Bat Biodiversity (Vespertilioniformes: Order Chiroptera) in Some Tropical and Arid-Subtropical Regions of Pakistan

BAT BIODIVERSITY (VESPERTILIONIFORMES: ORDER CHIROPTERA) IN SOME TROPICAL AND ARID-SUBTROPICAL REGIONS OF PAKISTAN

Arshad Javid 2007-VA-516

Lahore

A THESIS SUBMITTED IN THE PARTIAL FULFILLMENT OF THE REQUIREMENT FOR THE DEGREE

DOCTOR OF PHILOSOPHY

WILDLIFE AND ECOLOGY

UNIVERSITY OF VETERINARY AND ANIMAL SCIENCES LAHORE-PAKISTAN

2011 The Controller Examinations, University of Veterinary and Animal Sciences, Lahore.

We the members of the Supervisory Committee, certify that the contents and form of thesis submitted by Mr. ARSHAD JAVID have been found satisfactory and recommend that it should be processed for the evaluation by the External Examiner for the award of the degree.

SUPERVISORY COMMITTEE

CHAIRMAN ______DR. MUHAMMAD MAHMOOD-UL-HASSAN

CO-SUPERVISOR ______PROF. DR. MIRZA AZHAR BEG

MEMBER ______DR. MUHAMMAD ALI NAWAZ

MEMBER ______PROF. DR. MUHAMMAD AKRAM

DEDICATED TO MY FATHER CH. WALI MUHAMMAD (LATE) (MAY HIS SOUL REST IN ETERNAL PEACE)

MY BELOVED MOTHER RUQAYYA BIBI (MAY SHE LIVE A HEALTHY AND LONG LIFE) & MY BROTHERS AND SISTERS

CONTENTS NO. TITLE PAGE 1. Acknowledgements v 2. List of Tables vi 3. List of Figures xi CHAPTERS I. INTRODUCTION 1 Objectives 3 II. REVIEW OF LITERATURE 4 Species account 13 Rhinolophus blasii 13 Rhinopoma hardwickii 14 Taphozous nudiventris 15 Taphozous perforatus 16 Scotoecus pallidus 17 Scotophilus heathii 18 S. kuhlii 19 Pipistrellus ceylonicus 20 P. javanicus 21 P. pipistrellus 22 P. tenuis 23 Hypsugo savii 24 III. MATERIALS AND METHODS 26 Study Area 26 SA 1 - The Margalla Hills National Park (MHNP) 26 SA 2 - The Chinji National Park (CNP) 29 SA 3 - The Lal Suhanra National Park (LSNP) 34 SA4 - The Central Punjab (CP) 36 Sampling Strategy 42 External Morphology 43 Cranial Measurements 44 Bacular Measurements 44 Echolocation 46 Sound analysis 48 IV. RESULTS 50 PART I. BAT SURVEY AND ABUNDANCE 50 SUB-AREA 1 53 Pakistan Museum of Natural History (PMNH) 53 Marghzar Zoo 53 Rawal Town 53 National Agricultural Research Council (NARC) 56

Rattowal 56 Tanaza Dam, Kherimoorat 56 Loi Bher Wildlife Park 56 SUB-AREA 2 56 Kanhatti Garden 59 Sodhi Wildlilfe Sanctuary 59 Sodhi Rest House 59 Shrine of Baba Mehdi 59 Dalwal Village 59 Uchhali Lake 59 Khabbeki Lake 59 SUB-AREA 3 60 Mojgarh 60 Derawar Fort 60 Fish Hatchery Bahawalpur 60 Noor Mahal 63 Marot fort 63 SUB-AREA 4 63 Kalian Daas 63 Government College Gojra 63 Gojra Grave Yard 63 Fish Hatchery Manawa 66 Shalimar Garden 66 Pattoki 66 Head Balloki 66 Chhanga Manga 66 Rasul Nagar 66 Ali Pur Chathha 66 COMBINED ABUNDANCES AND DIVERSITY 66 Monthly captures and seasonal abundances 66 Spatial Variation 69 PART II. SPECIES DISTRIBUTION 73 Family Rhinolophidae 73 Rhinolophus blasii 73 Family Rhinopomatidae 73 Rhinopoma hardwickii 73 Family Emballonuridae 73 Taphozous nudiventris 73 T. perforatus 73 Family Vespertilionidae 73

Scotoecus pallidus 73 Scotophilus heathii 73 S. kuhlii 78 Pipistrellus ceylonicus 78 P. javanicus 78 P. pipistrellus 78 P. tenuis 78 Hypsugo savii 78 PART III. MORPHOLOGY 79 Family Embellonuridae 79 Taphozous nudiventris 79 Principal Component Analysis 81 T. perforatus 85 Family Vespertilionidae 88 Scotoecus pallidus 88 Scotophilus heathii 91 Principal Component Analysis 95 S. kuhlii 99 Pipistrellus ceylonicus 102 P. javanicus 105 P. pipistrellus 113 P. tenuis 116 Hypsugo savii 120 PART IV. ECHOLOCATION CALLS 127 V DISCUSSION 134 GENERAL DISCUSSION 134 BAT SURVEY AND ABUNDANCE 136 SPECIES DISTRIBUTION 141 Family Rhinolophidae 141 Rhinolophus blasii 141 Family Rhinopomatidae 142 Rhinopoma hardwickii 142 Family Emballonuridae 142 Taphozous nudiventris 142 T. perforatus 143 Family Vespertilionidae 143 Scotoecus pallidus 143 Scotophilus heathii 143 Scotophilus kuhlii 144 Pipistrellus ceylonicus 144

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P. javanicus 144 P. pipistrellus 145 P. tenuis 145 Hypsugo savii 146 MORPHOLOGY 147 Family Emballonuridae 147 Taphozous nudiventris 147 T. perforatus 147 Family Vespertilionidae 149 Scotoecus pallidus 149 Scotophilus heathii 149 S. kuhlii 152 Pipistrellus ceylonicus 152 P. javanicus 155 P. pipistrellus 155 P. tenuis 155 Hypsugo savii 159 ECHOLOCATION 164 SUMMARY 167 Future Recommendations 171 LITERATURE CITED 174

ACKNOWLEDGEMENTS

I situate my sincere and humble thanks before Almighty Allah, who created the universe and bestowed the mankind with knowledge and wisdom to search for its secrets.

I wish to express sincere gratitude to my hardworking, responsive and praiseworthy supervisor

Dr. Muhammad Mahmood-ul-Hassan, Chairman and Associate Professor, Department of

Wildlife and Ecology, University of Veterinary and Animal Sciences, Lahore for his affectionate supervision, demonstrative guidance, sympathetic behavior and tremendous help not only in accomplishment of present study but also in every aspect of my life.

I am also grateful to the members of my supervisory committee, Prof. Dr. Mirza Azhar Beg,

Eminent Educationist, Department of Zoology, University of Arid Agriculture, Rawalpindi, Dr.

Muhammad Ali Nawaz, Assistant Professor, Department of Wildlife and Ecology and Prof. Dr.

Muhammad Akram, Chairman, Department of Poultry Production, University of Veterinary and

Animal Sciences, Lahore for their skillful guidance and critical insight.

Special thanks are extended to all my friends especially members of the PhD club and colleagues for their wishes and cordial cooperation during the studies. Lastly, I wish to thank my family especially my father, mother, my brothers and sisters who always pray for my success. They taught, loved and supported me to achieve higher goals in life. Their concern in me can never be fully returned but will always be remembered.

This pioneering study on bats in Pakistan would have not been possible without the financial assistance of the Higher Education Commission of Pakistan (HEC Project/20-1033/R&D) and the Rufford Small Grants Commission UK (RSG 12.4.09). the monetary support provided by both these funding agencies is gratefully acknowledged.

Arshad Javid

LIST OF TABLES Table Title Page No. No.

1. GPS location of seven sites sampled for bats at Margalla Hills National 30 Park and adjacent areas (SA1) from June 2009 to May 2011. GPS location of seven sites sampled for bats at Chinji National Park 2. and adjacent areas (SA2) during a period extending from June 2009 to 32 May 2011. 3. GPS location of five sites sampled for bats at Lal Suhanra National 37 Park and adjacent areas (SA1) from June 2009 to May 2011. 4. GPS location of eleven sites sampled for bats in four districts of central 39 Punjab that jointly form SA4 from June 2009 to May 2011. 5. Time frame and survey intensity in four sub-areas sampled for bats 51 from June 2009 to May 2011. Number of days spent in the field in each month in the four sub-areas 6. and the total number of days spent in the field from June 2009 to May 52 2011. Species, sex, number, age, netting index and relative abundance of the 7. bats collected from seven sampling stations in SA1from June 2009 to 54 May 2011 (n is the number of bats captured from each station). Species, sex and number of bats collected from seven sampling stations 8. in SA1 from June 2009 to May 2011 (n is the number of bats captured 55 from each station). Species, sex, number, age, netting index and relative abundance of the 9. bats collected from seven sampling stations in SA2 from June 2009 to 57 May 2011 (n is the number of bats captured from each station). Species, sex and number of bats collected from seven sampling stations 10. in SA2 from June 2009 to May 2011 (n is the number of bats captured 58 from each station). Species, sex, number, age, netting index and relative abundance of the 11. bats collected from five sampling stations in SA3 from June 2009 to 61 May 2011 (n is the number of bats captured from each station). Species, sex and number of bats collected from five sampling stations 12. in SA3 from June 2009 to May 2011 (n is the number of bats captured 62 from each station). Species, sex, number, age, netting index and relative abundance of the 13. bats collected from eleven sampling stations in SA4 from June 2009 to 64 May 2011 (n is the number of bats captured from each station). Species, sex and number of bats collected from eleven sampling 14. stations in SA4 from June 2009 to May 2011 (n is the number of bats 65 captured from each station). 15. Monthly capture/activity patterns of the twelve bat species captured 67 from some arid subtropical and tropical regions of Pakistan.

Combined seasonal relative abundance (%) of the twelve bat species 16. captured from the four SAs (n is the number of bat; N is the total 68 numer of bats captured in any season). 17. Combined seasonal diversity of various bat species captured from the 70 four SAs. Combined relative abundance (%) of the twelve bat species captured 18. from four sub-areas (SAs) from June 2009 to May 2011 (n is number 71 of bats captured; N is the total bats captured in from any sub-area). 19. Locality related diversity of various bat species captured from all the 72 four sub-areas. Combined mean body mass (g) and external body measurements (mm) 20. of Taphozous nudiventris captured from Margalla Hills National Park 80 (SA1) and Lal Suhanatra National Park (SA3) from June 2009 to May 2011 (n is the number of specimens). Mean cranial measurements (mm) of Taphozous nudiventris captured 21. from Rattowal (SA1) from June 2009 to May 2011 (n is the number of 82 specimens). Mean bacular measurements (mm) of Taphozous nudiventris captured 22. from Rattowal (SA1) from June 2009 to May 2011 (n is the number of 83 specimens). Mean body mass (g) and external body measurements (mm) of 23. Taphozous perforatus captured from Margalla Hills National Park 86 (SA1) and Lal Suhanara National Park (SA3) from June 2009 to May 2011 (n is the number of specimens). Cranial measurements (mm) of Taphozous perforatus captured from 24. Rattowal (SA1) from June 2009 to May 2011 (n is the number of 87 specimens). Mean bacular measurements (mm) of Taphozous perforatus captured 25. from SA1 from June 2009 to May 2011 (n is the number of 89 specimens). Mean body mass (g) and external body measurements (mm) of 26. Scotoecus pallidus captured from SA3 from June 2009 to May 2011 (n 90 is the number of specimens). Mean cranial measurements (mm) of Scotoecus pallidus captured from 27. Bahawalpur Fish Hatchery (SA3) from June 2009 to May 2011 (n is 92 the number of specimens). Mean bacular measurements (mm) of Scotoecus pallidus captured from 28. Bahawalpur Fish Hatchery (SA3) from June 2009 to May 2011 (n is 93 the number of specimens). Combined mean body mass (g) and external body measurements (mm) of Scotophilus heathii captured from Margalla Hills National Park 29. (SA1), Chinji National Park and (SA2), Lal Suhanara National Park 94 (SA3) and some localities of Central Punjab (SA4) from June 2009 to May 2011 (n is the number of specimens). 30. Combined mean cranial measurements (mm) of Scotophilus heathii 96 captured from Margalla Hills National Park (SA1), Chinji National

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Park (SA2), Lal Suhanara National Park (SA3) and Central Punjab (SA4) from June 2009 to May 2011 (n is the number of specimens). Combined mean bacular measurements (mm) of Scotophilus heathii 31. captured from Margalla Hills National Park (SA1), Lal Suhanara 97 National Park (SA3) and Central Punjab (SA4) from June 2009 to May 2011 (n is the number of specimens). Combined mean body mass (g) and external body measurements (mm) 32. of Scotophilus kuhlii captured from Margalla Hills National Park 101 (SA1) and Central Punjab (SA4) from June 2009 to May 2011 (n is the number of specimens). Combined mean cranial measurements (mm) of Scotophilus kuhlii 33. captured from Margalla Hills National Park (SA1) and Central Punjab 100 (SA4) from June 2009 to May 2011 (n is the number of specimens). Mean bacular measurements (mm) of Scotophilus kuhlii captured from 34. Pakistan Museum of Natural History (SA1) from June 2009 to May 104 2011 (n is the number of specimens). Combined mean body mass (g) and external body measurements (mm) 35. of Pipistrellus ceylonicus captured from Margalla Hills National Park 106 (SA1) and Central Punjab (SA4). Combined mean cranial measurements (mm) of Pipistrellus ceylonicus 36. captured from Margalla Hills National Park (SA1) and some localities 107 of Central Punjab (SA4) from June 2009 to May 2011 (n is the number of specimens). Combined mean bacular measurements (mm) of Pipistrellus ceylonicus 37. captured from Margalla Hills National Park (SA1) and Central Punjab 108 (SA4) from June 2009 to May 2011 (n is the number of specimens). Combined mean body mass (g) and external body measurements (mm) 38. of Pipistrellus javanicus captured from Margalla Hills National Park 110 (SA1), Chinji National Park (SA2) and Central Punjab (SA4) from June 2009 to May 2011 (n is the number of specimens). Combined mean cranial measurements (mm) of Pipistrellus javanicus 39. captured from Chinji National Park (SA2) and Central Punjab (SA4) 111 from June 2009 to May 2011 (n is the number of specimens). Combined mean bacular measurements (mm) of Pipistrellus javanicus 40. captured from SA1, SA2 and central Punjab SA4 from June 2009 to 112 May 2011 (n is the number of specimens). Combined mean body mass (g) and external body measurements (mm) 41. of Pipistrellus pipistrellus captured from Margalla Hills National Park 114 (SA1), Lal Suhanara National Park (SA3) and Central Punjab (SA4) from June 2009 to May 2011 (n is the number of specimens). Combined mean cranial measurements (mm) of Pipistrellus pipistrellus captured from Margalla Hills National Park (SA1), Chinji 42. National Park (SA2), Lal Suhanara National Park (SA3) and Central 115 Punjab (SA4) from June 2009 to May 2011 (n is the number of specimens).

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Combined mean bacular measurements (mm) of Pipistrellus 43. pipistrellus captured from Margalla Hills National Park (SA1), Chinji 117 National Park (SA2) and Central Punjab (SA4) from June 2009 to May 2011 (n is the number of specimens). Mean body mass (g) and external body measurements (mm) of 44. Pipistrellus tenuis captured from SA1 from June 2009 to May 2011 (n 118 is the number of specimens). Mean cranial measurements (mm) of Pipistrellus tenuis captured from 45. Margalla Hills National Park (SA1) from June 2009 to May 2011 (n is 119 the number of specimens). Mean bacular measurements (mm) of Pipistrellus tenuis captured from 46. Margalla Hills National Park (SA1) from June 2009 to May 2011 (n is 121 the number of specimens). Mean body mass (g) and external body measurements (mm) of 47. Hypsugo savii captured from Margalla Hills National Park (SA1) and 122 Chinji National Park (SA2) from June 2009 to May 2011 (n is the number of specimens). Mean cranial measurements (mm) of Hypsugo savii captured from 48. Margalla Hills National Park (SA1) from June 2009 to May 2011 (n is 124 the number of specimens). Bacular measurements (mm) of Hypsugo savii captured from SA1from 49. June 2009 to May 2011 (n is the number of specimens). 125

50. Social calls emitted by T. nudiventris and T. perforatus recorded at 128 Derawar Fort and Mojgarh in Lal Suhanra National Park (SA3). 51. Call parameters of some vespertilionid bats recorded from some arid 130 subtropical and tropical regions of Pakistan. Mean external body and cranial measurements of Taphozous 52. nudiventris (I = Roberts (1997), II = Bates and Harrison (1997), III = 148 Present study). Mean external body and cranial measurements of Taphozous 53. perforatus (I = Roberts (1997), II = Bates and Harrison (1997), III = 150 Present study). 54. Mean external body and cranial measurements of Scotoecus pallidus (I 151 = Roberts (1997), II = Bates and Harrison (1997), III = Present study). 55. Mean external body and cranial measurements of Scotophilus heathii (I 153 = Roberts (1997), II = Bates and Harrison (1997), III = Present study). 56. Mean external body and cranial measurements of Scotophilus kuhlii (I 154 = Roberts (1997), II = Bates and Harrison (1997), III = Present study). Mean external body and cranial measurements of Pipistrellus 57. ceylonicus (I = Roberts (1997), II = Bates and Harrison (1997), III = 156 Present study). 58. Mean external body and cranial measurements of Pipistellus javanicus 157 (I = Bates and Harrison (1997), II = Present study). 59. Mean external body and cranial measurements of Pipistellus 158 pipistrellus (I = Roberts (1997), II = Bates and Harrison (1997), III =

Present study). 60. Mean external body and cranial measurements of Pipistellus tenuis (I = 160 Bates and Harrison (1997), II = Present study). 61. Mean external body and cranial measurements of Hypsugo savii (I = 161 Bates and Harrison (1997), II = Present study). Descriptive parameters of echolocation calls of the Egyptian tomb bat (T. perforatus) from Pakistan. Explanation: n – number of individual calls analyzed (in parentheses number of call sequences from which 62. 166 calls were obtained); Fmax – start frequency; Fmin – end frequency; FmaxE – frequency with maximum energy (peak frequency); PD – pulse duration; IPI – inter-pulse interval; bold upper lines – mean±SD, lower lines – range.

LIST OF FIGURES Figure Page Title No. No. Map of Pakistan showing the location of 30 sampling sites where any bat roost was located or mist nets were erected in the four sub-areas. 1. 27 Of these seven were located in SA1, seven in SA2, five in SA3 and eleven in SA4. GPS location of five sites sampled for bats at Margalla Hills National Park (n = 7) and adjacent areas ( n =2) which jointly form SA1 during 2. the period extending from June 2009 to May 2011 (the map of 31 Pakistan showing the location of Margalla Hills National Park is given in the inset). GPS location of seven sites sampled for bats at Chinji National Park (n = 6) and adjacent areas ( n =1) which jointly form SA2 during the 3. 33 period extending from June 2009 to May 2011 (the map of Pakistan showing the location of Chinji National Park is given in the inset). GPS location of five sites sampled for bats at Lal Suhanra National Park and adjacent areas which jointly form SA3 during the period 4. 38 extending from June 2009 to May 2011 (the map of Pakistan showing the location of Lal Suhanra National Park is given in the inset). GPS location of eleven sites sampled for bats at Gujranwala (n = 2), Lahore (n = 3), Kasur (n =3) and Toba Tek Singh ( n = 3) which 5. jointly form SA1during the period extending from June 2009 to May 40 2011 (the map of Pakistan showing the location of Margalla Hills National Park is given in the inset). Dorsal (top), ventral (middle) and lateral (bottom) view of the bat skull showing definition of various cranial measurements (GTL = Greatest skull length, CBL Condylo-basal length, CCL = Condylo- canine length, ZB = Zygomatic breadth, BB = Breadth of the 6. 45 braincase, IC = interorbial constriction, PC = Postorbital contriction, 3 M = Mandible length, C-M = Maxillary toothrow length, C-M3= Mandibular toothrow length, M3-M3 = Posterior palatal width, C1-C1 = Anterior palatal length after Bates and Harrison, 1997). Dorsal (above) and lateral (below) view of the baculum of bat defining parameters recorded for measuring bacula of the bats (TLB = 7. Total bacular length, SL = Shaft length, PBL = Proximal branch 47 length, DBL = Distal branch length, PBW = Proximal branch width, DBW = Distral branch width, BH = Bacular height). Distribution map of bats belonging to Family Rhinolophidae recorded 8. 74 from four sub-areas from June 2009 to May 2011. Distribution map of bats belonging to Family Rhinopomatidae 9. 75 recorded from four sub-areas from June 2009 to May 2011. Distribution map of bats belonging to Family Emballonuridae 10. recorded from four sub-areas from June 2009 to May 2011. 76

Distribution map of bats belonging to Family Vespertilionidae 11. 77 recorded from four sub-areas from June 2009 to May 2011. Photograph of the bacula of T. nudiventris, captured from SA3 (a) 12. showing its shape and (b) position of baculum in penis of T. 83 nudiventris. Principal component analysis of the two Taphozous nudiventris populations recoded from SA1 and SA3 (Scree plot (a), scatter plot 13. 84 (b) and the Table of Eigen values (c) for the first four components is shown above. PC 1 loads size while PC 2 loads locality). (a) Baculum of T. perforatus captured from Sub-area 1 and (b) shows 14. 89 its position within the penis. 15. Dorsal view of the baculum of Scotoecus pallidus captured from SA3 93 Dorsal view of the bacula of Scotophilus heathii (1. Bat lab 47NARC, 2. Bat lab 4BFH, 3. Bat lab 8BFH, 4. Bat lab 13BFH, 5. Bat lab 14 16. 98 BFH, 6. Bat lab 39PMNH, 7. Bat lab 54Gojra, 8. Bat lab 114 RTown, 9. Bat lab 115 RTown, 10. Bat lab102 Pattoki). Principal component analysis of the four Scotophilus heathii populations recoded from SA1 (PMNH = P; Rawal Town = R; NARC = N), SA2 (Dalwal = D), SA3 (Bahawalpur Fish Hatchery =B) and 17. 100 SA4 (Gojra = G and Pattoki = Pt) Scree plot (a), scattered plot (b) and the Table of eigen values (c) for the first four components is shown above. PC 1 loads size while PC 2 loads locality). Dorsal view of the baculum of Scotophilus kuhlii captured from SA1 18. 104 from June 2009 to May 2011. Dorsal view of the bacula of Pipistrellus ceylonicus (a), P. javanicus 19. 126 (b), P. pipistrellus (c), P. tenuis (d) Hypsugo savii (e). 20. The sonograms of the flight calls of Taphozous spp. 129 21. Representative sonograms of Scotophilus heathii. 131 22. Representative sonograms of Pipistrellus spp. 132

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CHAPTER I

INTRODUCTION

This study on the diversity of bats (Suborder Vespertilioniformes) was conducted in four

sub-areas (SAs) that were located in tropical and arid sub-tropical regions of Pakistan. These

included the Margalla Hills National Park (SA1), the Chinji National Park (SA2), the Lal

Suhanra National Park (SA3) and four districts of central Punjab i.e. Gujranwala, Lahore, Toba

Tek Singh and Kasur that jointly form sub-area 4 (SA4). The former two National Parks i.e. the

SA1 and the SA2 are located in areas of Pakistan that are less pinfluenced by monsoons in the arid sub-tropical region while the third, the Lal Suhanra National Park (SA3) is situated in the tropical sand dune region of the Upper Indus Plains typified by Cholistan desert.

The Margalla Hills National Park (MHNP) was set up in 1979 to provide refuge to

Himalayan goral (Naemorhedus goral: Cetartiodactyla), Common Muntjac (Muntiacus muntjak:

Artiodactyla) and Leopard (Panthera pardus: Carnivora). The bat fauna of MHNP is not well

documented (Mahmood-ul-Hassan et al., 2009). Nawaz et al., (2007) have declared it home to

six bat species which include Pteropus giganteus, Rousettus leschenaultii, Eptesicus serotinus,

Myotis muricola, Megaderma lyra and the Scotophilus heathii but their sources remain

unconfirmed. Other sources state that MHNP is home to Hipposideros fulvus (Hinton and

Thomas, 1926, Sinha, 1980), S. heathii (Roberts, 1997), Scotophilus kuhlii (Hinton and Thomas,

1926), Myotis longipes (Ellerman and Morrison-Scott, 1951) and Pipistrellus tenuis (Roberts,

1997).

