larger than the Microchiroptera. However, there is a considerable overlap in body size. The weight of the Megachiropterans varies from 10g to 1.5 kg and the forearm length (i.e. the length of the radius and ulna, which is generally used to indicate the size of a bat) ranges from 35-230mm. The wingspan or spread of the wings of the largest fruit bat, vampyrus is about 170cm. Yet the majority of Megachiropetrans are relatively small. There are 42 genera of Megachiropterans and the maximum forearm length is less than 70mm in about 50% (24 genera). The maximum Fore Arm Length (FAL) exceeds 1 meter only in 7 genera, which include the genus Pteropus.

The largest bat, Golden-crowned Flying Fox (Acerodon jubatus)

A Field Guide to the Bats of Sri Lanka 21 The smallest bat, Kitti’s Hog-nosed Bat (Craseonycteris thonglongyai)

The largest Microchiropteran as measured by forearm length is the false , Vampyrum spectrum, which attains a wingspan of 60 - 100cm and body mass of 145–190g. This is also the largest carnivorous bat in the world. The smallest bat, Kitti’s Hog-nosed bat found in , Craseonycteris thonglongyai, (probably the smallest mammal in the world as well) has a body weight of just 2g.

In the eyes of many people, bats are not colourful animals and the general notion is that bats are dark coloured. Although this is true in most cases, there are some brightly coloured bats, such as the painted bat ( picta), which typically haunts banana plantations and is found in several parts of Sri Lanka. Its entire body, apart from parts of the wing, is bright orange or tawny red in colour, whereas the middle parts of the wing are conspicuously black. The colour may even vary among individuals and even though the typical fur colour of the Rufous (Rhinolophus rouxii) is dull brown, the colour of some individuals may vary from yellow to bright orange. Albinism in bats is rather rare, but albino bats have been seen on rare occasion.

Rufous horseshoe bat (Rhinolophus rouxii) colour variations- Left Dull brown form (HS), Right yellow to bright orange form (TP)

22 A Field Guide to the Bats of Sri Lanka Figure 3 - Above Left and Right Microchiroptera bat Below Right Megachiroptera bat

A B

C D

Figure 4 - Left Different types of bat tails A. Free tail, B. Mouse tail, C. Full membrane, E F D. Sheath tail, E. Short tail, F. tail lacking

A B C D

Figure 5 - Above Different types of noseleafs - A. Triangular, B. Truncate, C. Heart shape, D. Square shape, Below Different types of ears - A. Large rounded, B. Pointed, C. Broad, D. Rounded, E. Triangular

A B C D E

A Field Guide to the Bats of Sri Lanka 23 2.2. Reproduction and Social Life

Bats are a group of highly social animals. Generally bats are gregarious, which means they live in large congregations and in fact they form the largest known mammalian congregations, some of which exceed several millions of individuals (e.g. Davis cave in Mexico: 20 million bats). However, all bats do not live in such groups and many species live in smaller groups. In species that are considered “solitary”, the mother raises her young alone. Thus, the mother and her dependant young (generally a single pup) is the basic social unit of bats, and depending on the species, this may range in size from dozens of individuals to millions.

Bats exhibit numerous specialisations in female reproductive morphology and cyclicity. Nearly all temperate bat species show a single peak of reproduction to avoid harsh weather conditions such as winter. Early mating prior to hibernation occurs in almost all species of temperate bats regardless of taxonomic affiliation. Thus, in most species that live in temperate countries, some form of sperm storage has evolved and the young are born after the winter when weather conditions are more favourable. Seasonal peaks of reproduction are also common in tropical bat species, even though the fluctuation of weather conditions is not as severe as in temperate regions. For tropical insectivorous species, rainfall is the most important environmental determinant in reproductive seasonality. The reproduction of these species is often synchronised with prominent peaks in rainfall, as insects are more abundant in rainy seasons.

Generally many species of bats give birth to a single pup in one breeding season, although twinning is also occasionally seen at least in some species. In Sri Lanka, twining has been observed in a few species such as: tenuis, P. ceylonicus and heathii.

Nursery colony of Hipposideros lankadiva, Sri Lanka (DR)

24 A Field Guide to the Bats of Sri Lanka Index Scientific Names Common Names

Chaerephon plicatus 122 Black-Beared Tomb bat 114 Cynopterus brachyotis 58 Cantor’s Leaf-nosed bat 72 Cynopterus sphinx 56 Chocolate bat 92 affinis 92 Common flying fox 50 tickelli 94 Dusky Leaf-nosed bat 74 Hipposideros ater 74 Egyptian Free-tailed bat 120 Hipposideros fulvus 76 Fulvous fruit bat 54 Hipposideros galeritus 72 Fulvous Leaf-nosed bat 76 Hipposideros lankadiva 68 Great Yellow bat 96 Hipposideros speoris 70 Greater False Vampire bat 82 Kerivoula hardwicki 106 Hardwicke’s forest bat 106 Kerivoula picta 104 Indian Pipistrelle 88 Megaderma lyra 82 Indian Pygmy bat 86 Megaderma spasma 80 Kelaart’s Pipistrelle 90 fuliginosus 108 Large Leaf-nosed bat 68 Murina cyclotis 102 Lesser Dog-faced fruit bat 58 Myotis hasseltii 100 80 Pipistrellus ceylonicus 90 Lesser Woolly Horseshoe bat 62 Pipistrellus coromandra 88 Lesser Yellow bat 98 Pipistrellus tenuis 86 Long-Winged Tomb bat 112 Pteropus giganteus 50 Painted bat 104 Rhinolophus beddomei 62 Pouch-bearing Tomb bat 116 Rhinolophus rouxii 64 Round-eared Tube-nosed bat 102 Rousettus leschenaulti 54 Rufous Horseshoe bat 64 Saccolaimus saccolaimus 116 Schneider’s Leaf-nosed bat 70 Scotophilus heathi 96 Schreiber’s Long-fingered bat 108 Scotophilus kuhlii 98 Short-nosed fruit bat 56 Tadarida aegyptiaca 120 Tickell’s bat 94 longimanus 112 Van Hasselt’s mouse-eared bat 100 Taphozous melanopogon 114 Wrinkled-lipped Free-tailed bat 122

A Field Guide to the Bats of Sri Lanka 141