The Chinji National Park (CNP) was established in 1987 to conserve the Punja urial

(Ovis vignei punjabiensis: Artiodactyla) population and Olea ferruginea-Acacia modesta type

vegetation which are characteristic features of this park. Previous studies indicate that SA2

harbours Rhinopoma microphyllum (Lindsay, 1927ii), R. hardwickii (Lindsay, 1927iii),

Taphozous nudiventris (Linday, 1927ii, Roberts, 1997), Hipposideros cineraceus (Roberts,

1997), S. heathii (Roberts, 1997) and P. tenuis (Roberts, 1997).

The Lal Suhanra National Park was established in 1972 to save the blackbuck (Antelop

cervicapra: Cetartiodactyla) and Indian rhinoceros (Rhinoceros unicornis: Perissodactyla

populations that had gone extinct from this area and were later re-introduced in this park. Roberts

(1997) is the only source of information on the bat fauna of this National Park that reports T. nudiventris and S. heathii from the park.

Like the other three SAs, the bat fauna of the land area between Ravi and Chenab

“Rachna Doab” in which SA4 is located is also the least known. Dobson (1876) reported

Scotoecus pallidus from this area. Taber et al. (1967) conducted an extensive survey of a large part of SA4 and recorded S. heathii, Pipistrellus ceylonicus, P. mimus and P. kuhlii from this region. Other species recorded from SA4 include Scotozous dormeri (National Museum of

Natural History (USNM), Washington) in Bates and Harrison, 1997), Megaderma lyra

(Roberts, 1997), Rhinolophus blasii (Roberts, 1997), Rhinopoma microphyllum (Roberts, 1977)

and P. tenuis (Roberts, 1997).

Bats constitute 28% of the mammalian fauna of the country (Roberts, 1997) but the exact

number of chiropteran taxa occurring within Pakistan has been a matter of debate (Roberts,

1997; Bates and Harrison, 1997; Walker and Molur, 2003; Wilson and Reeder, 2005). Although

54 species of bats have been reported from the country, many new taxa can be identified if the

cryptic bat species are accurately identified as it has been done in other parts of the world (see

e.g. Jones and van Parijs, 1993; Davidson-Watts et al., 2006). The biology and ecology of almost

all bat species in Pakistan is poorly known. At present, there is no specialized bat biologist in the

country and there is a serious lack of taxonomic capacity to identify bats on the basis of their

morphological features. There are only a few who are either interested or properly equipped to undertake taxonomic work on bats. Similarly, bats are rarely considered in either environmental policies or educational projects. In many cases, the only information about a species is based on the original description since it has not been collected subsequently (Mahmood-ul-Hassan et al.,

2009).

According to the reports from International Union for the Conservation of Natural

Resources (IUCN), bats are facing substantial threats from habitat loss and fragmentation all over the world. Species with relatively small geographic ranges and those that are ecologically specialized tend to be at greatest risk (Jones et al., 2003). Given their importance as top order predators and pest control agents, their value as bio-indicators and their vulnerability (Jones et al., 2009), monitoring bats to assess population trends over longer time scales is urgently needed.

Since, after the last couple of centuries no long-term studies on bats is available from the region where Pakistan is located today, the present study was designed to explore bat biodiversity

(Vespertilioniformes: Order Chiroptera) in some tropical and arid sub-tropical regions of

Pakistan. The specific objectives of this study are as follows.

Objectives

1. To make an extensive survey of the study area to explore the diversity of various chiropteran species belonging to order Vespertilioniformes.

2. To re-define geospatial distribution and habitat preferences of various vesper bats of study area.

3. To describe the morphology, external body, cranial and bacular measurements of various vesper bats inhabiting the study area.

4. To record and analyze echolocation calls of these bat species.

CHAPTER II

REVIEW OF LITERATURE

Mammals in general exhibit a host of adaptations which enable them to exploit a variety of habitats on land, water and air (Kalko, 1997; Ramirez-Pulido et al., 2005). Chiroptera, one of the

26 mammalian orders, is remarkable for its high diversity and broad geographic distribution

(Simmons, 2005). Bats comprise 1150 recognized extant species (Schipper et al., 2008) and are major contributors to mammalian biodiversity (Meyer et al., 2010). They constitute almost 20% of mammalian species worldwide (Simmons, 2005). Bats possess a multitude of features that craft them excellent bioindicators of human-induced changes and are best suited to gauge climate change and habitat quality across the globe due to their cosmopolitan distribution combined with high functional and taxonomic diversity (Jones et al., 2009). Moreover, many species fulfill key ecosystem services, particularly in tropical ecosystems, as pollinators, seed dispersers, and control agents of arthropod populations (e.g. Kalka et al., 2008; Kelm et al., 2008; Lobova et al.,

2009; von Helversen and Winter, 2003; Williams-Guillén et al., 2008; Mahmood-ul-Hassan et al., 2010). Bats also have an excellent ability to respond to a wide range of global phenomena and environmental stressors such as urbanization, agricultural intensification, forest disturbances

(e.g. logging, oil extraction), habitat loss and fragmentation, global climate change, and overhunting (Meyer et al., 2010). Finally, the responses of bats to habitat disturbance are often associated with those of other taxa (e.g. Bass et al., 2010; Jones et al., 2009). They are the only mammals capable of a sustained flight crossing physical barriers that the other mammals cannot

(Fenton, 1992; Wilson and Reeder, 1993; Hutson et al., 2001). They are polyphagous in their food niche and range from primary consumers such as seed, pollen, leaf or fruit eaters (e.g.

Pteropus, Rousettus and Cynopterus spp.) to top predators in certain ecosystems and consume

4 arthropods small mammals (e.g. Megaderma lyra) and even fish (e.g. Myotis vivesi) (Beolens et al., 2009; Fenton, 1992; Hill and Smith, 1985) .

Once the order Chiroptera was sub-divided into two sub Orders - the Megachiroptera and the Microchiroptera (Koopman, 1993; Bates and Harrison, 1997; Wilson and Reeder, 2005). The former was represented by only one family (Pteropodidae) restricted only to the Old World tropics of Africa and Asia while the latter included 17 families (Rhinopomatidae,

Emballonuridae, Craseonycteridae, Nycteridae, Megadermatidae, Rhinolophidae,

Hipposideridae, Noctilionidae, Mormoopidae, Phyllostomidae, Natalidae, Furipteridae,

Thyropteridae, Myzopodidae, Vespertilionidae, Mystacinidae and Molossidae) (Mahmood-ul-

Hassan and Nameer, 2006). However, a recent molecular phylogenetic and phylogeographic analysis of the bats has changed the above taxonomic perspective leading to a new arrangement of the Order Chiroptera. Based on modern analytical techniques Dr. Emma Teeling and her team have proposed an entirely new classification of chiropterans (see summary in Jones and Teeling,

2006). The megabats (Megachiroptera) are now grouped with rhinolophoids and their relatives

(families Rhinolophidae, Hipposideridae, Craseonycteridae, Rhinopomatidae and

Megadermatidae) into the suborder Yinpterochiroptera while the remaining bat families constitute the subrder Yangochiroptera (Mahmood-ul-Hassan et al., 2009). Hutcheon and Kirsch

(2006) suggest that Peteropodiformes is a more appropriate name for the “Yinpterochiroptera” and Vespertilioniformes for Emballonuridae, Nycteridae and the “Yangochiropterans” which include Myzopodidae, Mystacinidae, Phyllostomidae, Mormoopidae, Noctilionidae,

Furipteridae, Thyropteridae, Natilidae, Molossidae, Miniopteridae and Vespertilionidae. The

Vespertilionidae (403 species) and the Phyllostomidae (160 species) are the largest families with a worldwide distribution found in the Nearctic and Neotropical regions (Hutson et al., 2001).

The Order Vespertilioniformes is represented by three families, seventeen genera and thirty three

species in Pakistan (Mahmood-ul-Hassan et al., 2009).

The bats are well acknowledged for their ecological services throughout the Europe and

America (Mickelburgh et al., 1992; Fujita and Tuttle, 1991). However, the importance of the positive role of bats in the ecosystem in Southeast Asia was recognized in 1998 when the

Malaysian government passed a wildlife protection ordinance which included protection of all

species of bats (Gumal and Racey, 1999). Bates and Harrison (1997) made the most comprehensive and up-to-date revision of the Chiroptera of the Indian subcontinent and enlisted

119 species of bats belonging to eight families and 37 genera from India, Pakistan, Nepal, Sri

Lanka, Maldives, Afghanistan, Tibet, and Northern Myanmar. Many of these bat species are now striving hard to survive in the region. Out of these 119 species, 110 are recorded within the

Indian limits (Bates and Harrison, 1997). As far as their status in the world is concerned, 51

species are least concerned, one is data deficient and one is near threatened (International Union

for the Conservation of Nature, 2011).

In India, two species of bats viz., the Wroughton’s free-tailed bat (Otomops wroughtonii) and Salim Ali’s fruit bat (Latidens salimalii) are highly protected and are on schedule 1 of wildlife (protection) act 1972. Furthermore, persistent efforts by globally renowned bat biologists and non-governmental organizations in India have resulted in providing legal protection to all 13 species of pteropodid bats (Singaravelan et al., 2009; Mahmood-ul-Hassan,

2010).The realization of the role of bats in the agricultural economy of India can be documented from the fact that owing to its feeding habits in Bihar, the Indian false vampire (Megaderma lyra) is considered as a good friend of farmers who reward it by food in bad weather and call it goddess Laxmi (Sinha, 1986). Bats are given no protection by law in Pakistan and many species

are hunted for their body fat to be used by local health practitioners to cure rheumatic pains

(Roberts, 1997).

Pakistan has a rich bat biodiversity which stems from its unique geographic position in

the world. The bat fauna of the country is composed of a blend of three of the world’s six

biogeographic regions – the Oriental, the Palaearctic and the Ethiopian. Pakistan lies at the western end of the South Asian subcontinent which is a suture between the Indo-Malayan and the Palaearctic regions whereas it is connected to the Ethiopian region through a land mass towards the southwest. The Indus demarcates a boundary between the Indo-Malayan and

Palaearctic regions as Indo-Malayan forms are predominant in the east of the Indus while the mountains of the north and west hold the Palaearctic forms. The Palaearctic species include a mixture of those common to a large part of Eurasia, along with resemblances to the Middle East,

West Asia (Afghanistan and Iran), Central Asia and Tibet. The unification elements from different origins ensure a unique and diverse mixture of fauna and flora (Roberts, 1997;

Mahmood-ul-Hassan et al., 2009).

Anthropogenic activities are adversely affecting biodiversity at all scales. Habitat degradation and fragmentation have specifically brought many bat species towards the risk of extinction. Long term monitoing of the bats is thus necessary to understand negative human impact on bat diversity in the world. (Meyer et al., 2010).

The changed patterns of rainfall, humidity and temperature along with anthropogenic changes in the habitat have altered distribution ranges of many bat species over the past few dacades. Pereswiet-Soltan (2007) on the basis of Mediterranean, temperate, continental, hemiboreal and milder climatic zones differentiated European bat fauna into three categories i.e. thermophilic species, species adapted to milder climatic conditions and species of cold climate.

He found that a certain structure and abundance of bat fauna correspond to each climatic zone.

From the three groups, those of milder climate have extended their distribution latitudinally, in the West of the Palaearctic Region. The tropical landscapes are currently fragmented to varying degrees. They either have gaps or have had their natural vegetation replaced. Thus, in tropical landscapes consist of different sizes of habitat fragments of natural or anthropogenically modified vegetation (i.e., crops, pastures, human settlements, and secondary vegetation) to which the movement of many bat species is restricted. Such changes do not affect all species equally as some are favor by these modifications while others are not (Moreno and Halffter, 2001).

Bat conservation is impossible without accurate species identification. Worldwide, scientists are using a multitude of techniques for enlisting bat faunas of different regions.

Although recent advances in molecular techniques have provided tools for investigating phylogenetic variations between morphologically similar species (Russo et al., 2006), these techniques are expensive and results are not immediately available (Weller et al., 2007).

Analysis of the echolocation call characteristics have also been used to distinguish sympatric bat species (Jones and van Parijs, 1993; Jacobs et al., 2006) but the bat detectors are too expensive to be purchased in many developing countries. It is due to these reasons that bat biologists in most parts of the world, especially in the underdeveloped countries, are using characters such as forehead slope, dorsal pelage sheen, and behavior of the bats to discriminate species (Harris,

1974; Nagorsen and Brigham, 1993; Verts and Carraway, 1998). Bat identification on the basis of external morphology and measurements of different skull parameters (Hill and Smith, 1984;

Vaughan et al., 2000; Jacobs et al., 2006) is still a highly reliable technique in most instances.

Use of character matrices and identification keys are authentic tools to identify different chiropteran species (Daniel, 2009; Srinivasulu et al., 2010).

In many instances morphological species identification has been confirmed by modern molecular techniques. Mitochondrial cytochrome b sequences in Miniopterus schreibersii (Kuhl,

1817) from the Oriental-Australasian areas revealed that the Oriental-Australasian M. schreibersii diverged from the Spanish M. schreibersii. The Chinese/Japanese specimens were also separated from the Australian specimens. These molecular results were in conformity with a previous report based on morphological characters by Maeda (1982) which suggested that

Miniopterus schreibersii in Europe, Asia and Australia should be regarded as three distinct species, named Miniopterus schreibersii, M. fuliginosus and M. oceanensis (Tian et al., 2004).

Bacular morphology has been successfully used in the taxonomy of some mammalian groups especially when the species are difficult to differentiate on the basis of general morphology (e.g. Didier, 1954; Rabeder, 1976; Strelkov, 1989a; Baryshnikov and Abramov,

1997; Benda and Tsytsulina, 2000). More than 160 Palaearctic vespertilionids viz. Plecotus,

Barbastella, and Myotis (Selysius subgenus) which are morphologically very similar have been differentiated on the basis of this attribute (Thomas, 1915; Chaine, 1926; Topal, 1958; Hill and

Harrison, 1987; Strelkov, 1989a; 1989b; Yoshiyuk, 1989; Smirov, 2000; Strelkov, 1989a, 1989b;

Benda and Tsytsulina, 2000; Kruskop and Lavrenchenko, 2000; Tsytsulina, 2001). Taphozous nudiventris magnus from T.n. kachhensis are also differentiated through this distinctive feature

(Asan and Albayrak, 2007). Baculum length in various bat species is positively associated with relative male body mass but this association has not been proved substantially through phylogenetic testing (Hosken et al., 2001). Age peculiarities of baculum shape have been studied in detail in Rodentia and Carnivora (e.g., Friley, 1949; Callery, 1951; Heidt, 1970;

Tarasov, 1974, 1984; Aksenova, 1980) but studied only sparingly in Chiroptera (Maeda, 1978).

Correlations between increasing condylobasal length, forearm length and baculum length were

found in Nyctalus aviator (Maeda, 1978) Murina silvatica, Vespertilio superans, Myotis

macrodactylus, and Rhinolophus cornutus (Yoshiyuk, 1989) but V. superans aged from 10 to

202 days showed no such relationship (Yoon et al., 1990). The comparison of the shape and size of bacula in Myotis myotis and Myotis blythi depicted that it is not the size but shape of the baculum that distinguishes these two species (Albayrak and Asan, 2001).

Almost all of the Pteropodiformes (with the exception of most pteropodids) and all

Vespertilioniformes use echolocation calls emitted through the mouth or nose to locate and hunt prey (Altringham, 1996). Effective monitoring of echolocation calls is important in the studies

on the ecology and conservation of bats. The structure of echolocation calls varies because of the

influences of morphology, age, geographical variation, foraging habitat and foraging mode. It

varies not only between species, but also within species. Ultrasonic bat detectors have been

increasingly used to study bat behaviors (Feng et al., 2000; Zhang et al., 2000; Jones et al.,

2006). Moreover, more and more researchers are using bat detectors to record echolocation calls

and to classify bats based on known calls (Fukui et al., 2004; Rydell et al., 2002; Vaughan et al.,

1997b).

Jones et al. (1993) studied echolocation call structure, flight morphology and feeding

behavior of three hipposiderid bats (Astellia tridens, Hipposideros caffer and H. ruber:

Chiroptera: Hipposideridae). They found that all the species emitted brief CF/FM (Constant

frequency/Frequency Modulated) echolocation calls. The frequency of the calls varied mainly by

sex and age differences, juveniles used lower frequencies than adults and males were lower in

frequency than females. In addition it was found that among adults CF frequency was related to

forearm length in a polynomial manner.

Sun et al. (2008) recorded one hundred and thirty-eight echolocation calls of 63 free- flying individuals of five bat species (Rhinolophus ferrumequinum, Myotis formosus, Myotis ikonnikovi, Myotis daubentoni and Murina leucogaster) by using an ultrasonic bat detector

(D980) in Zhi’an village of Jilin Province, China. According to the frequency-time spectra, these calls were categorized into two types: Frequency Modulated/Constant Frequency (FM/CF) (R. ferrumequinum) and FM (M. formosus, M. ikonnikovi, M. daubentonii and M. leucogaster).

Sonograms of the calls of R. ferrumequinum could easily be distinguished from those of the other four species. For the calls of the remaining four species, six echolocation call parameters, including starting frequency, ending frequency, peak frequency duration, longest inter-pulse interval and shortest inter-pulse interval, were examined by stepwise discriminant analysis. The results show that 84.1% of calls were correctly classified, which indicates that these parameters of echolocation calls play an important role in identifying bat species. These parameters can be used to test the accuracy of general predictions based on bats’ morphology in the same forest and can provide essential information for assessing patterns of bat habitat use.

Russo and Jones (2002) described spectral and temporal features of echolocation calls emitted by 22 bat species (three rhinolophids, 18 vespertilionids and the molossid Tadarida teniotis) from Italy. Time expanded recordings of calls from 950 bats of known identity were examined. Rhinolophus ferrumequinum, R. hipposideros, R. euryale and T. teniotis could be identified by measuring the call frequency of highest energy.

O’Farrell (1997) identified nineteen bat species on the basis of differences in the time/frequency characteristics of their echolocation calls. Species foraging in open appear to use loud calls that can be detected at a greater distance than species that forage in clutter and use calls of low intensity.

The variation of echolocation calls can cause the researcher to sometimes incorrectly

identify the species of bats according to calls. Moreover the calls which are recorded in one

region or in a habitat may not be applicable to other regions, or to other habitats (Fukui et al.,

2004).

All the echolocating bats use a variety of roosting and foraging habitats. Forest habitats

of both the tropical and temperate areas provide them excellent refuge. They often prefer to roost

in the vicinity of aquatic habitats which are the breeding grounds of the most insect species.

Buildings, crevices of walls and ceilings, lofts and chimneys of the old abandoned buildings and

disused wells are the commonest bat roosts. Bark and hollows of the old trees in sub-urban and peri-urban areas also provide them a secure refuge (Roberts, 1997; Anonymous, 2010).

The tropical landscapes are currently fragmented to varying degrees and have gaps or have had their natural vegetation replaced. Thus, in these landscapes there are different sizes of habitat fragments of natural or anthropogenically modified vegetation (i.e., crops, pastures, human settlements, and secondary vegetation) to which the movement of many species of both plants and animals is restricted. These changes do not affect all species equally since some are favored by these modifications (Moreno and Halffter, 2001).

Out of 24 microchiropteran speies previously known from Singapore only 15 were observed indicaing a rapid decline of species diversity in the country Pottie et al. (2005). Purohit and Vyas (2006) randomly sampled bats from various roosting sites in and around Jodhpur, the

Great Indian Desert, to explore the sex ratio of different species of bats. Females were dominated in the population of Rhinopoma microphyllum, R. hardwickii, Taphozous nudiventris and

Rhinolophus lepidus while in Pteropus giganteus, Taphozous perforatus and Scotophilus heathii

males dominated. Male and females shared equal percentages in populations of Pipistrellus

tenuis of this study area.

Loss of their natural habitat by increased human population and human activities such as

deforestation, use of pesticides, industrial activities, loss of buildings or alteration in the design of their roofs and deliberate anthropogenic disturbance are the major causes of their population decline throughout the world. Even the minor alterations in the habitat such as the loss of key

landscape elements for example tree lines, hedgerows, and canals that are used regularly by bats

during flight result in the abandonment of their roosts and maternity colonies. On the basis of

these qualities, bats in Pakistan may also serve as good indicators of ecosystem health (Jones et

al., 2009).

SPECIES ACCOUNTS

Rhinolophus blasii is considered to be a marginal species in Pakistan and perhaps represented in

the country by its subspecies R. b. meyeroemi (Corbet and Hill, 1992). The population status of

the species in the Indian subcontinent is unknown (Bates and Harrison, 1997). Z. B. Mirza and T.

J. Roberts collected a single specimen of the species in 1968 from Lahore (Roberts, 1997) which is preserved in the museum of the Department of Zoology at the University of the Punjab,

Lahore. The species was included in “Lower Risk: near threatened’ in the 1996 IUCN Red List

of threatened Animals (Baillie and Groombridge, 1996). The species is considered to be in

Lower Risk category (IUCN, 2003), near threatened in South Asia (Walker and Molur, 2003)

and Least Concern (IUCN, 2008; Mahmood-ul-Hassan et al., 2009).

Body measurements. Body and cranial measurements of the specimens captured from the area

averaged as head and body length 33 mm, forearm length 41 mm, tail length 21 mm and ear

length 15 mm (Roberts, 1997). Bates and Harrison (1997) have described the morphometrics of

the specimens from Iran (after DeBlase, 1980) having forearm length 47. 8 mm (43 – 50 mm),

tail length 26.6 mm (21.0 – 35. 0 mm), hind foot length 13.3 mm (8.0 – 11.0 mm) and ear length

20.0 mm.

Cranial measurements. Various cranial measurements of R. blasii captured from the study area averages as greatest skull length 19.7 mm (19.1 - 20.1 mm), condylocanine length 16.5 mm (16.0

– 17.0 mm), zygomatic breadth 9.2 mm (8.8 – 9.5 mm), breadth of the braincase 8.6 mm (8.2 –

9.1 mm), post orbital constriction 2.3 mm (2.2 – 2.8 mm), mandibular length 12.2 mm (11.9 –

12.7 mm) (Deblase, 1980).

Rhinopoma hardwickii has been reported from Rohtas in Salt Range, from around Karachi and

Landhi in southern Sindh and Karchat Hills near Hyderabad. The species is rare and locally distributed. This bat uses the same type of diurnal roosts as R. microphyllum. The species is considered to be in “Low Risk category” (Mahmood-ul-Hassan et al., 2009).

Body measurements. Nine specimens of R. hardwickii were captured from Sindh and northern

Punjab. The average head and body length ranged from 55-69 mm (mean 62 mm), tail length 67

mm long (ranged between 57-77 mm), hind foot and ear length averaged 11.5 mm (range 9-15

mm) and 18 mm (range 16-20 mm) respectively. The average forearm length was found to be 60

mm (range 60–67 mm).

Bates and Harrison (1997) collected specimens of R. harrdwickii from Pakistan. The average

head and body length was 66.6 mm (55.0 – 73.0 mm), tail length 66.8 mm (56.0 – 78.0 mm),

hind foot length 13.4 mm (11.0 – 15.0 mm), forearm length 59.2 (52.9 – 64.0 mm), ear length

19.3 mm (17.0 – 21.0).

Cranial measurements. Cranial measurements of the R. hardwickii specimens captured from

India and Pakistan had average greatest skull length 18.7 mm (17.5 – 19.7 mm), condylocanine

length 16.5 mm (15.5 – 17.5 mm), zygomatic breadth 10.9 mm (10.1 – 11.7 mm), breadth of the

braincase 8.2 mm (7.8 – 8.5 mm), postorbital constriction 2.8 mm (2.5 – 3.2 mm) and average mandible length was noted as 12.8 mm (11.8 – 13.6 mm) (Bates and Harrison, 1997).

Taphozous nudiventris is a very large bat without any clavicular collar of paler hairs as born by

Taphozus perforatus and have a well developed gular pouch in adult males consisting of a crescentic flap of skin about 10-11 mm wide across its orificeis. It is a typical sheath-tailed bat, with a dog-like face, long semi-naked muzzle, without noseleaf, comparatively large eyes and widely spaced triangular ears. Adult female specimens can be recognized by the presence of a crescentic mark (Roberts, 1997; Bates and Harrison, 1997). The species is distributed in

Mauritania, Senegal, and Guinea-Bissau to Djibouti, Egypt, Jordan, NE Turkey, south to

Tanzania and east to Burma.

In Pakistan this is wide spread and common. In Punjab, specimens have been collected from

Multan (Roberts, 1997), Salt Range (Lindsay, 1927; Roberts, 1997) and Bahawalpur (Roberts,

1997). It occurs throughout Sindh from Sukker and Kahairpur in the north to Jaccobabad and

Thatta in the south (Wroughton, 1916; Siddiqui, 1961; Roberts, 1997) however, the species has

not been collected from any mountainous region of Baluchistan or Khyber Pakhtunkhwa

(Roberts, 1997; Mahmood-ul-Hassan et al., 2009). The species is kept in “Least Concern”

category (IUCN, 2008).

Body measurements. Thirteen specimens of T. nudiventris were captured from Punjab and

Sindh provinces. Their average forearm length was 71 mm, head and body length 92 mm (range

86-98 mm), tail length 36.5 mm (range 34-42 mm), hind foot 14 mm (range 11-18 mm), and the

ear 22 mm (range 19-23 mm). The maximum body mass recorded was 50 g (Roberts, 1997).

According to Bates and Harrison (1997) the body measurements of the specimens captured from

India were as average head and body length 96.3 mm (90.0 – 105.0 mm), tail length 32.6 mm

(22.0 – 46.0 mm), hind foot length 14.6 mm (11.0 – 18.0 mm), forearm length 74.3 (71.0 – 80.0

mm), ear length 22.0 mm (18.0 – 25.0).

Cranial measurements. Various cranial measurements T. nudiventris samples were averaged as

the greatest skull length 25.8 mm (22.5 – 27.9 mm), condylocanine length 23.4 mm (21.6 – 25.6

mm), zygomatic breadth 15.9 mm (14.4 – 17.8 mm), breadth of the braincase 11.5 mm (10.6 –

12.7 mm), postorbital constriction 8.1 mm (7.0 – 8.6 mm),mandible length 20.1 mm (18.2 – 21.5

mm) (Bates and Harrison, 1997).

Taphozous perforatus is smaller in size than T. nudiventris and can be differentiated in the field

by the presence of fur both on the dorsal and ventral side of the body and extending up to the tip

of the tail. Another important feature is the presence of a radialo-metacarpel pouch and traces of

a gular pouch. The wings are long and narrow and the tips are intricately folded inward when the

bat is at rest. The species is distributed in Mauritania and Senegal to Botswana, Mozambique,

Somalia, Djibouti, and Egypt; S Arabia; Jordan; S Iran; Pakistan; NW India. This species is uncommon and limited in distribution in Pakistan. The species was recorded from Jatti near the

Thatta district by the Zoological Survey of Pakistan (Siddiqui, 1961). This species has not been

recorded from elsewhere in Pakistan (Roberts, 1977). The species is not threatened currently but

may be vulnerable to human disturbance of roosts (Bates et al., 1994b) IUCN categorizes the

species as “Least Concern” (Mahmood-ul-Hassan et al., 2009).

Body measurements. The average forearm length varies from 59-63 mm (mean 62.3 mm) with

the tail length 21-27 mm. The hind foot and ear length averages 11.5 mm and 18 mm

respectively (Roberts, 1997).

Bates and Harrison (1997) collected T. perforatus samples from India. Their head and body length averaged 74.5 mm (71.0 – 80.0 mm), tail length 24.0 mm (20.0 – 28.0 mm), hind foot length 10.1 mm (8.2 – 12.5 mm), forearm length 61.3 (59.2 – 63.8 mm), length of 3rd metacarpal

55.4 (53.7 – 57.2 mm), length of 1st phalanx of 3rd metacarpal 19.0 mm (18.3 – 19.7 mm), ear length 18.0 mm (14.0 – 20.0).

Cranial measurements. Various cranial measurements recorded average greatest skull length

20.5 mm (19.9 – 21.5 mm), condylocanine length 19.0 mm (18.4 – 19.7 mm), zygomatic breadth

11.7 mm (11.5 – 12.1 mm), breadth of the braincase 9.3 mm (9.2 – 9.6 mm), interorbital constriction 5.8 mm (5.5 – 6.2 mm), mandible length 15.0 mm (14.6 – 15.6 mm) (Bates and

Harrison, 1997).

Scotoecus pallidus is endemic to the Indian subcontinent and has a local and restricted distribution in Pakistan. It was first described by Dobson in 1867 from a specimen collected from

Mian Mir (Lahore). Further collections were made from different regions of northern Sindh

(Kashmore and Mirpur in Jacobabbad, Larkana, Sukker and Dadu Districts) and Punjab

(Muzaffargarh and Sialkot). Its population status is uncertain and deserves further study

(Mahmood-ul-Hassan et al., 2009).

Body measurements. The average head and body length of the S. pallidus was recorded as 54 mm, tail length 37 mm, hind foot length 8 mm, ear length 13 mm (Roberts, 1997). While the average head and body length was measured as 52.8 mm, tail length 36.9 mm, hind foot length

8.3 mm, forearm length 36.2 mm, 5th metacarpal length 33.7 mm, 4th metacarpal length 34.2 mm,

3rd metacarpal length 34.6 mm, ear length 12.8 mm (Bates and Harrison, 1997).

Cranial measurements: Various cranial measurements include average greatest skull length

15.1 mm, condylocanine length 14.1 mm, zygomatic breadth 10.5 mm, breadth of the braincase

7.7 mm, post-orbital constriction 4.3 mm and mandibular length 11.4 mm (Bates and Harrison,

1997).

Scotophilus heathii is geographically distributed in Afghanistan to South China, including

Hainan Isl, south to Sri Lanka, Vietnam, Cambodia, Thailand and Burma. In Pakistan the species

is common and widespread throughout the Indus plains. It has been collected from Kohat

(NWFP), Islamabad city, Multan, Lahore and Sialkot districts (Punjab), Kashmoor, Sakkur,

Jacobabad, Mirpur Sakro, Dadu, Landi, Malir, Karachi (Sindh) (Wroughton, 1916a; Lindsay,

1926; Siddiqui, 1960; Taber et al., 1967; Walton, 1974; Roberts, 1997). IUCN categorizes the

species as “Least Concern” (IUCN, 2008).

Body measurements. Walton (1974) captured twenty six specimens of S. heathii from Malir,

Karachi and Mirpur Sakro. The morphometric measurements include average head and body length 60 mm, tail length 13 mm and forearm length 58 mm. Roberts (1997) collected six bat specimens from south-west Punjab and Karachi and mentioned various morphometric measurements of six bat specimens as head and body length averaged 55 mm (range 51 – 60 mm), tail length 12 mm (range 11 – 13 mm), ear length 16 mm (range 14 – 17 mm).

Bates and Harrison (1997) collected data from Pakistan, India and Sri Lanka. Average head and body length of sixty three bat specimens was found to be 82.5 (range 67.0 – 93.0), tail length of these sixty three specimens was (59.1 43.0 – 71.0), hind foot length of fifty eight S. heathii bat

specimens was found to be 12.0 (9.0 – 15.0 mm), forearm length of sixty four specimens was

averaged as 60.7 (range 55.4 – 65.8 mm), length of the 5th metacarpal of sixty one specimens

averaged 54.6 mm (range 50.3 – 59.9 mm), length of the 4th metacarpal of sixty one bat

specimens was 58.2 mm (range 54.0 – 63.9 mm), average length of 3rd metacarpal of sixty one

specimens was found to be 59.4 mm (range 53.7 – 68.4 mm), ear length of sixty three bat

specimens averaged 16.9 mm (range 13.0 – 20.2 mm).

Cranial measurements. Average greatest skull length of sixty three specimens was measured as

23.4 mm (range 21.7 – 25.2 mm), condylocanine length 20.2 mm (19.0 – 21.3 mm), zygomatic breadth of sixty two specimes was counted as 15.6 mm (14.5 – 16.9 mm), average breadth of the braincase 10.2 mm (9.7 – 10.8 mm), post-orbital constriction of sixty six bat specimens 5.5 mm

(range 5.2 – 5.9 mm), mandibular length of sixty six S. heathii specimens averaged 16.3 (range

14.8 – 18.0).

Scotophilus kuhlii is uncommon in Pakistan with a very restricted distribution. The species is present only in southern Sindh (Roberts, 1997). Geographiccaly this species is distributed in

Bangladesh, Pakistan to Taiwan, south to Sri Lanka, Burma, Cambodia, W Malaysia, Java, Bali,

Nusa Tenggara (Indonesia), southeast to Philippines and Aru Isles (Indonesia) (Mahmood-ul-

Hassan et al., 2009). The species is categorized as ‘Least Concern’ (IUCN, 2008).

Body measurements. Walton (1974) captured S. kuhlii specimens from Mirpur Sakro and mentioned various morphometric measurements of three bat samples. Their average head and body length was 49 mm, tail length 13 mm, forearm 49 mm and the ear length was 17 mm.

Roberts (1997) collected bat samples from Karachi. The average morphometric measurements of four specimens have an average head and body length of 52 mm (range 47 – 58 mm), tail length

11 mm and ear length was 17 mm.

Bates and Harrison (1997) collected data from Pakistan, India and Sri Lanka. The average head and body length was measured as 69.8 mm (47) (range 60.0 – 78.0 mm), tail length, 47.5 mm

(46) (range 40.0 – 65.0 mm), hind foot length 10.0 mm (33) (range 8.0 – 13.0 mm), forearm length 49.0 mm (47) (range 44.0 – 56.4 mm), length of 5th metacarpal length 45.0 mm (41) (42.1

– 53.9 mm), 4th metacarpal length 48.3 mm (43) (range 43.7 – 57.2 mm), 3rd metacarpal length

48.8 mm (47) (range 44.4 – 58.8 mm), ear length 13.5 mm (46) (range 9.0 – 17.0 mm).

Cranial measurements. The cranial measurements include average greatest skull length was

19.6 mm (49) (range 18.7 – 20.4 mm), condylocanine length 17.3 mm (49) (range 16.3 – 18.0

mm), zygomatic breadth 13.0 mm (43) (range 12.4 – 13.7 mm), breadth of the braincase 8.9 mm

(49) (range 8.8 – 9.4 mm), post-orbital constriction 4.7mm (49) (4.4 – 5.1 mm) and mandibular

length 13.7mm (48) (12.9 – 14.4 mm).

Pipistrellus ceylonicus seems to be common in Pakistan. The species is particularly abundant

around Karachi and Thatta districts in Sindh. It is likely to be present in warmer southern

latitudes in the Indus plains (Mahmood-ul-Hassan et al., 2009). Taber et al. (1967) collected it from Lyallpur (Faisalabad) and Khanewal. It is relatively large sized pipistrelle. Its ears, naked areas of the face, wings and inter femoral membranes are uniform dark brown. The adult body mass ranges between seven to eight grams (Madhavan, 1971). The muzzle in this species often has glandular swellings between the eyes (Roberts, 1977). This species is distributed in Pakistan,

India, Sri Lanka, Bangla Desh, Burma, Kwangsi and Hainan (China), Vietnam, Borneo

(Mahmood-ul-Hassan et al., 2009). This species is categorized as “Least Concern” (IUCN,

2008).

Body measurements. Roberts (1997) measured five specimens and found that the head and body length averaged 51 mm (range 48- 56 mm) and the average tail length was 38 mm (range

36-44 mm). The hind foot and ear length averaged 9 mm and 12 mm respectively. Average forearm length was 35 mm. The body measurements recorded by Bates and Harrison (1997) are as follows. Head and body length is 53.5 mm (45.0 – 64.0 mm), tail length 38.2 mm (30.0 – 45.0 mm), hind foot length 8.3 mm (6.0 – 11.0 mm), forearm length 37.2 (33.0 – 42.0 mm), wingspan

251.1 (227 – 262 mm), length of 5th metacarpal 33.6 (30.7 – 36.7 mm), length of 4th metacarpal

35.1 (32.6 – 38.8 mm), length of 3rd metacarpal 35.8 (33.0 – 39.5 mm), ear length 12.2 mm (9.5

– 14.0).

Cranial measurements. Greatest skull length 15.0 mm (14.4 – 15.8 mm), condylocanine length

13.7 mm (13.1 – 14.3 mm), zygomatic breadth 9.8 mm (9.2 – 11.0 mm), breadth of the braincase

7.3 mm (6.8 – 7.8 mm), postorbital constriction 4.0 mm (3.7 – 4.3 mm), mandibular length 11.2

mm (10.6 – 12.0 mm) (Bates and Harrison, 1997).

Pipistrellus javanicus is distributed in East Afghanistan, North Pakistan, North and Central

India, South and East Tibet (China), Burma, Thailand, Vietnam, Through SE Asia to Lesser

Sunda Isles and Philippines; perhaps Australia. No literature is available on the distribution of

this species in Pakistan however a single specimen was collected from Gharial, Murree Hills

(Mahmood-ul-Hassan et al., 2009). The species falls in ‘Least Concern’ category (IUCN, 2008).

Body measurements. The average head and body length of specimens collected from Pakistan,

India, Nepal and Bangladesh is 47.1 mm (40.0 – 55.0 mm), tail length 33.9 mm (26.0 – 40.0 mm), hind foot length 6.0 mm (3.0 – 8.0 mm), forearm length 33.2 (30.0 – 36.0 mm), length of

5th metacarpal 31.4 (29.0 – 33.4 mm), length of 4th metacarpal 32.6 (29.9 – 34.7 mm), length of

3rd metacarpal 33.0 (25.9 – 34.8 mm) and the ear length was 11.8 mm (5.0 – 15.0 mm) (Bates and Harrison, 1997).

Cranial measurements. The average cranial measurements of the P. javanicus include greatest skull length 13.6 mm (13.0 – 14.6 mm), condylocanine length 12.4 mm (11.9 – 13.1 mm), zygomatic breadth 8.5 mm (8.2 – 9.0 mm), breadth of the braincase 6.6 mm (6.3 – 7.1 mm), postorbital constriction 3.7 mm (3.3 – 4.3 mm) and mandibular length 9.9 mm (9.3 – 10.7 mm)

(Bates and Harrison, 1997).

Pipistrellus pipistrellus. The taxonomic status of this bat species is unknown from Pakistan. The

species is distributed in British Isles, S Denmark, W Europe to the Volga and Caucasus,

Morocco; Greece, Turkey, Israel and Lebanon to Afghanistan, Kashmir, Kazakhstan, Pakistan,

Burma, Sinkiang (China), perhaps Korea, Japan and Taiwan. The Brisitsh Museum has one

specimen that was collected from Kashmir in the beginning of 19th century. Two other specimens were collected from Gilgit by an expedition carried out by University of Marryland in 1965

(Robers, 1997). The species has a restricted range in the Indian subcontinent (Bates and

Harrison, 1997) and seems to be common in Pakistan as there has been no further field studies on bats in Kashmir or Gilgit (Roberts, 1997). The species is considered to be ‘Least Concern’

(IUCN, 2008). P. pipistrellus is a small sized bat with the tail length smaller than head and body length. The extreme tip of the tail protrudes from interfemoral membrane. The interfemoral membrane is supported by well developed calcars and a post-calcarial lobe is present. The legs are short and feet are small. The pararhinal glandular swellings on the muzzle are well developed. The wings are narrow. The fifth digit is considerably longer than the metacarpal bone of the fourth digit. The ears are short and broad, slightly tapered and rounded at the tip. The tragus is almost ‘banana shaped. Pipistrelles in Europe have recently been shown to comprise two cryptic species – P. pipistrellus which echolocates with most energy around 45 kHz, and P. pygmaeus, with most energy at 55 kHz (Jones and Parijs, 1993).

Body measurements. The body measurements of two P. pipistrellus specimens captured from

Gilgit include head and body length ranging from 42- 45 mm, forearm length 27 – 33 mm with, tail length 33 - 34 mm, hind foot length 7 mm and ear length 11 - 12 mm (Roberts, 1997).

The body measurements recorded by Bates and Harrison (1997) are as follows. Average head and body length is 44.0 mm (40.0 – 48.0 mm), tail length 32.9 mm (29.0 – 35.0 mm), hind foot

length 6.1 mm (6.0 – 7.0 mm), forearm length 31.0 (30.0 – 31.6 mm), length of 5th metacarpal

28.9 (28.4 – 29.8 mm), length of 4th metacarpal 29.6 (28.7 – 30.8 mm), length of 3rd metacarpal

29.9 (29.5 – 31.0 mm) and ear length 11.1 mm (10.5 – 12.0).

Cranial measurements. The Greatest skull length of P. pipistrellus was 12.1 mm (11.9 – 12.5

mm), Condylocanine length measured 10.9 mm (10.4 – 11.3 mm), Zygomatic breadth was 7.6

mm (7.2 – 7.9 mm), Breadth of the braincase was 6.1 mm (5.9 – 6.3 mm), postorbital constriction 3.3 mm (3.2 – 3.5 mm) and the mandibular length 8.4 mm was (7.9 – 8.7 mm)

(Bates and Harrison, 1997).

Pipistrellus tenuis is the smallest pipistrelle found within the subcontinent with an average

forearm length of 27.7 mm. The species is hard to differentiate from smaller individuals of P.

coromandra on the basis of forearm length. Its body mass averages about 2 grams

(Gopalakrishna and Karim, 1972). The species is distributed in Afghanistan to the Moluccas; S

China, Laos, Vietnam, Cocos eeling Isles and Christmas Isle (Indian Ocean). The species has

been recorded from Malakand (Roberts, 1997), Chitral (Sinha, 1980), Multan and Chaklala

(Hinton and Thomas, 1926), Chakri, Gambat, Sukkur (Siddiqui, 1961), Karachi, Malir (Walton,

1974). The species is considered as ‘Least Concern’.

Body measurements. The body measurements of the specimen from India, Pakistan, Nepal and

Sri Lanka are as head and body length is 39.1 mm (33.0 – 45.0 mm), tail length 28.9 mm (20.0 –

35.0 mm), hind foot length 5.3 mm (3.0 – 7.0 mm), forearm length 27.7 (25.0 – 30.2 mm), length

of 5th metacarpal 25.9 (23.5 – 28.5 mm), length of 4th metacarpal 26.4 (23.7 – 29.2 mm), length

of 3rd metacarpal 26.7 (23.9 – 29.7 mm), ear length 9.7 mm (5.0 – 11.0) (Bates and Harrison,

1997) .

Cranial measurements. Greatest skull length 11.5 mm (10.7 – 12.1 mm), Condylocanine length

10.2 mm (9.3 – 10.7 mm), Zygomatic breadth 7.4 mm (7.3 – 7.6 mm), Breadth of the braincase

6.0 mm (5.6 – 6.3 mm), postorbital constriction 3.3 mm (2.9 – 3.7 mm), mandible length 7.9 mm

(7.2 – 8.3 mm) (Bates and Harrison, 1997).

Hypsugo savii The distribution and status of this species is unknown in Pakistan and based on literature survey it can be stated that it may be present in Pakistan (Mahmood-ul-Hassan et al.,

2009). The distribution and population status of this species is unknown in the Indian subcontinent (Bates and Harrison, 1997). This bat has not been shown by Simmons (2005) to be present in Pakistan, however Roberts (1997) and Walker and Molur (2003) have shown this species to be present in East Punjab, India where it is Vulnerable. It is a medium sized pipistrelle with the tail length which is significantly shorter than the head and body. The muzzle is blackish and nearly naked. The ears are relatively large and with broadly rounded tips; each ear having a broad tragus. The interfemoral membrane is essentially naked except for a few hairs adjacent to the tail and body. This species is distributed in France, Portugal, Spain, Italy, S Switzerland,

Austria, E Hungary, Balkan Countries, Morrocco, N Algeria, and the Canary Isles. (Spain) and

Cape Verde Isls through the Crimia and Caucasus, Turkey, Lebanon, Syria, Israel, Iran,

Kazakhistan, Turkemenistan, Uzebekistan, Kyrgyzstan, Tajikistan, Afghistan to N India and

Burma (Wilson and Reeder, 2005; Mahmood-ul-Hassan et al., 2009). IUCN 2008 categorized the species as ‘Least Concern’.

Body measurements. The body measurements of an extralimital specimen are as follows. Head and body length is 52.0 mm (47.0 – 60.0 mm), tail length 33.0 mm (30.0 – 35.0 mm), hind foot length 7.1 mm (6.4 – 8.0 mm), forearm length 31.0 (32.1 – 38.0 mm), wingspan 235.5mm (226.0

– 251.0 mm), length of 5th metacarpal 30.5 (29.1 – 31.3 mm), length of 4th metacarpal 31.5

(30.2 – 34.0 mm), length of 3rd metacarpal 31.9 (30.4 – 33.2 mm), ear length 12.1 mm (10.0 –

14.0) (Bates and Harrison, 1997).

Cranial measurements. The greatest skull length 14.0 mm (13.6 – 14.4 mm), Condylocanine length 12.8 mm (12.4 – 13.3 mm), Zygomatic breadth 8.7 mm (8.5 – 9.1 mm), Breadth of the braincase 6.7 mm (6.6 – 6.8 mm), postorbital constriction 3.6 mm (3.5 – 3.7 mm), mandible length 9.8 mm (9.6 – 10.3 mm) (Bates and Harrison, 1997).

CHAPTER III

MATERIALS AND METHODS

This study was conducted in those arid subtropical and tropical regions of the country which included less pronounced monsoon influenced areas of the Salt Range, the Upper Indus Plains and the sand dune areas typified by the Cholistan (Roberts, 1997). The present study is the first organized attempt to describe the poorly known bat fauna of these regions across a latitudinal gradient (Figure 1). Bat surveys were conducted in two protected areas situated in the arid subtropical region i.e. the Margallah Hills National Park (MHNP) and the Chinji National Park

(CNP), and in another, the Lal Suhanara National Park (LSNP), situated in the tropical sand dune region of the Upper Indus Plains. Some areas lying adjacent to these national parks were also sampled for bats. The MHNP, the CNP and the LSNP were designated as sub-area 1 (SA1), sub- area 2 (SA2) and sub-area 3 (SA3), respectively. In addition to these protected areas, bat samples were also collected from some non-protected areas of Gujranwala, Lahore, Tob Tek Singh and

Kasur districts. These four districts collectively constituted sub-area 4 (SA4). These sub-areas were selected to maximize the chances of capture of as many bat species inhabiting arid- subtropical and tropical habitats of Pakistan as possible. Some detail of each of these four SAs is given in the following section.

STUDY AREA

SA 1 - The Margalla Hills National Park (MHNP)

The Margalla Hills National Park (073°73.32'E, 33°4159.61'N) is 40 km in length and forms the northeastern edge of Islamabad. It spreads in a roughly east–west direction and its altitude varies from 465 to 1600 m (Shinwari and Khan, 2000). The park was a wildlife sanctuary under the

West Pakistan Wildlife Protection Ordinance, 1959 and was declared as national park on April

Figure 1. Map of Pakistan showing the location of 30 sampling sites where any bat roost was located or mist nets were erected in the four sub-areas. Of these seven were located in SA1, seven in SA2, five in SA3 and eleven in SA4.

27, 1980 under Section 21(1) of the Islamabad Wildlife (Protection, Conservation and

Management) ordinance, 1979 and is placed in the World Conservation Union (IUCN)

Management Category V (Protected Landscape). The topography of the MHNP is rugged, with

numerous valleys and steep slopes. The soil type is alkaline (pH 7.3-8.73) except at Pharrheela

Rest House which is slightly acidic (pH 6.82). Electric conductivity of the soil ranges from 0.86

dsm to 3.43 dsm at Saidpur and Rawal Lake areas respectively (Awan et al., 1992). Rocks dating

back to the Jurassic and Triassic ages have been recorded (Hagler Bailly Pakistan, 2007). The

Margalla hills are an extension of the Himalayan range and form the northern boundary of the

Potohar plateau. The area is drained by the River Kurang and its tributaries, which flow into the

Soan River (http://www.wildlifeofpakistan.com/national_parks.html).

The MHNP has been included in the sub-tropical scrub forest (Champion et al., 1965).

The average maximum temperature is 34.3°C while the average minimum temperature is 3.4°C

(range 15 ºC - 40 ºC). Rain fall occurs in the monsoon and winter, the average being 1200 mm

per year. The park is under pressure because of illegal settlements, quarries, fires, extensive tree

cuttings, urban encroachment, and pollution (Shinwari and Khan, 2000). Vegetation of MHNP is

largely supported by the monsoon rains while its flora, is the remnant of the natural communities

from the great Indo-Himalayan ecosystem extending north-east into the greater Indian sub-

continent and Southeast Asia. The Margalla Hills represent a contact zone with the arid Irano-

Saharan ecosystem. It is here that the natural ranges of several plants from both regions overlap.

The southern aspects of these hills have thin soils and marginal rainfall supporting a short stature

and xeric broad-leaved deciduous and evergreen forests. Along with that grows a diverse shrub

under story. Tree stature diversity however increases in ravines. Pines are dominant in the

northern aspects and in elevations higher than 1400 m, where it is cooler and precipitation is

more effective. Groves of oaks are still found in some places. The dominant trees of MNHP, in

alphabetical order include Acacia catechu, A. camus, A. nilotica, A. modesta, Bauhinia

variegate, Butea monosperma, Maytenus rayleanus, Olea ferruginea, Phyllanthus emblica, Pinus

roxburgii and Quercus leucotrichophora (Hijazi, 1984; Akbar, 1988; Khattak and Ahmed, 1990;

Khan, 1994; Shinwari et al., 1996). The Park was setup to provide refuge to the gray goral,

barking deer and the leopard (Panthera pardus). Rhesus monkeys (Macaca mulatta), jackals

(Canis aureus), wild boars (Sus scrofa), porcupines (Hystrix indica), mongoose (Herpestes

fuscus) and the pangolin (Manis crassicaudata) or scaly anteater (Smutsia temminckii) exists in

the area (Masud, 1979). Various sampling stations at Margalla Hills National Park and its

adjacent areas with GPS location of each are mentioned in Table 1 and graphically represented in

Figure 2. These sampling stations include Pakistan Museum of Natural History (PMNH),

Marghzar Zoo, Rawal Town, National Agricultural Research Council (NARC), Rattowal, Tanaza

Dam and Loi Bher Wildlife Park.

SA 2 - The Chinji National Park (CNP)

The CNP is a part of the Salt range which is located in the central-north region of the province of

Punjab in Pakistan. It is situated mainly in the districts of Chakwal and Khushab with the larger

part in the former. In the east-west direction these low hills extend for over 130 kilometres and about 50 kilometres in the north-south direction. The “Khabeki” and the “Uchhali” are the two major salt water lakes of this park that lie in the south of Salt Range. The “Khabeki” is one kilometer wide and two km long while the “Uchhali” that lies at the base of Sakaser (altitude

1522m) covers an area of 943 hectares (Birdlife International, 2011). After Murree, the Soon

Valley is an important hill station of the Punjab (http://www.forumforfree.com/forums/index.php?m

forum=forumpk&showtopic=3641). The sampling stations at CNP with their GPS coordinates are

given in Table 2 and graphically represented in Figure 3.

Table 1. GPS location of seven sites sampled for bats at Margalla Hills National Park and adjacent areas (SA1) from June 2009 to May 2011.

Netting stations GPS Location Latitude Longitude Pakistan Museum of Natural History 33º43.194 073º03.631 Marghzar Zoo 33º44.037 073º03.562 Rawal Town 33º40.966 073º07.108 National Agricultural Research Council 33º39.892 073º07.108 Rattowal 33º28.644 072º43.638 Tanaza Dam, Kherimoorat 33º27.594 072º44.066 Loi Bher Wildlife Park 33º34.632 073 º 07.53

Figure 2. GPS location of five sites sampled for bats at Margalla Hills National Park (n = 7) and adjacent areas ( n =2) which jointly form SA1during the period extending from June 2009 to May 2011 (the map of Pakistan showing the location of Margalla Hills National Park is given in the inset).

Table 2. GPS location of seven sites sampled for bats at Chinji National Park and adjacent areas (SA2) during a period extending from June 2009 to May 2011.

Netting stations GPS Location Latitude Longitude Kanhatti Garden 32º40.711 072º14.818 Sodhi Wildlife Sanctuary 32º34.098 072º13.428 Sodhi Rest House 32º34.550 072º16.425 Shrine of Baba Mehdi 32º35.427 072º20.724 Khabbeki Lake 32º37.185 072º12.654 Dalwal 32º42.725 072º53.065 Uchaali Lake 32º30.329 072º00.305

Figure 3. GPS location of seven sites sampled for bats at Chinji National Park (n = 6) and adjacent areas ( n =1) which jointly form SA2 during the period extending from June 2009 to May 2011 (the map of Pakistan showing the location of Chinji National Park is given in the inset).

The climate is one of the extremes in both winters and summers. During winters, the temperature

goes below the freezing point while in summers the mean maximum daily temperature is above

40ºC. July and August are the rainiest months (monsoon showers). Some rainfall is also received

in January and February but rains are erratic in general often coming in a few storms. The

average annual rainfall decreases from east to west, and consequently the vegetation is more

luxuriant in eastern part than in the west (Said, 1956). The main tree and shrub (non-woody)

species are Olea ferruginea, Acacia modesta, Pistacia integerrima, Dodonaea viscosa, Capparis

aphylla, Tecoma undulata, Gymnosporia royleana, Monotheca buxifolia and Zizyphus

nummularia (Sheikh, 1987, 1993; Said, 1956). On hotter southern aspects Acacia modesta is the

principal tree species where Olea ferruginea may be found scattered and in sheltered places

(Said, 1956; Sheikh, 1987). Both the main tree species i.e. Olea ferruginea and Acacia modesta

are slow growing. Olea ferruginea attains a height of about 3 meters in 30 years and Acacia

modesta reaches about 3.5 meters height at this age (Said, 1956). Under good site conditions a fully mature Acacia modesta tree can grow up to 9 m in height and a mature Olea ferruginea tree up to 9-12m tall (Sheikh, 1993). Various sampling stations at CNP include Kanhatti Garden,

Sodhi Wildlife Sanctuary, Sodhi Rest House, Shrine of Baba Mehdi, Khabbeki Lake, Dalwal village and Uchhali Lake.

SA 3 - The Lal Suhanra National Park (LSNP)

Lal Suhanra (29°12' to 29°28'N; 71°48' to 72°08'E) was established in 1972. It is the oldest national park of the country (Ahmad, 2007) that spans over 153,000 acres and is notable for the diversity of its landscape. Although fragments of all the mjor habitat types exist in this park desert flora and fauna are the key components of this park. The park occupies an area of

127,480 acres out of which 16.5% is covered by irrigated plantations, 79.8% by desert, and 3.7%

by wetlands. Abbasia Canal also traverses across the park that has created a semi-wetland type habitat in the amidist of the desert. Sand dunes measuring between 1 and 6 meters in height are

the predominant landscape feature of this park (Hussain, 2008).

On the basis of parent material, soil and vegetation, the Lal Suhanra National Park is

divided into two regions. The Lesser Cholistan lies in the north and borders canal-irrigated areas

covering about 7770 km2 while the Greater Cholistan region comprises 18130 km2 and

constitutes the southern flank of the park. The Lesser Cholistan consists of ‘Dahars’ which are

large saline, hard and compact areas that alternate with low sandy ridges. Sand dunes are

stabilized, semi-stabilized or shifting, while the valleys are mostly covered with sand. The soils

are classified as either saline or saline sodic, with pH ranging from 8.2 to 8.4 and 8.8 to 9.6,

respectively. The Greater Cholistan is a wind sorted sandy desert and comprises river terraces,

large sand ridges and less interdunal plain areas (Baig et al., 1980; Akbar et al., 1996; Arshad et

al., 2003).

Cholistan desert is characterized by xerophytic species that are adapted to extreme

weather conditions. Eastern part of this desert that receives 200 mm rainfall (annual average)

presents comparatively better vegetation cover than the southern part that receives only 100 mm

rainfall (annual average). Plant distribution in both these areas is mainly affected by soil topography and chemical composition and is manifested by the association of certain plant

species to certain soils. Haloxylon recurvum, H. salicornicum and Suaeda fruticosa dominate

compact saline ‘dahars’, whereas Salsola baryosma, Sporobolus ioclados, Aeluropus lagopoides,

Capparis decidua, Cymbopogon jwarancusa, Ochthochloa compressa and Prosopis cineraria are the indicator species of ‘dahars’ having some sandy cover. Similarly, Calligonum polygonoides, Aerva javanica, Panicum turgidum and Lasiurus scindicus are most common at

sand dunes (Chaudhry, 1992; Arshad et al., 2003). Various sampling sites at Lal Suhanara

National Park include Mojgarh, Derawar Fort, Behawalpur Fish Hatchery, Noor Mahal and

Marot Fort. The GPS coordinates of all these sites are mentioned in Table 3 and represented

graphically in Figure 4.

SA4 - The Central Punjab (CP)

Four districts (Lahore, Kasur, Gujranwala and Toba Tek Singh) in central Punjab were

sampled for collection of bat samples. Various sampling sites at all the four districts are

mentioned in Table 4 and graphically represented in Figure 5.

Lahore (31o20 and 31o50 N and 74o05 and 74o37 E) is the second most populated city of

Pakistan and is situated on the left bank of river Ravi. The population pressure caused the

expansion of the city to the lower alluvial plain and unavailability of ground water is one of the

most adverse consequences. Undulating and low alluvial plains are the characteristics of the area

(Naeem et al., 2007).

Eastern side of the city is surrounded by International Indian border, western side is surrounded by Degh nala, Muridke town lies on the northern side while the Hudiara drain is situated on the southern side of the city (Sergey, 2001). The soil type is late Pleistocene silty loess (Harland, 1970). The development of the canal irrigation system gave rise to extensive agricultural areas and gardens and caused expansion of the city at the expense of Sub Tropical

Thorn Forest ecozone (Champion et al., 1965; Chaudhry, 1990). Indigenous trees were grown

along the roadsides and canals. Mostly, these trees were Mulberry, Pipal, Banyan, Jaman, etc.

(Masood, 2004; WWF-P, 2008).

Table 3. GPS location of five sites sampled for bats at Lal Suhanra National Park and adjacent areas (SA3) from June 2009 to May 2011.

Netting stations GPS Location Latitude Longitude Mojgarh 29º01.132 072º08.427 Derawar Fort 28º46.045 071º20.210 Fish Hatchery, Bahawalpur 29º23.186 071º38.148 Noor Mahal 29º22.695 071º40.132 Marot Fort 29º10.615 072º26.075

Figure 4. GPS location of five sites sampled for bats at Lal Suhanra National Park and adjacent areas which jointly form SA3 during the period extending from June 2009 to May 2011 (the map of Pakistan showing the location of Lal Suhanra National Park is given in the inset).

Table 4. GPS location of eleven sites sampled for bats in four districts of central Punjab that jointly form SA4 from June 2009 to May 2011.

Exact locality GPS Location Latitude Longitude Kalian Daas (T.T. Singh) 31º12.037 072º40.487 Government College, Gojra (T.T. Singh) 31º09.024 072º41.077 Gojra Grave Yard (T.T. Singh) 31º09.263 072º40.058 Badian (Lahore) 31°29.223 074°24.632 Manawa (Lahore) 31º35.647 074º27.660 Fish Ponds, Pattoki (Kasur) 31º02.487 073º52.536 Shalimar Garden, Lahore (Lahore) 31º35.333 074º22.863 Head Balloki (Kasur) 31º12.790 073º51.901 Chhanga Manga (Kasur) 31º04.879 074º00.128 Rasul Nagar (Gujranwala) 32º19.424 073º46.623 Ali Pur Chathha (Gujranwala) 32º11.272 074º09.361

Figure 5. GPS location of eleven sites sampled for bats at Gujranwala (n = 2), Lahore (n = 3), Kasur (n =3) and Toba Tek Singh ( n = 3) which jointly form SA1during the period extending from June 2009 to May 2011 (the map of Pakistan showing the location of Margalla Hills National Park is given in the inset).

Kasur is situated on southeast side of Lahore and is one of the oldest industrial cities of Pakistan

(http://en.wikipedia.org/wiki/Kasur). Although there are significant changes in lithologies having

soil contents of silt, loamy clay, clay and sandy, the clay particles increase with distance from

Ravi river. The average annual rainfall is about 650 mm, but rains are more frequect during

monsoon (Ashraf et al., 2010).

Gujranwala (32°.16 N, 74°.18 E) lies 226 m above sea level and is the seventh largest

city in Pakistan (World Gazetteer, 2010). Hot and moist climate prevails during summer and

colded during winter season. The city receives average annual rainfall of 950 mm but the rainfall

is more frequent during monsoon. The temperature varies from 4°C during winter to above 40°C

during summer (Qadir et al., 2008; Ullah et al., 2009).

District Toba Tek Singh (30°33 to 31°2 degree N and 72°08 to 72°48′ E) is district

comprises of three tehsils i.e. Gojra, Kamalia, and Toba Tek Singh (T.T. Singh). River Ravi

floods the lower parts of the district during monsoon. The temperature of the district varies between 6 °C during winter to 40.7°C during summer. Average annual rainfall is 254–381 mm

(Tauseef-ur-Rehman et al., 2011).

The diversity of the bat fauna of these four SAs was ascertained by identifying various bat specimens up to species level. Although molecular techniques are being widely used now a

days, simple morphological techniques that form the basis of most authentic taxonomic studies

on bats of this region and elsewhere in the world were employed (see for example Dobson, 1876,

1877, DeBlase, 1980, Bates and Harrison, 1997, Roberts, 1997, Dietz and Helversion, 2004,

Srinivasulu et al., 2010 etc.) due to budgetary constraints.

Bats were identified up to species on the basis of their external body and cranial and

bacular measurements, and compared with the available literature (Bates and Harrison, 1997,

Roberts, 1997, Mahmood-ul-Hassan et al., 2009 and Sirinivasulu et al., 2010 are the main source

of information on bats of the study area were consulted for this purpose). The bat echolocation

calls were also recorded to develop sonograms of various bat species of the study area. The echolocation calls of bats of Pakistan have never been recorded and thus the accurate

identification of bats species from their calls might be wrong. It is only due to the dearth of

literature on this aspect of Pakistani bat sound calls, fiand this pioneering study will aid future

bat biologists to identify various bat species on the basis of their calls and help in population assessment and conservation studies. The population of bats of the each sampling site was also estimated. The details of each of these methods are described as follows.

SAMPLING STRATEGY

Exploratory visits were made to locate as many bat roosts in all the sub-areas as possible.

Potential bat roosts such as old and undisturbed buildings, ruins, abandoned wells, farm houses, tree groves and forest plantations were thoroughly searched. Local people were also interviewed for gleaning maximum information about the exact location of various bat roosts. An ultrasonic detector (Pettersson Ultrasound Detector D 1000X) was used to maximize the chances of locating a bat roost. Once located, the global position of each roost was determined using a

Garmin etrax H Global Position System (GPS).

A total of six viz., 12 m (n = 1), 9 m (n =2) and 6 m (n = 3) long high quality, deep black,

UV stable and strong mist nets (Ecotone 716/6, 716/9 and 716/12) were used to capture bats.

Each of these five shelved, 16×16 mm mesh sized and 2.5 m high net was erected either in “L” or “V” shape at strategic positions on a pair of 3 m long bamboo poles in such a way that the last shelf of each net remained one foot above the ground. The total mist net area in each session was

120 m2. The nets were ready to operate half an hour before sunset. All the nets were opened

42 simultaneously at sunset and continued to operate, depending on the weather conditions, for two hours after sunset. Nets were checked continuously to disentangle any captured bat. The sampling effort remained the same throughout the study.

The population of the bats at a particular subarea was assessed by calculating the netting index and relative abudance of various species. The neting index and relative abundance were calculated by using following formulae.

Netting index of ith species = (ni/Total net area  hrs) 100

Overall netting index = (N/Total net area  hrs) 100

Where

ni = Number of individuals of ith species, and

N = Total number of individuals of all the bat species captured

While

Relative abundance (%) of ith species = (ni/N) 100

The spatial and temporal changes in diversity of bat fauna were also investigated.

Combined relative abundances of the bats captured in spring (March, April and May), summer

(June, July and August), autumn (September, October and November) and winter (December,

January and February) were used to find out temporal changes in bat diversity. PAST statistical package was used to calculate various diversity indices.

EXTERNAL MORPHOLOGY

Each disentangled bat was placed in a separate cotton bat bag during mist netting and at the completion of a netting session, each bat was weighed up to 0.1 g (Pesola balance 10050,

Swiss made) euthanized and preserved in a plastic bottle in absolute alcohol. Field number, sex, age, exact locality and district of capture of each bat was noted on the plastic bottle. After three

43 consecutive nights of mist netting in a subarea, all the specimens were brought to the laboratory and autopsied after recording their external body measurements using a digital vernier caliper (0-

150 mm). These measurements include head and body length, ear length, tragus length, forearm length, claw length, thumb length, length of each metacarpal including its phalanges, wing span, penis length, tibia length, calcar length, hind foot, tail length, and free tail length (in case the tail is completely or partially free of uropatagium)., These measurements were used to ascertain the species of the bat captured from a particular subarea and compared with those already available in the literature. The age of each captured bat specimen was determined following Dietz (2005).

CRANIAL MEASUREMENTS

Skulls were prepared for recording cranial measurements of each bat specimen. For this purpose, eye balls, tongue and excessive flesh was removed from each skull. The brain tissue was macerated and removed using forceps and cotton and cranial cavity was washed with a jet of water. Skulls thus cleaned were kept overnight in a dilute solution (0.2 % of Potassium

Hydroxide (KOH)). After being thoroughly washed in tap water again, the skulls were kept in absolute alcohol for a night before being transferred to acetone for another night. Each of the dry skulls was stored in a properly labeled vial padded with cotton. The greatest length of skull

(GTL), condylo-basal length (CBL), condylo-canine length (CCL), zygomatic breadth (ZB), breadth of braincase (BB), post-orbital constriction (PC), mandible length (M), maxillary toothrow length (C-M3), mandible toothrow length (C-M3), posterior palatal width (M3-M3), anterior palatal width (C1-C1) were measured (Figure 6 ).

BACULAR MEASUREMENTS

The penis of the male bats was cut down as close to the surface of the body as possible so that the baculum is not damaged. The cut penis was be placed in a test tube half filled with cold water

Figure 6. Dorsal (top), ventral (middle) and lateral (bottom) view of the bat skull showing definition of various cranial measurements (GTL = Greatest skull length, CBL Condylo-basal length, CCL = Condylo-canine length, ZB = Zygomatic breadth, BB = Breadth of the braincase, IC = interorbial constriction, PC = Postorbital contriction, M 3 = Mandible length, C-M = Maxillary toothrow length, C-M3= Mandibular toothrow length, M3-M3 = Posterior palatal width, C1-C1 = Anterior palatal length after Bates and Harrison, 1997)

45 and boiled for two minutes. The boiled penis was then transferred to a plastic tube containing 5%

KOH and a pinch of alizarin red powder. After 24 hours, the stained bacula were dissected out of the tissue under a dissecting microscope with fine foreceps and stored in glycerin in a labeled test tube (Bates et al., 2005). The baculum comprises of a bony (proximal) and a cartilaginous

(distal) segment (Lidicker, 1968). Once a stained baclum is obtained, various bacular measurements and ratios were recorded following Lidicker and Yang (1986) under a microscope at 30X magnification using a stage and an ocular micrometer. Only those bacula were used in this study which are complete and their cartilaginous portions are intact. Following measurements ratios and indices were taken from each baculum (Figure 7).

Simpson (1996) was followed for statistical analysis of data on bacular measurements.

Total bacular length (TLB)

Shaft length (SL)

Proximal branch length (PBL)

Distal branch length (DBL)

Proximal branch width (PBW)

Distal branch width (DBW)

Bacular height (BH)

ECHOLOCATION

Recordings conditions and equipment

Recordings were made under three conditions;

(i). During emergence from roosts where bats of known identity occurred. Each site was visited only once to avoid pseudo-replication (Hulbert, 1984). Calls were recorded 20-30 m away from

Figure 7. Dorsal (above) and lateral (below) view of the baculum of bat defining parameters recorded for measuring bacula of the bats (TLB = Total bacular length, SL = Shaft length, PBL = Proximal branch length, DBL = Distal branch length, PBW = Proximal branch width, DBW = Distral branch width, BH = Bacular height).

47 the roost exit, so the usually broadband calls emitted immediately after emergence not included in our dataset.

(ii). When bats released from the hand after capture. The bats were mist-netted at foraging sites while leaving the roost, on a few occasions, captured inside the roost. Some bats were released in clutter, others in the open. As a result, both calls affected by cluttered environments and those typically emitted in the open were represented in the sample.

(iii). At foraging sites. In this regard the bats were recorded in free-flight. Each individual was recorded at a different site to eliminate the risk of pseudo-replication. The species were recorded in flight and identified by examining the species-specific structure of their social calls (Barlow and Jones, 1997 a,b; Russo and Jones, 1999, 2000).

Sound Analysis

None of the bats found in Pakistan has ever been recorded. A Patterson D1000X was used to record bat sounds. The recordings were analyzed with the software Bat-Sound 4

(Pettersson Elektronik AB) using a sample frequency of 44.1 kHz, with 16 bits/sample, and a

512 point FFT with a Hamming window for analysis. At least two echolocation calls (pulses) selected at random from each bat was analyzed for all species.

Five parameters were measured from each call; start frequency (SF, the frequency value measured at the beginning of the call), end frequency (EF, the frequency value measured at the end of the call), frequency of maximum energy (FMAXE), duration (D) and inter-pulse interval

(IPI, the time interval between 2 consecutive calls). D and IPI (ms) were measured from oscillograms, FMAXE (kHz) from power spectra, and all other spectral parameters (kHz) from spectrograms. In some bat species, the highest energy may be in either the fundamental or in the second harmonic (e.g. Parsons and Jones, 2000); therefore FMAXE (kHz) was taken from the

harmonic with the highest energy while other measurements were taken from the fundamental.

For most of the species, measurements were taken from the fundamental harmonic containing most energy, i.e. always from the fundamental in most of the species. Mean and standard

deviation for all the pulses analyzed from a single long recording was determined and the values

were compared with those available in literature. In the absence of a bat call library, only a small

proportion of the recorded calls assigned to any particular bat species.

CHAPTER IV

RESULTS

PART I. BAT SURVEY AND ABUNDANCE

An extensive survey of the study area was conducted from June 2009 to May 2011 during which a total of 26 sampling efforts were made (Table 5, 6) and 65 days were spent in the field

(Table 6). The number of days in each sampling effort varied from one to three. These sampling efforts involved both searching bat roosts and strategic sites (sampling station) for erecting mist nets during the day and erecting mist nets for capturing bats in the evening. Bat surveys were labor intensive and logistically hungry and time consuming: suitable sampling stations were searched from dawn until dusk, and once a site was finalized mist nets were erected at that particular sampling station prior to sunset. The main criteria for selecting any a sampling station involved three factors viz., (a) it was secure for all the field workers to work until late at night,

(2) there was a bank of some water body or edge of a plantation nearby, and (c) it was close to the camping station. Mist netting was mostly done at the banks of water bodies that were situated within the premises of some governmental organizations where security staff was available all the night long. In spite of security concerns, attempts were made to sample all habitat types present in the four sub-areas. The survey intensity ranged from 3 days in July and November to 8 days in April while on the average 2.67 days were spent in each month in collecting bat samples

(Table 6).

The survey was conducted in following four Sub-Areas (SAs).

1. Margalla Hills National Park and Adjacent Areas (SA1)

2. Chinji National Park and Adjacent Areas (SA2)

3. Lal Suhanra National Park and Adjacent Areas (SA3)

Table 5. Time frame and survey intensity in four sub-areas sampled for bats from June 2009 to May 2011.

Year Sub-area SA1 SA2 SA3 SA4 18-20 Jun - 17-19 Aug - 2009 28-30 Sep - 20-22 Nov - 17-19 Dec 22-24 Oct - - 26-27 Mar 08-10 Jan 26 -28 Feb 19-21 Mar 17-19 Jul 23-25 Apr 17-19 May 16-17 Apr 2010 - 20-22 Aug 10-12 Sep 17-18 Jun - - - 8 - 9 Oct - - - 8-9,31 Dec 25 Jan 23 Jan - 21-23 Jan - 24-25 Apr - 26 -27 Feb 2011 - - - 18 Apr - - - 15-16 May

Table 6. Number of days spent in the field in each month in the four sub-areas and the total number of days spent in the field from June 2009 to May 2011. SA1 SA2 SA3 SA4 Combined 2009 2010 2011 2009 2010 2011 2009 2010 2011 2009 2010 2011 2009 2010 2011 Total Jan. - 1 3 1 - - - 3 3 4 7 Feb. - - - - 3 - - 2 3 2 5 Mar. - 2 ------3 - 5 - 5 Apr. - - 3 2 - - 2 1 5 3 8

May - - - - 3 - - 2 3 2 5 Jun. 3 ------2 3 2 5 Jul. - 3 ------3 3 Aug. - - - 3 3 - - - 3 3 6

Sept. 3 - - - - 3 - - - 3 3 - 6 - - Oct. - - 3 - - - - 2 3 2 5

Nov. - - - - 3 - - - 3 - 3 Dec. 3 ------3 3 3 6 9 5 1 3 9 3 6 9 12 8 18 35 12 65 15 15 15 20 65

1. Four districts of the central Punjab (SA4) which included Hafizabad, Lahore, Kasur and

Toba Tek Singh districts. Bat captures and trap indices for each species at each sampling

station in the four SAs is described as follows.

The bat captures from each of these areas is described as follows.

SUB-AREA 1

A total of 3600 m2hrs of mist netting effort resulted in the capture of 72 bats representing two families, four genera and nine species (Table 7). The overall netting index was 1.64. The netting index for two Taphozous spp. was not calculated as these Emballonurids were captured from their roost with the help of hand nets. Pipistrellus pipistrellus (netting index = 0.53; n =19; 11♂,

8♀) was the most captured while Hypsugo savii and Scotophilus kuhlii (netting index = 0.03) were the least captured among nine bat species. Scotophilus heathii (n = 18; 9♂, 9♀) and P. ceylonicus (n =15; 6♂, 9♀) were the two other captured species. Netting index for these two species was 0.50 and 0.42, respectively. Eleven Taphozous nudiventris and two T. perforatus that were hiding in their day time roosts were also captured from this SA (Table 8). The number of bats captured at each of seven sampling stations is described as follows.

1. Pakistan Museum of Natural History (PMNH): Mist nets were erected from 18 to 19

June, 2009 at this station (Table 8) and 20 bats belonging to three genera and five species

were captured. These included, in order of their abundance, Pipistrellus ceylonicus (n =

14), Scotophilus heathii (n = 2) and P. javincus (n = 2), and S. kuhlii (n =1) and Hypsugo

savii (n =1). Of these, there were twelve adult males and eight adult females.

2. Marghzar Zoo. No bat was captured from this station on June 20, 2009.

3. Rawal Town: Three netting efforts were made at Rawal Town Fish hatchery during the

whole study and a total of 21 bats were captured. Of these 21 adult bats, twelve were

Table 7. Species, sex, number, age, netting index and relative abundance of the bats collected from seven sampling stations in SA1from June 2009 to May 2011 (n is the number of bats captured from each station).

Species N Age Netting % Relative ♂ ♀ index abundance Taphozous nudiventris 7 4 Adult - 15.3 T. perforatus 2 0 Adult - 2.8 Scotophilus heathii 9 9 Adult 0.50 25.0 S. kuhlii 1 0 Adult 0.03 1.4 Pipistrellus pipistrellus 11 8 Adult 0.53 26.4 P. ceylonicus 6 9 Adult 0.42 20.8 P. javanicus 1 2 Adult 0.08 4.2 P. tenuis 2 0 Adult 0.06 2.8 Hypsugo savii 1 1 Adult 0.03 1.4

Number of bat species recorded= 9 Number of individuals= 72 Total netting efforts= 3600m2 hrs Overall netting index= 1.64

Table 8. Species, sex and number of bats collected from seven sampling stations in SA1 from June 2009 to May 2011 (n is the number of bats captured from each station).

Month Netting stations n Species (n Sex) Scotophilus heathii (2♂) Scotophilus kuhlii (1♂) Pakistan Museum of Natural History 20 Pipistrellus ceylonicus (6♂, 8♀) Jun.2009 Pipistrellus javanicus (1♂, 1♀) Hypsugo savii (1♂) Marghzar Zoo - - Scotophilus heathii (5♂) Rawal Town 8 Pipistrellus pipistrellus (2♂) Pipistrellus tenius (1♂) Scotophilus heathii (3♀) Sep. 2009 National Agricultural Research 9 Pipistrellus pipistrellus (2♂, 3♀) Council (NARC) Pipistrellus tenius (1♂) Taphozous nudiventris (3♂, 2♀) Rattowal 6 Taphozous perforatus(1♂) NARC - - Dec.2009 Tanaza Dam, Kherimoorat - - Loi Bher Wildlife Park - - Scotophilus heathii (1♂, 1♀) NARC 9 Mar 2010 Pipistrellus pipistrellus (4♂, 3♀) Tanaza Dam, Kherimoorat - - Scotophilus heathii (5♀) Rawal Town 6 Pipistrellus javanics (1♀) Jul. 2010 Tanaza Dam, Kherimoorat - - Taphozous nudiventris (4♂, 2♀) Rattowal 7 Taphozous perforatus (1♂) Scotophilus heathii (1♂) Jan. 2011 Rawal Town 7 Pipistrellus ceylonicus (1♀) Pipistrellus pipistrellus (3♂, 2♀)

male and nine were females. The catch included five S. heathii, two P. pipistrellus and

one P. tenuis during September 2009, five S. heathii and one P. javanicus during July

2010 and five P. pipistrellus, one P. ceylonicus and one S. heathii during 2011.

4. National Agricultural Research Council (NARC): NARC was netted during resulting

in capture of nine bats each was captured from this station during September 2009 and

March 2010 while none was capturedduring December 2009. The captured specimens

belonged to two genera and three species (Table 8).

5. Rattowal. A total of thirteen bat specimens were captured from this station. Of these, six

were captured during September, 2009 and seven during July, 2010. These included

eleven T. nudiventris and two T. perforatus.

6. Tanaza Dam, Kherimoorat. No bat was captured from this station during December

2009, March 2010 and July 2010.

7. Loi Bher Wildlife Park: Mist nets for bats were erected in Loi Bher Wildlife Park

during December 2010 but no bat specimen was captured from this station.

SUB-AREA 2

This sub-area included the Chinji National Park and its adjacent areas. Since civilians were not

permitted to carryout research activities in a large proportion of the area of this National Park and entry rights were reserved solely for the security agencies, only a fragment of this National

Park was actually sampled for bats. A total of 15 bat specimen belonging to three genera and four species were captured (Table 9). A total of 3600 m2hrs of netting efforts were employed at

seven sampling stationsat SA2 (Table 10). Overall mist netting index was 0.42. Scotophilus heathii

was the most captured species (netting index = 0.19; 4♂, 3♀) while P. javanicus was the second in order of its abundance (netting index = 0.17; 3♂, 3♀). Netting index for P. pipistrellus (1♂ = 1) and H. savii

(1♀ = 1) was the same (0.03).

Table 9. Species, sex, number, age, netting index and relative abundance of the bats collected from seven sampling stations in SA2 from June 2009 to May 2011 (n is the number of bats captured from each station).

Species N Age Netting % Relative ♂ ♀ index abundance Scotophilus heathii 3 4 Adult 0.19 46.7 Pipistrellus pipistrellus 1 - Adult 0.03 6.7 P. javanicus 3 3 Adult 0.17 40.0 Hypsugo savii - 1 Adult 0.03 6.7

Number of bat species recorded= 4 Number of individuals= 15 Total netting efforts= 3600 m2 hrs Overall netting index= 0.42

Table 10. Species, sex and number of bats collected from seven sampling stations in SA2 from June 2009 to May 2011 (n is the number of bats captured from each station).

Month Netting stations N Species (nSex) Shrine of Baba - - Mehdi Oct. 2009 Khabbeki Lake - - Dalwal Village - - Uchhali Lake - - Kanhatti Garden - - Jan. 2010 Uchhali Lake - - Khabbeki Lake - - Dalwal Village 13 Scotophilus heathii (3♂, 4♀) Pipistrellus pipistrellus(1♂) Apr. 2010 Pipistrellus javanics (3♂, 2♀) Uchhali Lake 2 Pipistrellus javanicus (1♀) Hypsugo savii (1♀) Khabbeki Lake - - Aug. 2010 Kanhatti Garden - - Uchhali Lake - - Jan. 2011 Kanhatti Garden - - Sodhi Wildlife - - Sanctuary Apr. 2011 Sodhi Wildlife - - Sanctuary Sodhi Rest House - -

Netting stations at SA2 included Kanhatti garden, Sodhi Wildlife Sanctuary, Sodhi Rest House,

Shrine of Baba Mehdi, bank of Khabbeki lake, Dalwal village and bank of Uchhali lake (Table

10). The entire spring catch belonged to Vespertilionidae (Table 10). Percentage relative abundance was maximum for S. heathii (46.67%).

1. Kanhatti Garden: Mist netting at Kanhatti garden was done thrice (January and August

2010, and January 2011) but no bat was captured during those netting efforts.

2. Sodhi Wildlilfe Sanctuary: In spite of through searching at Sodhi Wildlife Sanctuary,

no bat roost was located. Mist nets were erected on April 24, 2009 but no specimen was

captured.

3. Sodhi Rest House: On April 25, 2009 mist netting was done at Sodhi Rest House. No

specimen was captured from this station.

4. Shrine of Baba Mehdi: Mist netting efforts on the bank of a spring near shrine of Baba

Mehdi were also futile.

5. Dalwal Village: This area was sampled twice, once during October 2009 and the then

during April 2010. The first effort remained unproductive while thirteen bats were

captured during the second which included Scotophilus heathii (n = 7), Pipistrellus

pipistrellus (n = 1), and Pipistrellus javanicus (n = 5).

6. Uchhali Lake: This Lake was mist netted thrice during the whole study period i.e. during

January 2010, April, 2010 and August 2010. P. javanicus and a Hypsugo savii were

captured during April 2010 while no bat was captured during the other two visits.

7. Khabbeki Lake. Mist netting at the bank of this lake was done thrice i.e. during October

2009, April 2010 and August 2010 was also useless. But no bat was captured.

SUB-AREA 3

A total of 43 bats belonging to 3 families, 5 genera and 6 species were captured from this sub- area. The total netting effort was 3600 m2hrs with overall mist netting index of 0.47. Rhinopoma hardwickii, Taphozous nudiventris and T. perforatus were captured from their daytime roosts with the help of hand net. Pipistrellus pipistrellus was the most recorded (netting index = 0.33) while Scotoecus pallidus (netting index = 0.06) was the least recorded among those captured through mist nets. Taphozous nudiventris (34.9%) and Pipistrellus pipistrellus (27.9%) were the most abundant bat species of this sub-area (Table 11). Bat captures from each roost are described as follows.

1. Mojgarh. Here a shrine named Shrine of Abdul Maroof Shah provided a roost to almost 300

Taphozous spp. The shrine was visited thrice during this study. On August 17, 2009, nine

bats were captured of which eight (6 ♂ and 2 ♀) were T. nudiventris and one (♀) was T.

perforatus. During the subsequent visits to this roost on November 2009 and May 2010 no

bat was observed as the shrine had been cleaned and decorated after flushing all the bats from

this roost.

2. Derawar Fort. About 500 emballonurids were counted in this ruined fort on August 18,

2009. Of these eight were captured (seven T. nudiventris and one T. perforatus) with the help

of a hand net. Afterwards, the fort was visited four times (i.e. on November, 2009, February,

May and September, 2010) but no bat was recorded.

3. Fish Hatchery Bahawalpur. This locality was sampled four times and a total of 24 bats

were mist netted (Table 12). The netting effort during month of August, 2009 however

remained unproductive.

Table 11. Species, sex, number, age, netting index and relative abundance of the bats collected from five sampling stations in SA3 from June 2009 to May 2011 (n is the number of bats captured from each station).

Species N Age Netting index % Relative ♂ ♀ abundance R. hardwickii 2 0 Adult - 4.7 T. nudiventris 10 5 Adult - 34.9 T. perforatus 2 0 Adult - 4.7 S. pallidus 1 1 Adult 0.06 4.7 S. heathii 10 0 Adult 0.28 23.3 P. pipistrellus 6 6 Adult 0.33 27.9

Number of bat species recorded= 6 Number of individuals= 43 Total netting efforts= 3600 m2 hrs Overall netting index= 0.47

Table 12. Species, sex and number of bats collected from five sampling stations in SA3 from June 2009 to May 2011 (n is the number of bats captured from each station).

Month Netting stations n Species (n Sex) Mojgarh 9 Taphozous nudiventris (6♂,2 ♀) Taphozous perforatus (1♂) Aug. 2009 Derawar Fort 8 Taphozous nudiventris (4♂, 3♀) Taphozous perforatus (1♂) Bahawalpur Fish Hatchery - - Mojgarh - - Nov. 2009 Derawar Fort - - Noor Mahal 2 Rhinopoma hardwickii (2♂) Derawar Fort - - Fish Hatchery, Bahawalpur 14 Scotoecus pallidus (1♂, 1♀) Feb. 2010 Scotophilus heathii (7♂) Pipistrellus pipistrellus (3♂, 2♀) Marot Fort - - Mojgarh - - May 2010 Derawar Fort - - Fish Hatchery, Bahawalpur 3 Pipistrellus pipistrellus (2♂, 1♀) Derawar Fort - - Fish Hatchery, Bahawalpur 7 Scotophilus heathii (3♂) Sep. 2010 Pipistrellus pipistrellus (1♂, 3♀) Marot Fort - -

4. Noor Mahal. Noor Mahal is an archeological site managed by Pakistan Army.Here a colony

of 30 R. hardwickii was observed roosting in cellars. Two male R. hardwickii were collected

from this roost with the help of a hand net. No further visits could be made as the entry rights

were reserved.

5. Marot fort: This fort was visited twice during the whole study and no direct or indirect sign

of the presence of bat was recorded.

SUB-AREA 4

Mist netting was also carried out in four districts of central Punjab which included

Gujranwala, Lahore, Kasur and Toba Tek Singh and a total of 52 bat samples belonging to two families, three genera and six species (Table 13) were recorded from 11 sampling stations.

Rhinolophus blasii (n = 3; netting index = 0.05), Scotophilus heathii (n= 18; netting index =

0.31), S. kuhlii (n= 4; netting index = 0.07), Pipistrellus pipistrellus (n= 20; netting index = 0.35) and P. ceylonicus (n= 7; netting index = 0.12) were recorded from SA4. The percent relative

abundance was highest for P. pipistrellus (32.3%) and the lowest for R. blasii (4.8%).

The station-wise bat data is given in Table 14 and described as follows.

1. Kalian Daas. Mist netting at Kalian Daas, (Toba Tek Singh) was done on March 19, 2010

and seven bats belonging to two species i.e. S. heathii (5♂, 1♀) and P. pipistrellus (1♂) were

captured.

2. Government College Gojra. Mist netting at Government Boys College Gojra was carried

out on March 20, 2010 and six bats, all of them were S. heathii (4♂, 2♀).

3. Gojra Grave Yard. No bat was captured on March 21, 2010 from Gojra Grave Yard. .

Table 13. Species, sex, number, age, netting index and relative abundance of the bats collected from eleven sampling stations in SA4 from June 2009 to May 2011 (n is the number of bats captured from each station).

Species N Age Netting index % Relative ♂ ♀ abundance Rhinolophus blasii 1 2 Adult 0.05 4.8 Scotophilus heathii 14 4 Adult 0.31 29.0 S. kuhlii 0 4 Adult 0.07 6.5 Pipistrellus pipistrellus 14 6 Adult 0.35 32.3 P. ceylonicus 3 4 Adult 0.12 11.3

Number of bat species recorded= 6 Number of individuals= 52 Total netting efforts= 3600m2 hrs Overall netting index= 0.90

Table 14. Species, sex and number of bats collected from eleven sampling stations in SA4 from June 2009 to May 2011 (n is the number of bats captured from each station).

Month Exact locality n Species (nSex) Kalian Daas 7 Scotophilus heathii (5♂, 1♀) Pipistrellus pipistrellus (1♂) Mar. 2010 Government College, 6 Scotophilus heathii (4♂, 2♀) Gojra Gojra Grave Yard - - Rasul Nagar, Gujranwala 12 Scotophilus heathii (2♂, 1♀) Apr. 2010 Pipistrellus pipistrellus (6♂, 3♀) Ali Pur Chathha 1 Pipistrellus pipistrellus (1♂) Badian 4 Scotophilus kuhlii (4♀) Jun. 2010 Manawa 7 Pipistrellus ceylonicus (3♂, 4♀) Badian - - Oct. 2010 Manawa - - Fish Ponds, Pattoki - - Dec. 2010 - - - - Badian - - Jan. 2011 Manawa 3 Rhinolophus blasii (1♂, 2♀) Shalimar Garden, Lahore - - Head Balloki - - Feb. 2011 Chhanga Manga - - Apr. 2011 Shalimar Garden, Lahore - - Fish Ponds, Pattoki 12 Scotophilus heathii (3♂) May 2011 Pipistrellus pipistrellus (6♂, 3♀)

4. Fish Hatchery Manawa. Seven P. ceylonicus (3♂, 4♀) were captured on June 18, 2010

from this station. No bat was mist netted on a subsequent visit during October, 2010 but

three R. blasii (1♂, 2♀) were captured with hand net on January 21, 2011.

5. Shalimar Garden. Mist netting at Shalimar Garden, Lahore was made during the months of

January and April, 2011 but no bat was captured.

6. Pattoki. This station was mist netted on December, 2010 and May, 2011. No bat was

captured during the first session while Scotophilus heathii (n= 3) and P. pipistrellus (n= 9)

were captured on the subsequent session. All the bats were adult. The sexes of these bats are

given in Table 14.

7. Head Balloki. No bat was captured from this station on February and May, 2011.

8. Chhanga Manga. No bat specimen was captured from this station.

9. Rasul Nagar. Three S. heathii (2♂, 1♀) and nine P. pipistrellus (6♂, 3♀) were capture from

this station on April 16, 2010 from this station.

10. Ali Pur Chathha. Mist netting at Ali Pur Chathha was made on April 17, 2010 and a single

male P. pipistrellus was captured.

COMBINED ABUNDANCE AND DIVERSITY

Monthly captures and seasonal abundances

Table 15 describes monthly capture pattern of the twelve bat species recorded from the study area. The combined relative abundance (%) of all bat species recorded from the arid subtropical and tropical regions of Pakistan is given in Table 16. Rhinolophus blasii and S. pallidus were captured only during winter, R. hardwickii and P. tenuis during autumn, and S. kuhlii was recorded only during summer.

Taphozous nudiventris and T. perforatus were captured during summer and autumn. Scotophilus

Table 15. Monthly capture/activity patterns of the twelve bat species captured from some arid subtropical and tropical regions of Pakistan.

Month of capture Species Jun Jul Aug Sep Oct Nov Dec Jan Feb Mar Apr May Rhinolophus blasii ***** Rhinopoma hardwickii ***** Taphozous nudiventris ***** ***** ***** Taphozous perforates ***** ***** ***** Scotoecus pallidus ***** Scotophilus heathii ***** ***** ***** ***** ***** ***** ***** ***** Scotophilus kuhlii ***** Pipistrellus ceylonicus ***** ***** Pipistrellus javanicus ***** ***** ***** Pipistrellus pipistrellus ***** ********** ***** ***** ***** Pipistrellus tenuis ***** Hypsugo savii ***** *****

Table 16. Combined seasonal relative abundance (%) of the twelve bat species captured from the four SAs (n is the number of bats; N is the total numer of bats captured in any season).

Family/Species Summer Autumn Spring Winter % Rel. abundance (n) Rhinolophidae - - - 12.50 (3) Rhinolophus blasii - - - 12.50(3) Rhinopomatidae - 6.25(2) - - Rhinopoma hardwickii - 6.25(2) - - Emballonuridae 39.34(24) 18.75(6) - - Taphozous nudiventris 34.43 (21) 15.63(5) - - Taphozous perforatus 4.92(3) 3.13(1) - - Vespertilionidae 60.66(37) 75.00(24) 100.00(65) 87.50(21) Scotoecus pallidus - - - 8.33(2) Scotophilus heathii 11.48(7) 34.38(11) 41.54(27) 33.33(8) Scotophilus kuhlii 8.20(5) - - - Pipistrellus ceylonicus 34.43(21) - - 4.17(1) Pipistrellus pipistrellus - 34.38(11) 47.69(31) 41.67(10) Pipistrellus javanicus 4.92 (3) - 9.23(6) - Pipistrellus tenuis - 6.25(2) - - Hypsugo savii 1.64(1) - 1.54(1) - N 61 32 65 24

68 heathii was the only species captured during all the four seasons.

Highest species richness was recorded during summer that was followed by autumn, winter and spring. The species dominance was found to be highest during spring followed by winter, autumn and summer. The maximum diversity (H’) was recorded during spring followed by summer, autumn and winter. The evenness (E) was highest during autumn and lowest during summer. The Menhinick index was highest during autumn followed by winter, summer and spring seasons. The Margalef index was highest in summer and lowest in spring season. The equitability, Fisher alpha and Berger-Parker values were highest during winter, autumn and spring respectively (Table 17).

Spatial Variation

Table 18 shows locality related variations in the percent relative abundance of the bat species captured from four sub-areas. The Table shows that R. blasii was captured only from SA4, R. hardwickii and S. pallidus from SA3 and P. tenuis from SA1while T. nudiventris, T. perforatus were recorded from from SA1 and SA3, S. kuhlii and P. ceylonicus from SA1 and SA4, H. savii and P. javanicus from SA1 and SA2. Scotophilus heathii and P. pipistrellus were recorded from all the four sub-areas. S. heathii was the most abundant of all the species.

Species richness was maximum at SA1 followed SA4 and SA3 while it was the lowest at

SA2. The dominance was highest for SA2 and lowest for SA1. Both Shannon and Simpson indices had maximum value at SA1followed by SA3 and SA4 while lowest at SA2. Evenness was highest at SA4 followed by SA3, SA2 and SA1 (Table 19).

Table 17. Combined seasonal diversity of various bat species captured from the four SAs.

Indices Summer Autumn Spring Winter Total Taxa_S 7 6 4 5 12 Individuals 61 32 65 24 182 Dominance_D 0.262 0.2695 0.4088 0.309 0.2059 Shannon_H 1.479 1.33 1.848 1.002 1.479 Simpson_1-D 0.738 0.7305 0.5912 0.691 0.7941 Evenness_e^H/S 0.674 0.7315 0.6811 0.7565 0.5289 Menhinick 0.8963 1.061 0.4961 1.021 0.8895 Margalef 1.46 1.443 0.7187 1.259 2.114 Equitability_J 0.7972 0.8255 0.7229 0.8266 0.7436 Fisher_alpha 2.04 2.18 0.9413 1.922 2.884 Berger-Parker 0.3443 0.3438 0.4769 0.4167 0.2912

Table 18. Combined relative abundance (%) of the twelve bat species captured from four sub- areas (SAs) from June 2009 to May 2011 (n is number of bats captured; N is the total bats captured in from any sub-area).

Species SA1 SA2 SA3 SA4 Combined Relative abundance (n) Rhinolophus blassi - - - 5.77(3) 1.65(3) Rhinopoma hardwickii - - 4.65(2) - 1.10(2) Taphozous nudiventris 15.28(11) - 34.88(15) - 14.29(26) Taphozous perforatus 2.78(2) - 4.65(2) - 2.20(4) Scotoecus pallidus - - 4.65(2) - 1.10(2) Scotophilus heathii 25.00(18) 46.67(7) 23.26(10) 34.62(18) 29.12(53) Scotophilus kuhlii 1.39(1) - - 7.69(4) 2.75(5) Pipistrellus ceylonicus 20.83(15) - - 13.46(7) 12.09(22) Pipistrellus pipistrellus 26.39(19) 6.67(1) 27.91(12) 38.46(20) 28.57(52) Pipistrellus javanicus 4.17(3) 40.00(6) - - 4.95(9) Pipistrellus tenuis 2.78(2) - - - 1.10(2) Hypsugo savii 1.39(1) 6.67(1) - - 1.10(2) N 72 15 43 52 182

Table 19. Locality related diversity of various bat species captured from all the four sub-areas.

Diversity indices MHNP Chinji LSNP CP Overall Taxa_S 9 4 6 5 12 Individuals 72 15 43 52 182 Dominance_D 0.2025 0.3867 0.2601 0.2951 0.2059 Shannon_H 1.762 1.083 1.491 1.367 1.848 Simpson_1-D 0.7975 0.6133 0.7399 0.7049 0.7941 Evenness_e^H/S 0.6473 0.7386 0.7402 0.7844 0.5289 Menhinick 1.061 1.033 0.915 0.6934 0.8895 Margalef 1.871 1.108 1.329 1.012 2.114 Equitability_J 0.802 0.7814 0.8321 0.8491 0.7436 Fisher_alpha 2.715 1.785 1.896 1.363 2.884 Berger-Parker 0.2639 0.4667 0.3488 0.3846 0.2912

PART II. SPECIES DISTRIBUTION

The distribution of each species in the study area is shown by its map (Figure 8 to 11) and described as follows.

Family Rhinolophidae

1. Rhinolophus blasii. This species was recorded from Manawa (31º35.647, 074º27.660) in

Lahore district (SA4) (see Figure 8).

Family Rhinopomatidae

2. Rhinopoma hardwickii. This species was recorded from Noor Mahal (29º22.695,

071º40.132) in Bahawalpur district (SA3) (see Figure 9).

Family Emballonuridae

3. Taphozous nudiventris. This species was recorded from Ratowal (33º28.644,

072º43.638) in Attock district, from shrine of Abdul Maroof Shah at Mojgarh (29º01.132,

072º08.427) and from Derwar Fort (28º46.045, 071º20.210) in Bahawalpur districts (see

Figure 10).

4. T. perforatus. This species was also found co-roosting with T. nudiventris at all the

above mentioned three localities (see Figure 10).

Family Vespertilionidae

5. Scotoecus pallidus. This species was recorded from Bahawalpur Fish Hatchery

(29º23.186, 071º38.148) in Bahawalpur district (SA3) (see Figure 11).

6. Scotophilus heathii. This is the most widely distributed bat of the study area. It was

recorded from PMNH (33º43.194, 073º03.631), Rawal Town (33º40.966, 073º07.108),

NARC (33º39.892, 073º07.108) in SA1, Dalwal (32º42.725, 072º53.065) in SA2,

Bahawalpur Fish Hatchery (29º23.186, 071º38.148) in SA3 and Kalian Daas (31º12.037,

Figure 8. Distribution map of bats belonging to Family Rhinolophidae recorded from four sub-areas from June 2009 to May 2011.

Figure 9. Distribution map of bats belonging to Family Rhinopomatidae recorded from four sub-areas from June 2009 to May 2011.

Figure 10. Distribution map of bats belonging to Family Emballonuridae recorded from four sub-areas from June 2009 to May 2011.

Figure 11. Distribution map of bats belonging to Family Vespertilionidae recorded from four sub-areas from June 2009 to May 2011.

072º40.487), Govt. College Gojra (31º09.024, 072º41.077), Rasul Nagar (32º19.424,

073º46.623) and Pattoki (31º02.487, 073º52.536) in SA4 (see Figure 11).

7. Scotophilus kuhlii. This species was recorded from PMNH (33º43.194, 073º03.631) in

SA1 and Badian (31°29.223, 074°24.632) in SA4 (see Figure 11).

8. Pipistrellus ceylonicus. This species was recorded from PMNH (33º43.194, 073º03.631)

and Rawal Town (33º40.966, 073º07.108) in SA1 and Manawa (31º35.647, 074º27.660)

in SA4 (see Figure 11).

9. P. javanicus. This species was captured from PMNH (33º43.194, 073º03.631) and Rawal

Town (33º40.966, 073º07.108) in SA1 and from Dalwal (32º42.725, 072º53.065) and

Ucchali Lake (32º30.329, 072º00.305) in SA2 (see Figure 11).

10. P. pipistrellus. This species was recorded from NARC (33º39.892, 073º07.108), Rawal

Town (33º40.966, 073º07.108) in SA1, Dalwal (32º42.725, 072º53.065) in SA2,

Bahawalpur Fish Hatchery (29º23.186, 071º38.148) in SA3 and Kalian Daas (31º12.037,

072º40.487), Rasul Nagar (32º19.424, 073º46.623), Ali Pur Chattha (32º11.272,

074º09.361) and Pattoki (31º02.487, 073º52.536) in SA4 (see Figure 11).

11. P. tenuis. This species was captured from Rawal Town (33º40.966, 073º07.108) and

NARC (33º39.892, 073º07.108) in SA1 (see Figure 11).

12. Hypsugo savii. This species was recorded from PMNH (33º43.194, 073º03.631) in SA1

and from Ucchali Lake (32º30.329, 072º00.305) in SA2 (see Figure 11).

PART III. MORPHOLOGY

FAMILY EMBALLONURIDAE

Two species of this family exist in Pakistan and both were recorded from the study area.

The external body, cranial and bacular measurements of both these species captured from the

study area are discussed as follows.

1. Taphozous nudiventris

Body mass and external body measurements. Table 20 describes mean body mass and

external body measurements of T. nudiventris. Combined mean body mass of 26 bats captured

from SA1 and SA3 was 50.80 g ± 0.426 (SD). The mean head and body length was 86.87 mm ±

5.556 (SD) while the ear was 14.70 mm ± 2.541 (SD) long. The mean tragus height was 4.23

mm ± 0.563 (SD). Mean thumb and forearm length was 8.40 mm ± 2.640 (SD) and 71.00 mm ±

1.464 (SD). Length of 3rd metacarpal, and its 1st and 2nd phalanges were 62.87 mm ± 2.834 (SD),

27.23 mm ± 0.942 (SD) and 28.47 mm ± 1.968 (SD), respectively.

The length of 4th metacarpal and its 1st and 2nd phalanges was 50.37 mm ± 2.125 (SD),

14.03 mm ± 2.546 (SD) and 8.23 mm ± 0.842 (SD) respectively. The length of 5th metacarpal

and 1st phalanx was 40.83 mm ± 2.416 (SD) and 14.80 mm ± 1.623 (SD), respectively. The

average wing span was 354.13 mm ± 36.073 (SD). Tibia, calcar, hind foot, tail and penis were

29.27 mm ± 1.163 (SD), 6.50 mm ± 2.196 (SD), 15.57 mm ± 1.474 (SD), 27.57 mm ± 12.187

(SD) and 6.08 mm ± 1.145 (SD) long, respectively. The ear (F cal = 8.620; df = 25), claw (Fcal =

=5.891; df = 25), and tail lengths (Fcal = 47.287; df = 25) of the two populations varied significantly at p<0.05 (Table 20).

Table 20. Combined mean body mass (g) and external body measurements (mm) of Taphozous nudiventris captured from Margalla Hills National Park (SA1) and Lal Suhanatra National Park (SA3) from June 2009 to May 2011 (n is the number of specimens).

SA1 SA3 Combined Statistics Body Parameters n= 11 n= 15 n= 26 F cal P Body mass 54.99±6.973(11) 48.47±14.508(15) 50.80±0.426(26) 2.314 0.152 Head and body length 87.43±4.903 (11) 82.08±20.556 (15) 86.87±5.556 (26) 0.126 0.728 Ear length 16.36±1.952 (7) 14.05±3.940 (8) 14.70±2.541 (15) 8.620 0.012** Tragus length 4.36±0.476 (7) 4.02±1.055 (8) 4.23±0.563 (15) 0.618 0.446 Thumb length 8.14±0.988 (7) 7.95±3.053 (8) 8.40±2.640 (15) 0.117 0.738 Claw length 2.29±.756 (7) 2.59±0.873 (8) 2.73±0.776 (15) 5.891 0.030* Forearm length 71.43±1.813 (11) 66.96±16.843 (15) 71.00±1.464 (26) 1.136 0.306 Length of 3rd metacarpal 63.30±3.099 (10) 59.38±14.744 (11) 62.87±2.834 (21) 0.280 0.606 1st phalanx on 3rd metacarpal 27.29±0.636 (10) 25.67±6.512 (11) 27.23±0.942 (21) 0.038 0.849 2nd phalanx on 3rd metacarpal 28.57±0.345 (10) 26.82±7.066 (11) 28.47±1.968 (21) 0.035 0.855 Length of 4th metacarpal 50.14±1.864 (10) 50.56±2.441 (11) 50.37±2.125 (21) 0.015 0.954 1st phalanx on 4th metacarpal 14.50±2.291 (10) 13.37±3.719 (12) 14.03±2.546 (22) 0.423 0.527 2nd phalanx on 4th metacarpal 8.43±0.450 (10) 7.79±2.049 (12) 8.23±0.842 (22) 0.689 0.421 Length of 5th metacarpal 40.50±2.198 (9) 38.54±9.633 (13) 40.83±2.416 (22) 0.227 0.642 1st phalanx on 5th metacarpal 14.21±2.177 (9) 14.02±3.413 (13) 14.80±1.623 (22) 1.807 0.202 Wing span 335.00±27.031 (8) 333.77±86.070 (9) 354.13±36.073 (17) 4.657 0.050 Tibia length 29.57±1.134 (8) 27.63±6.914 (9) 29.27±1.163 (17) 0.895 0.361 Calcar length 7.21±2.157 (8) 6.29±2.320 (9) 6.50±2.196 (17) 1.432 0.253 Hind foot length 15.93±1.790 (8) 14.78±3.621 (9) 15.57±1.474 (17) 0.778 0.394 Tail length 38.71±7.825 (8) 27.06±12.721 (9) 27.57±12.187 (17) 47.287 0.000** Penis length 6.20±0.758 (9) 5.71±1.773 (9) 6.08±1.145 (18) 0.325 0.578

Cranial measurements. Mean cranial measurements of two specimens of T. nudiventris

captured from the study area are mentioned in Table 21. The breadth of the braincase and the

zygomatic bone were 10.87 mm ± 0.180 (SD) mm and 14.15 mm ± 1.688 (SD), respectively.

The postorbital constriction was 5.18 mm ± 0.216 (SD) long. Condylo-canine and condylo-basal length was 19.27 mm ± 1.024 (SD) and 24.00 mm ± 1.257 (SD), respectively. The greatest skull, maxillary tooth row, mandibular tooth row and the mandible lengths were 26.16 mm ± 0.323

(SD), 8.75 mm ± 2.425 (SD), 10.08 mm ± 1.976 (SD) and 17.53 mm ± 1.149 (SD), respectively.

Anterior and posterior palatal width of the skull was 4.62 mm ± 1.365 (SD) and 11.16 mm ±

1.455 (SD), respectively.

Bacular measurements. Mean total baculum length of the two specimens was 0.58 mm ± 0.017

(SD) with shaft length of 0.54 mm ± 0.069 (SD). The proximal and distal lengths of the baculum were 0.04 mm ± 0.052 (SD) and 0.0 mm, respectively. The proximal and distal bacular widths were 0.25 mm ± 0.035 (SD) and 0.20 mm ± 0.000 (SD), respectively. The bacula were 0.10 mm

± 0.000 (SD) high (Table 22, Figure 12).

Principal Component Analysis. A comparison of the two bat populations through Principal

Component Analysis (PCA) using head and body length, ear length, tragus length, claw length, forearm length, length of third metacarpal, length of first phalanx on third metacarpal, length of second phalanx on third metacarpal, length of fourth metacarpal, length of first phalanx on fourth metacarpal, length of second phalanx on fourth metacarpal, length of fifth metacarpal, length of first phalanx on fifth metacarpal, wingspan, tibia length, calcar length, hind foot length and tail length as multiple variables showed that maximum variation was due to first three components

(Figure 13). The PCA suggests that the two populations are different from each other but the small data set does not confirm this generalization and needs to be studied with a large data set.

Table 21. Mean cranial measurements (mm) of Taphozous nudiventris captured from Rattowal (SA1) from June 2009 to May 2011 (n is the number of specimens).

Cranial Parameters n= 2 Breadth of braincase 10.87±0.180 Zygomatic breadth 14.15±1.688 Postorbital constriction 5.18±0.216 Condylo-canine length 19.27±1.024 Condylo-basal length 24.00±1.257 Greatest length of skull 26.16±0.323 Maxillary toothrow 8.75±2.425 Anterior palatal width 4.62±1.365 Posterior palatal width 11.16±1.455 Mandibular toothrow 10.08±1.976 Mandible length 17.53±1.149

Table 22. Mean bacular measurements (mm) of Taphozous nudiventris captured from Rattowal (SA1) and Mojgarh (SA3) from June 2009 to May 2011 (n is the number of specimens).

Bacular Parameters n= 2 Total length of baculum 0.58±0.017 Length of shaft 0.54±0.069 Length of proximal branch 0.04±0.052 Length of distal branch 0.00±0.000 Width of proximal branch 0.25±0.035 Width of distal branch 0.20±0.000 Height of baculum 0.10±0.000

(a) (b)

Figure 12. Photograph of the bacula of T. nudiventris, captured from SA3 (a) showing its shape and (b) position of baculum in penis of T. nudiventris.

(a)

90 80 70 60 50 40 igenvalue %

E 30 20 10 0 0 2 4 6 8 10 12 14 16 18 Component

25 (b) 20

Component 2 -80 -64 -48 -32 -16 16 32 48 64 -5

-10

-15

-20

Components Eigen value % Variance 1 1323.27 86.01 2 145.526 9.459 3 30.6951 1.9952 4 14.784 0.96107

Figure 13. Principal component analysis of the two Taphozous nudiventris populations recoded from SA1 and SA3 (Scree plot (a), scatter plot (b) and the Table of Eigen values (c) for the first four components is shown above. PC 1 loads size while PC 2 loads locality).

2. Taphozous perforatus

Body mass and external body measurements. Table 23 shows mean body mass and external

body measurements of four specimens of T. perforatus captured form two different sub-areas i.e.

Margalla Hills National Park and Lal Suhanara National Park. The mean body mass of all the

four specimens captured from the study area was 43.80 g ± 9.997 (SD). The head and body length measured 84.30 mm ± 5.450 (SD) long. The mean average ear length of all the four

specimens was 15.30 mm ± 2.110 (SD) with tragus length of 4.80 mm ± 1.789 (SD). The length

of thumb and that of claw was 7.20 mm ± 0.447 (SD) and 3.40 mm ± 0.418 (SD) respectively.

The forearm was 64.30 mm ± 3.457 (SD) long. The lengths of the 3rd metacarpal and its 1st and

2nd phalanges were 60.40 mm ± 4.436 (SD) and 25.40±6.417, respectively. The length of the 4th metacarpal and 1st and 2nd phalanges on 4th metacarpal were 50.00 mm ± 2.894 (SD), 13.80 mm

± 1.681 (SD) and 9.50 mm ± 1.500 (SD), respectively. The length of 5th metacarpal was 41.80

mm ± 4.919 (SD) while its 1st phalanx was 14.20 mm ± 2.139 (SD) long. Average wing span was

323.60 mm ± 33.269 (SD). Tibia length was measured 27.40 mm ± 0.652 (SD). The calcar, hind foot, tail and penis lengths were 6.30 mm ± 1.396 (SD), 14.60 mm ± 1.949 (SD), 22.10 mm ±

2.702 (SD) and 5.80 mm ± 2.080 (SD), respectively.

Cranial measurements. Table 24 presents different cranial measurements of a single specimen of T. perforatus. The breadth of the braincase and the zygomatic breadth were 10.01 mm and

12.01 mm respectively. The postorbital constriction was 5.54 mm while condylo-canine length and condylo-basal lengths were 19.58 mm and 20.86 mm respectively. The greatest length of skull was 22.24 mm, maxillary toothrow 7.32 mm, anterior palatal width 6.07 mm, posterior palatal width 10.13 mm, mandibular toothrow length 10.25 mm and the length of mandible was recorded as 16.25 mm.

Table 23. Mean body mass (g) and external body measurements (mm) of Taphozous perforatus captured from Margalla Hills National Park (SA1) and Lal Suhanara National Park (SA3) from June 2009 to May 2011 (n is the number of specimens).

SA1 SA3 Combined Body Parameters n= 2 n= 2 n= 4 Body mass 48.85±0.212 40.43±12.544 43.80±9.997 Head and body length 79.25±1.061 87.67±4.041 84.30±5.450 Ear length 16.25±2.475 14.67±2.082 15.30±2.110 Tragus length 6.00±2.828 4.00±0.000 4.80±1.789 Thumb length 7.50±0.707 7.00±0.000 7.20±0.447 Claw length 3.75±0.354 3.17±0.289 3.40±0.418 Forearm length 61.75±0.354 66.00±3.606 64.30±3.457 Length of 3rd metacarpal 59.50±7.071 61.00±3.606 60.40±4.436 1st phalanx on 3rd metacarpal 25.50±2.121 26.00±2.000 25.80±1.789 2nd phalanx on 3rd metacarpal 22.50±11.314 27.33±2.082 25.40±6.417 Length of 4th metacarpal 50.50±2.828 49.67±3.512 50.00±2.894 1st phalanx on 4th metacarpal 13.25±3.182 14.17±0.289 13.80±1.681 2nd phalanx on 4th metacarpal 10.75±1.768 8.67±0.577 9.50±1.500 Length of 5th metacarpal 46.00±5.657 39.00±1.732 41.80±4.919 1st phalanx on 5th metacarpal 13.25±3.889 14.83±0.289 14.20±2.139 Wing span 304.00±22.627 336.67±36.295 323.60±33.269 Tibia length 27.25±1.061 27.50±0.500 27.40±0.652 Calcar length 7.75±0.354 5.33±0.577 6.30±1.396 Hind foot length 14.00±0.000 15.00±2.646 14.60±1.949 Tail length 24.75±1.061 20.33±1.528 22.10±2.702 Penis length 6.25±1.768 5.50±2.598 5.80±2.080

Table 24. Cranial measurements (mm) of Taphozous perforatus captured from Rattowal (SA1) from June 2009 to May 2011 (n is the number of specimens).

Cranial Parameters n=1 Breadth of braincase 10.01 Zygomatic breadth 12.01 Postorbital constriction 5.54 Condylo-canine length 19.58 Condylo-basal length 20.86 Greatest length of skull 22.24 Maxillary toothrow 7.32 Anterior palatal width 6.07 Posterior palatal width 10.13 Mandibular toothrow 10.25 Mandible length 16.25

Bacular measurements. Various bacular measurements of a single specimen of T. peforatus

captured from the study area are given in Table 25 the baculum shape is shown in Figure 14. The

total length of the baculum was measured as 0.69 mm with shaft length of 0.61 mm. The proximal and distal bacular lengths were 0.07 mm and 0.00 mm, respectively. The proximal and distal breadths of baculum were 0.27 mm and 0.22 mm respectively. The bacular height was 0.07 mm.

FAMILY VESPERTILIONIDAE

Thirty species of this Family exist in Pakistan. Of these, eight were recorded from the study area. The external body, cranial and bacular measurements of both these species captured from the study area are discussed as follows.

1. Scotoecus pallidus

Body mass and external body measurements. The mean body mass of two S. pallidus was

11.50 g ± 0.566 (SD) with head and body length of 56.50 mm ± 3.536 (SD). The ear and tragus lengths were 9.50 mm ± 0.707 (SD) and 7.00 mm ± 0.000 (SD), respectively. The lengths of thumb and claw were 6.00 mm ± 0.000 (SD) and 2.50 mm ± 0.707 (SD), respectively. The forearm was 35.50 mm ± 0.707 (SD) long. The lengths of 3rd metacarpal and its 1st and 2nd phalanges were 33.75 mm ± 0.354 (SD), 12.00 mm ± 0.707 (SD) and 11.00 mm ± 0.707 (SD), respectively. Lengths of 4th metacarpal and its 1st and 2nd phalanges were 31.75 mm ± 0.354

(SD), 12.00 mm ± 0.707 (SD) and 11.00 mm ± 0.707 (SD), respectively. The 5th metacarpal was

32.25 mm ±0.354 (SD) long with its 1st phalanx length of 8.75 mm ± 0.354 (SD), respectively.

Mean wing span, tibia, calcar, hind foot, tail and penis lengths were 258.00 mm ± 2.828 (SD),

13.00 mm ± 0.000 (SD), 4.00 mm ± 0.000 (SD), 8.50 mm ± 0.707 (SD), 35.50 mm ± 3.536 (SD)

and 7.00 mm ± 0.00 (SD), respectively (Table 26).

Table 25. Bacular measurements (mm) of Taphozous perforatus captured from SA1 from June 2009 to May 2011 (n is the number of specimens).

Bacular Parameters n= 1 Total length of baculum 0.69 Length of shaft 0.61 Length of proximal branch 0.07 Length of distal branch 0.00 Width of proximal branch 0.27 Width of distal branch 0.22 Height of baculum 0.07

(a) (b)

Figure 14. (a) Baculum of T. perforatus captured from Sub-area 1 and (b) shows its position within the penis.

Table 26. Mean body mass (g) and external body measurements (mm) of Scotoecus pallidus captured from SA3 from June 2009 to May 2011 (n is the number of specimens).

Body Parameters n= 2 Body mass 11.50±0.566 Head and body length 56.50±3.536 Ear length 9.50±0.707 Tragus length 7.00±0.000 Thumb length 6.00±0.000 Claw length 2.50±0.707 Forearm length 35.50±0.707 Length of 3rd metacarpal 33.75±0.354 1st phalanx on 3rd metacarpal 12.00±0.707 2nd phalanx on 3rd metacarpal 11.00±0.707 Length of 4th metacarpal 32.25±0.354 1st phalanx on 4th metacarpal 11.50±0.707 2nd phalanx on 4th metacarpal 10.00±1.414 Length of 5th metacarpal 31.75±0.354 1st phalanx on 5th metacarpal 8.75±0.354 Wing span 258.00±2.828 Tibia length 13.00±0.000 Calcar length 4.00±0.000 Hind foot length 8.50±0.707 Tail length 35.50±3.536 Penis length 7.00± 0.00

± 0.700 (SD) mm and 9.91 mm ± 0.323 (SD), respectively. The post-orbital constriction was

4.37 mm ± 0.503 (SD). The condylo-canine and condylo-basal lengths were 14.69 mm ± 0.341

(SD) and 15.27 mm ± 0.323 (SD), respectively. The greatest length of skull was 15.46 mm ±

0.449 (SD), maxillary toothrow length was 4.34 mm ± 1.868 (SD), anterior palatal width 4.22 mm ± 0.862 (SD) mm, posterior palatal width 5.35 mm ± 1.275 (SD), the mandibular toothrow length 4.32 mm ± 1.365 (SD) and mandible length was measured as 9.64 mm ± 2.425 (SD)

(Table 27).

Bacular measurements. The total length of the baculum of a single S. pallidus captured from

SA3 was 5.0 mm with shaft length of 4.1 mm. The proximal and distal bacular lengths were 0.6

mm and 0.3 mm, respectively. The proximal and distal breadths of the baculum were 1.0 mm and

0.5 mm, respectively. The baculum height was 0.5 mm (Table 28, Figure 15).

2. Scotophilus heathii

Body mass and external body measurements. Body mass and external body measurements of

Scotophilus heathii specimens captured from all the four sub-areas are given in Table 29. The mean body mass of all forty nine specimens captured from the study area was 35.97 g ± 5.943

(SD). Head and body length was 79.46 mm ± 6.941 (SD). Mean ear and tragus lengths were

12.65 mm ± 1.622 (SD) and 7.65 mm ± 1.047 (SD), respectively. Thumb and claw lengths varied significantly among the four populations (Fcal= 4.165, P= 0.007 and Fcal= 2.861, P= 0.034, respectively) and were 6.93 mm ± 0.927 (SD) and 2.57 mm ± 0.717 (SD) long, respectively. The mean forearm length was 58.69 mm ± 2.929 (SD). The lengths of 3rd metacarpal, and 1st and 2nd phalanx on 3rd metacarpal were 55.02 mm ± 4.338 (SD), 19.93 mm ± 2.273 (SD) and 15.26 mm

± 2.301 (SD), respectively. Similarly length of 4th metacarpal and lengths of 1st and 2nd phalanx

on 4th metacarpal were 52.09 mm ± 7.182 (SD), 15.80 mm ± 1.203 (SD) and 12.30 mm ± 1.552

Table 27. Mean cranial measurements (mm) of Scotoecus pallidus captured from Bahawalpur Fish Hatchery (SA3) from June 2009 to May 2011 (n is the number of specimens). .

Cranial Parameters n= 2 Breadth of braincase 7.12±0.700 Zygomatic breadth 9.91±0.323 Postorbital constriction 4.37±0.503 Condylo-canine length 14.69±0.341 Condylo-basal length 15.27±0.323 Greatest length of skull 15.46±0.449 Maxillary toothrow 4.34±1.868 Anterior palatal width 4.22±0.862 Posterior palatal width 5.35±1.275 Mandibular toothrow 4.32±1.365 Mandible length 9.64±2.425

Table 28. Bacular measurements (mm) of Scotoecus pallidus captured from Bahawalpur Fish Hatchery (SA3) from June 2009 to May 2011 (n is the number of specimens).

Bacular Parameters n= 1 Total length of baculum 5.0 Length of shaft 4.1 Length of proximal branch 0.6 Length of distal branch 0.3 Width of proximal branch 1.0 Width of distal branch 0.5 Height of baculum 0.5

Figure 15. Dorsal view of the baculum of Scotoecus pallidus captured from SA3

Table 29. Combined mean body mass (g) and external body measurements (mm) of Scotophilus heathii captured from Margalla Hills National Park (SA1), Chinji National Park and (SA2), Lal Suhanara National Park (SA3) and some localities of Central Punjab (SA4) from June 2009 to May 2011 (n is the number of specimens).

SA1 SA2 SA3 SA4 Combined Statistics Body Parameters n= 18 n= 7 n= 10 n= 18 n= 53 F cal P Body mass 38.5±7.20(18) 32.5±2.81(7) 35.8±4.31(10) 34.8±6.84(18) 35.97±5.943 (53) 2.304 0.073 Head and body length 79.1±8.77(17) 77.3±4.73(7) 82.0±7.67(7) 78.2±3.87(18) 79.46±6.941 (49) 2.546 0.155 Ear length 11.7±1.91(9) 12.3±1.32(4) 13.5±1.51(8) 13.2±0.68(7) 12.65±1.622 (28) 1.779 0.152 Tragus length 7.4±1.27(9) 8.1±1.03(4) 7.3±1.16(8) 7.9±0.65(7) 7.65±1.047 (28) 0.502 0.800 Thumb length 6.4±1.24(12) 7.8±0.87(5) 6.9±0.58(5) 7.2±0.26(15) 6.93±0.927(n= 37) 4.165 0.007** Claw length 2.4±0.88(12) 2.4±0.85(5) 3.0±0.46(5) 2.3±0.52(15) 2.57±0.717 (37) 2.861 0.034* Forearm length 60.3±2.80(17) 56.6±2.10(7) 58.8±3.09(7) 57.5±2.53(18) 58.69±2.929 (49) 0.425 0.854 Length of 3rd metacarpal 53.8±7.29(12) 56.4±0.85(5) 55.1±1.88(5) 55.8±1.60(15) 55.02±4.338 (37) 0.332 0.913 1st phalanx on 3rd metacarpal 19.6±1.90(12) 20.9±1.11(5) 20.1±3.63(5) 19.6±0.92(15) 19.93±2.273 (37) 0.470 0.823 2nd phalanx on 3rd metacarpal 14.2±2.37(12) 17.0±1.08(5) 15.1±2.95(5) 15.9±0.86(15) 15.26±2.301 (37) 1.014 0.443 Length of 4th metacarpal 48.9±11.98(12) 54.1±0.75(5) 53.1±2.04(5) 54.1±2.22(15) 52.09±7.182 (37) 0.647 0.692 1st phalanx on 4th metacarpal 15.8±0.90(12) 15.4±2.46(5) 15.6±0.99(5) 16.3±0.88(15) 15.80±1.203 (37) 0.096 0.996 2nd phalanx on 4th metacarpal 12.7±0.75(12) 12.5±1.00(5) 11.8±2.60(5) 12.3±0.88(15) 12.30±1.552 (37) 0.304 0.928 Length of 5th metacarpal 49.3±5.98(15) 51.6±1.11(7) 49.9±2.95(7) 49.9±1.53(15) 49.96±3.813 (37) 0.534 0.777 1st phalanx on 5th metacarpal 11.4±0.81(15) 11.1±1.31(7) 11.0±0.80(7) 10.9±0.80(15) 11.15±0.864 (37) 0.572 0.748 Wing span 293.9±65.57(5) 341.8±19.81(3) 331.5±117.48(3) 347.8±53.39(7) 324.11±78.376 (44) 0.868 0.534 Tibia length 23.8±4.29(12) 24.1±1.31(5) 25.2±1.25(5) 23.3±1.99(15) 24.13±2.762 (44) 0.713 0.643 Calcar length 8.6±2.15(15) 8.1±1.70(7) 7.3±2.13(7) 8.7±1.47(15) 8.16±1.939 (44) 0.973 0.467 Hind foot length 12.5±2.16(15) 12.0±0.82(7) 11.7±2.08(7) 13.3±1.33(15) 12.39±1.700 (44) 1.083 0.404 Tail length 54.1±9.2(15) 61.7±6.4 52.7±6.03 53.6±5.71 55.00±7.360 (44) 1.160 0.364 Penis length 7.4±1.29(15) 8.1±1.03(7) 7.5±1.07(7) 7.9±0.66(15) 7.25±0.974 (18) 2.359 0.067

(SD), respectively. The 5th metacarpal and its 1st phalanx were 49.96 mm ± 3.813 (SD) and 11.15

mm ± 0.864 (SD) long, respectively. The mean wing span was 324.11 mm ± 78.376 (SD). Tibia,

calcar and hind foot were 24.3 mm ± 2.762 (SD), 8.16 mm ± 1.939 (SD) and 12.76 mm ± 2.799

(SD) long, respectively. The average tail and penis lengths were 55.00 mm ± 7.360 (SD) and

7.25 mm ± 0.974 (SD), respectively.

Cranial measurements. The mean breadth of braincase of all the twenty specimens captured from all the four susb-areas i.e. Margalla Hills National Park (n= 6), Chinji National Park (n= 3),

Lal Suhanara National Park (n= 8) and some areas of central Punjab (n= 3) was recorded as 9.99 mm ± 0.951 (SD), zygomatic breadth 14.31 mm ± 1.140 (SD), postorbital constriction 5.38 mm

± 0.441 (SD). The condylo-canine and condylo-basal lengths were 19.01 mm ± 1.251 (SD) and

19.74 mm ± 1.185 (SD) respectively. The greatest length of skull was 21.39 mm ± 1.378 (SD), maxillary toothrow 7.82 mm ± 0.578 (SD), anterior palatal width 6.82 mm ± 0.577 (SD), posterior palatal width 9.38 mm ± 0.636 (SD), mandibular toothrow length 8.41 mm ± 0.523

(SD) and the length of mandible was recorded as 16.08 mm ± 0.882 (SD) (Table 30).

Bacular measurements. Table 31 describes various bacular measurements of ten specimens captured from the study area SA 1, SA3 and SA4. Mean total bacular length was measured 1.76 mm ± 0.150 (SD) with shaft length of 1.51 mm ± 0.232 (SD). The proximal and distal baular lengths were 0.19 mm ± 0.078 (SD) and 0.04 mm ± 0.034 (SD), respectively while proximal and distal bacular breadths were 0.95 mm ± 0.120 (SD) and 0.46 mm ± 0.058 (SD), respectively. The bacular height was 0.29 mm ± 0.074 (SD) (Figure 16).

Principal Component Analysis. A comparison of the four bat populations through Principal

Component Analysis (PCA) using head and body length, ear length, tragus length, claw length, forearm length, length of third metacarpal, length of first phalanx on third metacarpal, length of

Table 30. Combined mean cranial measurements (mm) of Scotophilus heathii captured from Margalla Hills National Park (SA1), Chinji National Park (SA2), Lal Suhanara National Park (SA3) and Central Punjab (SA4) from June 2009 to May 2011 (n is the number of specimens).

SA1 SA2 SA3 SA4 Combined Cranial Parameters n=6 n=3 n=8 n=3 n=20 Breadth of braincase 9.77±0.333 11.03±1.272 9.53±0.573 10.96±1.742 9.99±0.951 Zygomatic breadth 14.79±0.968 15.41±0.213 13.77±1.131 13.35±0.697 14.31±1.140 Postorbital constriction 5.19±0.837 5.61±0.159 5.24±0.364 5.63±0.952 5.38±0.441 Condylo-canine length 10.25±5.021 19.42±1.686 18.82±2.088 17.90±1.000 19.01±1.251 Condylo-basal length 10.94±4.834 20.04±1.474 19.44±2.128 19.35±1.970 19.74±1.185 Greatest length of skull 18.57±6.069 21.03±2.386 20.80±2.027 21. 80±1.245 21.39±1.378 Maxillary toothrow 6.97±1.660 8.78±0.860 7.35±0.509 7.84±0.341 7.82±0.578 Anterior palatal width 6.39±1.339 6.52±0.864 6.80±0.581 6.62±0.018 6.82±0.577 Posterior palatal width 8.69±2.138 9.64±0.281 9.13±0.727 9.55±0.216 9.38±0.636 Mandibular toothrow 7.49±2.332 8.76±0.458 8.14±0.597 8.69±0.000 8.41±0.523 Mandible length 14.58±3.628 16.14±0.493 15.80±1.186 16.62±0.126 16.08±0.882

Table 31. Combined mean bacular measurements (mm) of Scotophilus heathii captured from Margalla Hills National Park (SA1), Lal Suhanara National Park (SA3) and Central Punjab (SA4) from June 2009 to May 2011 (n is the number of specimens).

SA1 SA3 SA4 Combined Bacular Parameters n= 4 n= 5 n= 1 n= 10 Total length of baculum 1.81±0.127 1.77±0.145 1.52 1.76±0.150 Length of shaft 1.59±0.212 1.54±0.150 1.03 1.51±0.232 Length of proximal branch 0.18±0.101 0.19±0.066 0.27 0.19±0.078 Length of distal branch 0.03±0.023 0.06±0.041 0.03 0.04±0.034 Width of proximal branch 0.93±0.155 0.99±0.089 0.83 0.95±0.120 Width of distal branch 0.46±0.023 0.46±0.084 0.44 0.46±0.058 Height of baculum 0.48±0.032 0.45±0.070 0.29 0.44±0.074

Figure 16. Dorsal view of the bacula of Scotophilus heathii (1. Bat lab 47NARC, 2. Bat lab 4BFH, 3. Bat lab 8BFH, 4. Bat lab 13BFH, 5. Bat lab 14 BFH, 6. Bat lab 39PMNH, 7. Bat lab 54Gojra, 8. Bat lab 114 RTown, 9. Bat lab 115 RTown, 10. Bat lab102 Pattoki).

second phalanx on third metacarpal, length of fourth metacarpal, length of first phalanx on fourth

metacarpal, length of second phalanx on fourth metacarpal, length of fifth metacarpal, length of

first phalanx on fifth metacarpal, wingspan, tibia length, calcar length, hind foot length and tail length as variables showed that maximum variation was due to first three components (Figure

17). The PCA suggests that the SA3 populations is different from other three populations but the small data set does not confirm this generalization and needs to be studied with a large data set.

3. Scotophilus kuhlii

Body mass and external body measurements. Table 32 describes mean body mass and external body measurements of five specimens of S. kuhlii captured from SA1 (n= 1) and SA4

(n= 4). The mean body mass of all the five specimens was 22.2 g ± 2.59 (SD), head and body length of was 72.10 mm ± 8.096 (SD). The ear and the tragus lengths were 12.10 mm ± 2.53

(SD) and 6.60 mm ± 0.74 (SD), respectively. Thumb and claw lengths were 6.42 mm ± 0.94

(SD) and 2.70 mm ± 0.45 (SD), respectively. The forearm was 49.40 mm ± 3.03 (SD) long.

Length of the 3rd metacarpal was 47.30 mm ± 4.64 (SD) with its 1st phalanx 18.10 mm ± 0.96

(SD) and 2nd phalanx 13.60 mm ± 0.65 (SD). The length of 4th metacarpal and lengths of its 1st and 2nd phalanges were 44.90 mm ± 3.11 (SD), 15.40 mm ± 1.64 (SD) and 10.70 mm ± 2.08

(SD), respectively. The 5th metacarpal was 41.10 mm ± 3.27 (SD) long with length of its 1st phalanx 9.30 mm ± 0.57 (SD). Wing span was 293.20 mm ± 35.48 (SD), tibia length 20.10 mm

± 1.64 (SD), calcar length 8.00 mm ± 1.17 (SD) while mean hind foot, tail and penis lengths were 10.60 mm ±0.65 (SD), 42.40 mm ± 4.04 (SD) and 5.00 mm (n= 1), respectively.

Cranial measurements. The mean breadth of the braincase and zygomatic breadth of all the five specimens were 9.59 mm ± 1.610 (SD) and 13.43 mm ± 0.847 (SD), respectively. Post orbital constriction was 4.73 mm ± 0.394 (SD). The condylo-canine and condylo-basal lengths were

(a)

80 70 60 50 40

Eigenvalue % 30 20 10 0 0 2 4 6 8 10 12 14 16 18 Component

B B (b) 1.6 B B P B B 1.2 B

0.8

0.4

-3.6 -3 -2.4 -1.8 -1.2 -0.6 0.6 1.2 Component 2 Component

R-0.4 Pt Pt N D N RP DD Pt G G -0.8 B N G D

-1.2

R -1.6

-2 (c) Component 1

Components Eigen value % Variance 1 1323.27 86.01 2 145.526 9.459 3 30.6951 1.9952 4 14.784 0.96107

Figure 17. Principal component analysis of the four Scotophilus heathii populations recoded from SA1 (PMNH = P; Rawal Town = R; NARC = N), SA2 (Dalwal = D), SA3 (Bahawalpur Fish Hatchery =B) and SA4 (Gojra = G and Pattoki = Pt) Scree plot (a), scattered plot (b) and the Table of eigen values (c) for the first four components is shown above. PC 1 loads size while PC 2 loads locality).

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Table 32. Combined mean body mass (g) and external body measurements (mm) of Scotophilus kuhlii captured from Margalla Hills National Park (SA1) and Central Punjab (SA4) from June 2009 to May 2011 (n is the number of specimens).

SA1 SA4 Combined Body Parameters n= 1 n= 4 n= 5 Body mass 26.00 21.25±1.708 22.2±2.59 Head and body length 86.00 68.75±3.202 72.10±8.096 Ear length 16.00 11.13±1.493 12.10±2.53 Tragus length 5.50 6.88±0.479 6.60±0.74 Thumb length 5.60 6.63±0.946 6.42±0.94 Claw length 2.50 2.75±0.500 2.70±0.45 Forearm length 44.50 50.63±1.493 49.40±3.03 Length of 3rd metacarpal 42.00 48.63±4.131 47.30±4.64 1st phalanx on 3rd metacarpal 18.50 18.00±1.080 18.10±0.96 2nd phalanx on 3rd metacarpal 13.00 13.75±0.645 13.60±0.65 Length of 4th metacarpal 39.50 46.25±0.866 44.90±3.11 1st phalanx on 4th metacarpal 18.00 14.75±0.866 15.40±1.64 2nd phalanx on 4th metacarpal 8.00 11.38±1.652 10.70±2.08 Length of 5th metacarpal 35.50 42.50±1.080 41.10±3.27 1st phalanx on 5th metacarpal 8.50 9.50±0.408 9.30±0.57 Wing span 240.00 306.50±22.338 293.20±35.48 Tibia length 17.50 20.75±0.866 20.10±1.64 Calcar length 7.00 8.25±1.190 8.00±1.17 Hind foot length 10.00 10.75±0.645 10.60±0.65 Tail length 38.00 43.50±3.697 42.40±4.04 Penis length 5.00 - 5.00±0

101

17.62 mm ± 0.462 (SD) and 18.01 mm ± 0.302 (SD), respectively. The greatest length of the

skull was 18.98 mm ± 0.613 (SD). The maxillary toothrow was 18.98 mm ± 0.613 (SD), anterior palatal width 6.27 mm ± 0.486 (SD), posterior palatal width 5.79 mm ± 0.356 (SD), mandibular

toothrow and mandible lengths were 7.53 mm ± 0.812 (SD) and 14.41 mm ± 1.173 (SD),

respectively (Table 33).

Bacular measurements. Bacular measurements of a single male S. kuhlii captured from SA1 are

given in Table 34 (Figure 18). The total length of the baculum was 1.74 mm with a shaft length

of 1.52 mm. The proximal and distal bacular lengths were 0.07 mm and 0.15 mm, respectively.

The proximal and distal bacular widths were 1.05 mm and 0.49 mm, respectively while the

baculum was 0.49 mm in height.

4. Pipistrellus ceylonicus

Body mass and external body measurements. Mean body mass and external body

measurements of twenty two P. ceylonicus captured from SA1 and SA4 are given in Table 35.

The mean body mass was 3.54 g ± 1.549 (SD) while head and body lengths were 63.60 mm ±

7.486 (SD). The ear and tragus lengths were 8.06 mm ± 1.724 (SD) and 4.70 mm ± 0.694 (SD),

respectively. Thumb and claw lengths were 3.89 mm ± 0.689 (SD) and 1.41 mm ± 0.350 (SD)

respectively. The length of forearm was 29.92 mm ± 2.492 (SD). The length of 3rd metacarpal

and 1st and 2nd phalanges on 3rd metacarpal was 25.85 mm ± 3.205 (SD), 12.18 mm ± 2.246 (SD)

and 11.90 mm ± 2.741 (SD), respectively. The lengths of 4th metacarpal and its 1st and 2nd phalanges were 25.49 mm ± 3.462 (SD), 11.05 mm ± 1.416 (SD) and 8.92 mm ± 1.272 (SD), respectively. The length of 5th metacarpal and 1st phalanx on 5th metacarpal was 25.52 mm ±

3.271 (SD) and 9.15 mm ± 1.565 (SD), respectively. Mean wing span, tibia length, calcar, hind

foot, tail and penis lengths were 192.85 mm ± 12.343 (SD), 12.35 mm ± 1.181 (SD), 5.35 mm ±

1.387 (SD), 6.34 mm ± 1.187 (SD), 25.68 mm ± 3.442 (SD) and 5.57 mm ± 0.753 (SD), respectively.

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Table 33. Combined mean cranial measurements (mm) of Scotophilus kuhlii captured from Margalla Hills National Park (SA1) and Central Punjab (SA4) from June 2009 to May 2011 (n is the number of specimens).

Cranial Parameters SA1 SA4 Combined n=1 n=4 n=5 Breadth of braincase 12.45 8.88±0.243 9.59±1.610 Zygomatic breadth 14.94 13.05±0.089 13.43±0.847 Postorbital constriction 4.75 4.72±0.455 4.73±0.394 Condylo-canine length 17.75 17.58±0.526 17.62±0.462 Condylo-basal length 18.06 18.00±0.347 18.01±0.302 Greatest length of skull 18.01 19.23±0.323 18.98±0.613 Maxillary toothrow 5.44 6.48±0.162 6.27±0.486 Anterior palatal width 5.56 5.84±0.385 5.79±0.356 Posterior palatal width 6.22 8.27±0.135 7.86±0.922 Mandibular toothrow 8.74 7.23±0.524 7.53±0.812 Mandible length 16.36 13.93±0.506 14.41±1.173

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Table 34. Bacular measurements (mm) of Scotophilus kuhlii captured from Pakistan Museum of Natural History (SA1) from June 2009 to May 2011 (n is the number of specimens).

Bacular Parameters n=1 Total length of baculum 1.74 Length of shaft 1.52 Length of proximal branch 0.07 Length of distal branch 0.15 Width of proximal branch 1.05 Width of distal branch 0.49 Height of baculum 0.49

Figure 18. Dorsal view of the baculum of Scotophilus kuhlii captured from SA1 from June 2009 to May 2011.

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Head and body length (Fcal= 5.342, P= 0.982), ear length (Fcal= 10.731, P= 0.004), tragus

height (Fcal= 7.301, P= 0.015), length of thumb (Fcal= 5.047, P= 0.037), claw length (Fcal=

6.683, P= 0.019), length of the 3rd metacarpal (Fcal= 7.762, P= 0.012), 2nd phalanx on 3rd metacarpal (Fcal= 30.865, P= 0.000), 1st phalanx on 4th metacarpal (Fcal= 16.578, P= 0.001), 1st phalanx on 5th metacarpal (Fcal= 6.735, P= 0.018), wing span (Fcal= 13.148, P= 0.002) and hind

foot length (Fcal= 7.763, P= 0.012) varied significantly among the two populations (Table 35).

Cranial measurements. The mean breadth of braincase for five specimens of P. ceylonicus captured from the study area was recorded as 6.55 mm ± 1.604 (SD). The zgomatic breadth was

9.62 mm ± 4.012 (SD). The post-orbital constriction was 4.04 mm ± 0.776 (SD). The condylo- canine and condylo-basal lengths were 11.68 mm ± 3.826 (SD) and 12.54 mm ± 4.236 (SD), respectively. The greatest length of the skull was 10.76 mm ± 0.257 (SD). The maxillary toothrow length was 5.01 mm ± 2.484 (SD), anterior palatal width 4.20 mm ± 1.649 (SD), posterior palatal width 5.88 mm ± 2.039 (SD), mandibular toothrow and the mandible lengths were 4.64 mm ± 2.067 (SD) and 9.28 mm ± 3.956 (SD), respectively (Table 36).

Bacular measurements. Mean total length of the bacula of four P. ceylonicus was 3.66 mm ±

1.190 (SD), with the shaft length of 2.93 mm ± 1.125 (SD). The proximal and distal bacular lengths were 0.39 mm ± 0.2 06 (SD) and 0.28 mm ± 0.015 (SD), respectively while the proximal and distal widths were 0.75 mm ± 0.183 (SD) and 0.37 mm ± 0.035 (SD), respectively. The

bacular height was 0.60 mm ± 0.188 (SD) (Table 37).

5. Pipistrellus javanicus

Body mass and external body measurements. Mean body mass of all the ten P. javanicus was

7.30 g ± 2.889 (SD) while head and body length was 52.00 mm ± 2.712 (SD). The ear and tragus

lengths were 10.20 mm ± 3.369 (SD) and 4.44 mm ± 0.863 (SD), respectively. Thumb and claw

105

Table 35. Combined mean body mass (g) and external body measurements (mm) of Pipistrellus ceylonicus captured from Margalla Hills National Park (SA1) and Central Punjab (SA4).

SA1 SA4 Combined Statistics Body Parameters n= 15 n= 7 n= 22 F cal P Body mass 3.54±1.997 3.53±0.495 3.54±1.549 0.001 0.982 Head and body length 60.75±6.824 67.88±6.643 63.60±7.486 5.342 0.033* Ear length 7.22±1.345 9.31±1.487 8.06±1.724 10.731 0.004** Tragus length 4.99±0.640 4.25±0.535 4.70±0.694 7.301 0.015** Thumb length 4.14±0.568 3.50±0.707 3.89±0.689 5.047 0.037* Claw length 1.55±0.334 1.19±0.259 1.41±0.350 6.683 0.019** Forearm length 30.58±2.952 28.31±2.404 29.92±2.492 3.244 0.088 Length of 3rd metacarpal 27.25±3.028 23.75±2.252 25.85±3.205 7.762 0.012** 1st phalanx on 3rd metacarpal 12.54±1.296 11.63±3.238 12.18±2.246 0.790 0.386 2nd phalanx on 3rd metacarpal 10.17±2.060 4.50±0.926 11.90±2.741 30.865 0.000** Length of 4th metacarpal 26.68±3.657 23.69±2.314 25.49±3.462 4.201 0.055 1st phalanx on 4th metacarpal 11.83±1.105 9.88±0.954 11.05±1.416 16.578 0.001** 2nd phalanx on 4th metacarpal 9.17±1.303 8.50±1.190 8.92±1.272 1.230 0.283 Length of 5th metacarpal 26.53±3.254 24.00±2.828 25.52±3.271 3.215 0.090 1st phalanx on 5th metacarpal 8.50±1.462 10.13±1.217 9.15±1.565 6.735 0.018** Wing span 140.17±31.562 186.38±20.955 192.85±12.343 13.148 0.002** Tibia length 12.74±1.217 11.75±0.886 12.35±1.181 3.899 0.064 Calcar length 5.29±1.777 5.44±0.496 5.35±1.387 0.050 0.825 Hind foot length 6.86±1.242 5.56±0.496 6.34±1.187 7.763 0.012** Tail length 25.29±3.708 26.25±3.151 25.68±3.442 0.359 0.556 Penis length 5.78±0.857 5.25±0.500 5.57±0.753 0.038 0.847

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Table 36. Combined mean cranial measurements (mm) of Pipistrellus ceylonicus captured from Margalla Hills National Park (SA1) and some localities of Central Punjab (SA4) from June 2009 to May 2011 (n is the number of specimens). .

SA1 SA4 Combined Cranial Parameters n= 3 n= 2 n= 5 Breadth of braincase 7.12±1.973 5.69±0.216 6.55±1.604 Zygomatic breadth 11.46±4.293 6.86±1.437 9.62±4.012 Postorbital constriction 4.26±1.000 3.71±0.216 4.04±0.776 Condylo-canine length 12.70±5.026 10.15±0.449 10.08±0.433 Condylo-basal length 13.65±5.580 10.87±0.503 10.54±0.418 Greatest length of skull 10.76±0.301 10.77±0.287 10.76±0.257 Maxillary toothrow 6.01±2.933 3.52±0.054 5.01±2.484 Anterior palatal width 4.79±2.026 3.30±0.000 4.20±1.649 Posterior palatal width 6.69±2.422 4.67±0.180 5.88±2.039 Mandibular toothrow 5.30±2.631 3.66±0.036 4.64±2.067 Mandible length 10.18±5.314 7.92±0.216 9.28±3.956

107

Table 37. Combined mean bacular measurements (mm) of Pipistrellus ceylonicus captured from Margalla Hills National Park (SA1) and Central Punjab (SA4) from June 2009 to May 2011 (n is the number of specimens).

SA1 SA4 Combined Bacular Parameters n= 3 n= 1 n= 4 Total length of baculum 3.83±1.394 3.14 3.66±1.190 Length of shaft 3.10±1.310 2.40 2.93±1.125 Length of proximal branch 0.32±0.191 0.59 0.39±0.206 Length of distal branch 0.29±0.062 0.27 0.28±0.051 Width of proximal branch 0.74±0.222 0.78 0.75±0.183 Width of distal branch 0.38±0.014 0.32 0.37±0.035 Height of baculum 0.63±0.220 0.51 0.60±0.188

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lengths were 5.25 mm ± 0.655 (SD) and 1.81 mm ± 0.372 (SD), respectively. The forearm was

35.13 mm ± 1.996 (SD) long. The length of 3rd metacarpal and its 1st and 2nd phalanges were

31.38 mm ± 2.264 (SD), 12.33 mm ± 1.465 (SD) and 10.19 mm ± 1.280 (SD) long, respectively.

The length of 4th metacarpal was 31.06 mm ± 2.847 (SD) with 1st phalanx 11.56 mm ± 1.522

(SD) and 2nd phalanx 9.56 mm ± 2.718 (SD), respectively. The length of the 5th metacarpal and

its 1st phalanx were 30.31 mm ± 2.902 (SD) and 9.25 mm ± 0.926 (SD), respectively. The wing

span was 192.13 mm ± 48.129 (SD), tibia length 12.96 mm ± 1.247 (SD), calcar length 6.69 mm

± 1.308 (SD), hind foot length 7.89 mm ± 0.785 (SD), tail length 30.38 mm ± 5.236 (SD) while

the penis was 7.33 mm ± 1.756 (SD) long. Locality wise non-significant variations were

observed among all the parameters except claw length (Fcal= 10.417, P= 0.016) (Table 38).

Cranial measurements. Mean cranial measurements of five P. javanicus captured from SA2

and SA4 are given in Table 39. Breadth of the braincase and zygomatic breadth was 7.31 mm ±

1.541 (SD) and 10.23 mm ± 2.830 (SD), respectively. Post-orbital constriction was 4.58 mm ±

0.811 (SD). The condylo-canine and condylo-basal lengths were 12.08 mm ± 4.583 (SD) and

13.01 mm ± 4.546 (SD), respectively. The greatest skull length was 13.01 mm ± 4.546 (SD). The

maxillary toothrow length was 5.35 mm ± 1.411 (SD). Anterior and posterior palatal widths were

4.93 mm ± 1.057 (SD) and 6.67 mm ± 1.995 (SD), respectively. Mandibular toothrow and length

of the mandible was 4.86 mm ± 1.218 (SD) and 10.29 mm ± 1.679 (SD), respectively.

Bacular measurements. Mean bacular measurements of P. javanicus captured from three sub-

areas SA1, SA2 and SA4 are given in Table 40. The mean total length of the bacula was 3.57

mm ± 0.860 (SD) with a shaft length of 2.77 mm ± 0.833 (SD). The proximal and distal bacular

lengths were 0.47 mm ± 0.218 (SD) and 0.29 mm ± 0.023 (SD), respectively. The mean

109

Table 38. Combined mean body mass (g) and external body measurements (mm) of Pipistrellus javanicus captured from Margalla Hills National Park (SA1), Chinji National Park (SA2) and Central Punjab (SA4) from June 2009 to May 2011 (n is the number of specimens).

SA1 SA2 SA4 Combined Body Parameters n= 3 n= 6 n= 1 n= 10 Body mass 5.77±2.380 9.40±1.631 3.5 7.30±2.889 Head and body length 54.33±2.517 51.25±1.190 48.0 52.00±2.712 Ear length 11.03±4.786 10.13±2.983 8.0 10.20±3.369 Tragus length 4.83±0.289 4.13±1.181 4.5 4.44±0.863 Thumb length 4.83±0.577 5.50±0.707 5.5 5.25±0.655 Claw length 1.50±0.000 2.13±0.250 1.5 1.81±0.372 Forearm length 34.33±2.754 35.25±1.555 37.0 35.13±1.996 Length of 3rd metacarpal 31.17±3.403 31.75±1.936 30.5 31.38±2.264 1st phalanx on 3rd metacarpal 12.37±2.470 12.38±0.946 12.0 12.33±1.465 2nd phalanx on 3rd metacarpal 10.00±2.000 10.38±1.031 10.0 10.19±1.280 Length of 4th metacarpal 30.50±4.770 31.75±1.555 30.0 31.06±2.847 1st phalanx on 4th metacarpal 11.83±2.754 11.50±0.408 11.0 11.56±1.522 2nd phalanx on 4th metacarpal 8.50±3.279 10.38±2.839 9.5 9.56±2.718 Length of 5th metacarpal 29.17±4.646 31.00±1.780 31.0 30.31±2.902 1st phalanx on 5th metacarpal 9.17±1.607 9.25±0.500 9.5 9.25±0.926 Wing span 156.67±47.184 222.00±37.630 179.0 192.13±48.129 Tibia length 13.57±1.102 12.50±1.472 13.0 12.96±1.247 Calcar length 7.00±2.291 6.38±0.479 7.0 6.69±1.308 Hind foot length 7.53±0.503 8.13±1.031 8.0 7.89±0.785 Tail length 27.33±7.638 33.25±1.708 28.0 30.38±5.236 Penis length - 7.25±2.475 7.5 7.33±1.756

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Table 39. Mean cranial measurements (mm) of Pipistrellus javanicus captured from Chinji National Park (SA2) and Central Punjab (SA4) from June 2009 to May 2011 (n is the number of specimens).

SA2 SA4 Combined Cranial Parameters n= 4 n= 1 n= 5 Breadth of braincase 7.67±1.508 5.84 7.31±1.541 Zygomatic breadth 10.19±3.265 10.41 10.23±2.830 Postorbital constriction 4.82±0.696 3.61 4.58±0.811 Condylo-canine length 12.62±5.099 9.88 12.08±4.583 Condylo-basal length 13.64±4.991 10.49 13.01±4.546 Greatest length of skull 14.74±4.548 11.48 14.09±4.201 Maxillary toothrow 5.85±0.993 3.35 5.35±1.411 Anterior palatal width 5.23±0.933 3.71 4.93±1.057 Posterior palatal width 7.18±1.886 4.62 6.67±1.995 Mandibular toothrow 5.36±0.541 2.84 4.86±1.218 Mandible length 11.04±0.127 7.29 10.29±1.679

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Table 40. Combined mean bacular measurements (mm) of Pipistrellus javanicus captured from SA1, SA2 and central Punjab SA4 from June 2009 to May 2011 (n is the number of specimens).

SA1 SA2 SA4 Combined Bacular Parameters (n= 2) (n= 1) (n= 1) (n= 4) Total length of baculum 3.10±0.156 4.85 3.23 3.57±0.860 Length of shaft 2.35±0.139 4.02 2.38 2.77±0.833 Length of proximal branch 0.39±0.346 0.51 0.56 0.47±0.218 Length of distal branch 0.31±0.017 0.27 0.27 0.29±0.023 Width of proximal branch 0.67±0.017 0.78 0.78 0.73±0.064 Width of distal branch 0.40±0.017 0.44 0.42 0.42±0.020 Height of baculum 0.51±0.035 0.54 0.59 0.54±0.040

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proximal and distal bacular widths were 0.73 mm ± 0.064 (SD) and 0.42 mm ± 0.020 (SD),

respectively. The baculum height was 0.54 mm ± 0.040 (SD).

6. Pipistrellus pipistrellus

Body mass and external body measurements. Pipistrellus pipistrellus was captured from all

the four sub-areas. The mean combined body mass and external body measurements of fifty two

P. pipistrellus captured from all the four sub-areas is given in Table 41. The combined mean

body mass of all the specimens captured from the study area was 3.78 g ± 0.530 (SD) and head

and length was 39.33 mm ± 2.690 (SD). The ear and tragus lengths were 8.81 mm ± 3.593 (SD)

and 4.28 mm ± 0.623 (SD), respectively. The thumb and claw lengths were 3.73 mm ± 0.656

(SD) and 1.39 mm ± 0.358 (SD), respectively. The forearm was 28.23 mm ± 1.264 (SD) long.

The length of the 3rd metacarpal was 25.27 mm ± 1.203 (SD) with its 1st phalanx 11.26 mm ±

0.855 (SD) and 2nd phalanx 8.90 mm ± 1.268 (SD). The 4th metacarpal and its 1st and 2nd phalanges were 24.81 mm ± 1.504 (SD), 10.53 mm ± 0.785 (SD) and 7.68 mm ± 0.927 (SD), respectively. The length of 5th metacarpal and its 1st phalanx was 24.63 mm ± 1.284 (SD) and

8.07 mm ± 0.654 (SD), respectively. The mean wing span, tibia length, calcar length, hind foot length, tail and penis lengths were 187.15 mm ± 28.431 (SD), 11.40 mm ± 0.935 (SD), 5.10 mm

± 1.261 (SD), 6.31 mm ± 0.863 (SD), 25.86 mm ± 3.396 (SD) and 6.40 mm ± 1.300 (SD),

respectively (Table 41).

Cranial measurements. Mean cranial measurements of P. pipistrellus from all the four sub-

areas are given in Table 42. The mean breadth of the braincase was 6.01 mm ± 0.428 (SD) while

the zygomatic breadth was 8.56 mm ± 2.501 (SD). The post-orbital constriction was 3.55 mm ±

0.196 (SD). The condylo-canine and condylo-basal lengths were 10.02 mm ± 0.583 (SD) and

10.88 mm ± 0.327 (SD), respectively. The greatest length of skull was 11.04 mm ± 0.342 (SD).

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Table 41. Combined mean body mass (g) and external body measurements (mm) of Pipistrellus pipistrellus captured from Margalla Hills National Park (SA1), Lal Suhanara National Park (SA3) and Central Punjab (SA4) from June 2009 to May 2011 (n is the number of specimens).

SA1 SA3 SA4 Combined Statistics Body Parameters (n=19) (n=12) (n=20) (n= 52) F cal P Body mass 3.82±0.549 4.30±0.200 3.70±0.542 3.78±0.530 1.171 0.339 Head and body length 37.50±13.635 42.67±2.309 39.14±3.988 39.33±2.690 1.205 0.245 Ear length 7.83±1.169 8.67±0.577 9.10±4.315 8.81±3.593 0.178 0.910 Tragus length 4.25±0.612 4.00±0.000 4.35±0.684 4.28±0.623 0.332 0.802 Thumb length 2.92±0.204 3.33±0.289 4.05±0.522 3.73±0.656 10.953 0.000** Claw length 1.08±0.376 1.50±0.000 1.45±0.350 1.39±0.358 1.976 0.141 Forearm length 28.58±1.497 29.00±0.000 27.98±1.270 28.23±1.264 0.927 0.441 Length of 3rd metacarpal 25.25±1.173 24.83±0.577 25.31±1.318 25.27±1.203 0.243 0.866 1st phalanx on 3rd metacarpal 11.42±0.917 11.17±0.764 11.19±0.887 11.26±0.855 0.351 0.789 2nd phalanx on 3rd metacarpal 8.83±0.516 8.67±0.289 8.93±1.519 8.90±1.268 0.107 0.955 Length of 4th metacarpal 24.75±1.173 24.33±0.289 24.83±1.713 24.81±1.504 0.292 0.831 1st phalanx on 4th metacarpal 10.33±0.753 10.50±0.000 10.60±0.875 10.53±0.785 0.161 0.922 2nd phalanx on 4th metacarpal 7.50±0.837 7.50±1.803 7.71±0.860 7.68±0.927 0.359 0.783 Length of 5th metacarpal 24.42±1.357 24.00±1.323 24.69±1.260 24.63±1.284 1.020 0.399 1st phalanx on 5th metacarpal 8.63±0.816 7.83±0.289 7.93±0.576 8.07±0.654 2.375 0.092 Wing span 162.00±26.069 191.00±3.606 195.79±26.118 187.15±28.431 3.787 0.022** Tibia length 11.33±1.033 11.33±0.577 11.40±0.995 11.40±0.935 0.140 0.935 Calcar length 4.50±0.548 5.33±1.528 5.31±1.337 5.10±1.261 1.247 0.312 Hind foot length 6.17±0.983 7.00±1.000 6.19±0.782 6.31±0.863 1.542 0.226 Tail length 25.17±4.916 23.00±1.732 26.36±3.016 25.86±3.396 1.087 0.372 Penis length 6.00±1.000 6.50±0.707 6.53±1.457 6.40±1.300 0.435 0.730

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Table 42. Combined mean cranial measurements (mm) of Pipistrellus pipistrellus captured from Margalla Hills National Park (SA1), Chinji National Park (SA2), Lal Suhanara National Park (SA3) and Central Punjab (SA4) from June 2009 to May 2011 (n is the number of specimens). .

Cranial Parameters SA1 SA2 SA3 SA4 Combined (n= 6) (n= 1) (n= 3) (n= 3) (n= 13) Breadth of braincase 6.05±0.311 6.88 5.62±0.195 5.80±0.237 6.01±0.428 Zygomatic breadth 9.47±2.986 8.45 7.79±2.494 7.47±0.847 8.56±2.501 Postorbital constriction 3.57±0.143 3.46 3.48±0.353 3.44±0.373 3.55±0.196 Condylo-canine length 10.18±0.292 10.40 10.02±0.509 9.65±1.037 10.02±0.583 Condylo-basal length 10.99±0.195 10.90 10.59±0.391 10.85±0.482 10.88±0.327 Greatest length of skull 11.02±0.346 11.06 10.95±0.432 11.42±0.475 11.04±0.342 Maxillary toothrow 3.59±0.221 3.42 3.46±0.103 3.46±0.444 3.56±0.197 Anterior palatal width 3.59±0.447 3.09 3.41±0.552 3.53±0.380 3.46±0.421 Posterior palatal width 4.89±0.262 4.42 4.81±0.301 4.73±0.068 4.80±0.258 Mandibular toothrow 3.51±0.385 3.59 3.21±0.053 3.70±0.745 3.43±0.393 Mandible length 8.18±0.874 7.75 7.14±0.659 8.00±0.267 7.87±0.802

115

The maxillary toothrow length, the anterior and posterior palatal widths were 3.56 mm ± 0.197

(SD), 3.46 mm ± 0.421 (SD) and 4.80 mm ± 0.258 (SD), respectively while the mandibular

toothrow length and the length of mandible was 3.43 mm ± 0.393 (SD) and 7.87 mm ± 0.802

(SD), respectively.

Bacular measurements. The mean total length of the eleven bacula of P. pistrellus was 3.19

mm ± 0.421 (SD) while the shaft of the bacula was 2.35 mm ± 0.245 mm long. The proximal and

distal bacular lengths were 0.59 mm ± 0.218 (SD) and 0.26 ± 0.075 (SD), respectively. The

proximal and distal bacular widths were 0.76 mm ± 0.055 (SD) and 0.38 mm ± 0.059 (SD). The

height of the baculum was 0.60 mm ± 0.156 mm (Table 43).

7. Pipistrellus tenuis

Body mass and external body measurements. Only two specimens of P. tenuis were captured

from SA1. The mean body mass was 4.25 g ± 1.061 (SD), while head and body length was 35.00

mm ± 2.828 (SD). The ear and tragus lengths were 7.50 mm ± 0.707 (SD) and 3.00 mm ± 0.000

(SD), respectively. The thumb and claw lengths were 3.50 mm ± 0.707 (SD) and 1.75 mm ± 0.354 (SD), respectively. The 3rd metacarpal and its 1st and 2nd phalanges were 25.25 mm ± 0.354 (SD), 10.75

mm ± 0.354 (SD) and 8.25 mm ± 1.061 (SD), respectively. The 4th metacarpal and its 1st and 2nd phalanges were 23.75 mm ± 1.061 (SD), 10.75 mm ± 0.354 (SD) and 8.00 mm ± 0.000 (SD) long, respectively. The length of the 5th metacarpal and its 1st phalanx was 23.75 mm ± 1.061

(SD) and 6.75 mm ± 0.354 (SD), respectively. The wing span, tibia, calcar, hind foot, tail and

penis lengths were 156.00 mm ± 21.213 (SD), 10.25 mm ± 1.768 (SD), 6.00 mm ± 1.414 (SD),

6.25 mm ± 1.768 (SD), 22.25 mm ± 3.182 (SD) and 4.50 mm ± 0.707 (SD), respectively (Table

44).

Cranial measurements. Various cranial measurements of a single P. tenuis captured from SA 1

are given in Table 45. The breadth of the braincase and zygomatic breadths were 6.30 mm and

116

Table 43. Combined mean bacular measurements (mm) of Pipistrellus pipistrellus captured from Margalla Hills National Park (SA1), Chinji National Park (SA2) and Central Punjab (SA4) from June 2009 to May 2011 (n is the number of specimens).

SA1 SA2 SA4 Combined Bacular Parameters (n= 2) (n= 1) (n= 8) (n= 11) Total length of baculum 3.26±0.035 0.345 3.15±0.489 3.19±0.421 Length of shaft 2.32±0.191 0.270 2.31±0.248 2.35±0.245 Length of proximal branch 0.65±0.191 0.049 0.58±0.246 0.59±0.218 Length of distal branch 0.27±0.069 0.029 0.25±0.085 0.26±0.075 Width of proximal branch 0.78±0.000 0.081 0.74±0.058 0.76±0.055 Width of distal branch 0.33±0.017 0.034 0.39±0.063 0.38±0.059 Height of baculum 0.64±0.208 0.078 0.57±0.150 0.60±0.156

117

Table 44. Mean body mass (g) and external body measurements (mm) of Pipistrellus tenuis captured from SA1 from June 2009 to May 2011 (n is the number of specimens).

NARC Rawal Town Combined Body Parameters Mean±SD n= 1 n= 1 (n= 2) Body mass 5.00 3.500 4.25±1.061 Head and body length 33.00 37.00 35.00±2.828 Ear length 7.00 8.00 7.50±0.707 Tragus length 3.00 3.00 3.00±0.000 Thumb length 4.00 3.00 3.50±0.707 Claw length 2.00 1.50 1.75±0.354 Forearm length 27.50 28.50 28.00±0.707 Length of 3rd metacarpal 25.50 25.00 25.25±0.354 1st phalanx on 3rd metacarpal 11.00 10.50 10.75±0.354 2nd phalanx on 3rd metacarpal 9.00 7.50 8.25±1.061 Length of 4th metacarpal 23.00 24.50 23.75±1.061 1st phalanx on 4th metacarpal 10.50 11.00 10.75±0.354 2nd phalanx on 4th metacarpal 8.00 8.00 8.00±0.000 Length of 5th metacarpal 23.00 24.50 23.75±1.061 1st phalanx on 5th metacarpal 7.00 6.50 6.75±0.354 Wing span 171.00 141.00 156.00±21.213 Tibia length 11.50 9.00 10.25±1.768 Calcar length 5.00 7.00 6.00±1.414 Hind foot length 7.50 5.00 6.25±1.768 Tail length 24.50 20.00 22.25±3.182 Penis length 5.00 4.00 4.50±0.707

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Table 45. Cranial measurements (mm) of Pipistrellus tenuis captured from Margalla Hills National Park (SA1) from June 2009 to May 2011 (n is the number of specimens).

Cranial Parameters n= 1 Breadth of braincase 6.30 Zygomatic breadth 7.80 Postorbital constriction 3.96 Condylo-canine length 9.42 Condylo-basal length 10.13 Greatest length of skull 10.19 Maxillary toothrow 3.78 Anterior palatal width 3.56 Posterior palatal width 5.23 Mandibular toothrow 4.27 Mandible length 7.82

119

7.80 mm, respectively. The post-orbital constriction was 3.96 mm. The condylo-canine and

condylo-basal lengths were 9.42 mm and 10.13 mm, respectively. The greatest length of the skull

was 10.19 mm. The maxillary toothrow length was 3.78 mm, anterior and posterior palatal

widths were 3.56 mm and 5.23 mm, respectively. Mandibular toothrow length and the length of

the mandible were 4.27 mm and 7.82 mm, respectively.

Bacular measurements. The total bacular length was 2.79 mm with shaft length of 2.08 mm.

The proximal and distal bacular lengths were 0.39 mm and 0.27 mm, respectively while

proximal and distal bacular widths were 0.61 mm and 0.34 mm, respectively. The height of the

baculum was 0.51 mm (Table 46).

8. Hypsugo savii

Body mass and external body measurements. Mean body mass and external body

measurements of two H. savii captured from SA1 and SA2 are given in Table 47. The mean

body was 6.90 g ± 4.243 (SD). The head and body length was 55.50 mm ± 19.092 (SD). The ear

length and the length of tragus was 9.50 mm ± 0.707 (SD) and 5.50 mm ± 0.707 (SD), respectively. The

lengths of the thumb and claw were 5.00 mm ± 1.414 (SD) and 2.00 mm ± 0.000 (SD), respectively.

The forearm was 36.75 mm ± 3.889 (SD). The length of 3rd metacarpal and its 1st and 2nd phalanges was 42.50 mm ± 12.728 (SD), 13.50 mm ± 2.121 (SD) and 10.90 mm ± 1.556 (SD), respectively. The length of the 4th metacarpal was 33.00 mm ± 2.828 (SD) with its 1st phalanx

13.00 mm ± 0.000 (SD) and 2nd phalanx 9.25 mm ± 0.354 (SD). The 5th metacarpal and the 1st phalanx on 5th metacarpal was 31.95 mm ± 2.758 (SD) and 10.25 mm ± 1.061 (SD), respectively.

Mean wing span, tibia, calcar, hind foot and tail lengths were 167.00 mm ± 32.527 (SD), 12.75

mm ± 1.061 (SD), 6.00 mm ± 1.414 (SD), 8.50 mm ± 0.707 (SD) and 33.50 mm ±6.364 (SD),

respectively.

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Table 46. Bacular measurements (mm) of Pipistrellus tenuis captured from Margalla Hills National Park (SA1) from June 2009 to May 2011 (n is the number of specimens).

Bacular Parameters n= 1 Total length of baculum 2.79 Length of shaft 2.08 Length of proximal branch 0.39 Length of distal branch 0.27 Width of proximal branch 0.61 Width of distal branch 0.34 Height of baculum 0.51

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Table 47. Mean body mass (g) and external body measurements (mm) of Hypsugo savii captured from Margalla Hills National Park (SA1) and Chinji National Park (SA2) from June 2009 to May 2011 (n is the number of specimens).

SA1 SA2 Combined Body Parameters n= 1 n= 1 n= 2 Body mass 3.9 9.9 6.90±4.243 Head and body length 69.0 42 55.50±19.092 Ear length 9.0 10 9.50±0.707 Tragus length 5.0 6 5.50±0.707 Thumb length 4.0 6 5.00±1.414 Claw length 2.0 2 2.00±0.000 Forearm length 39.5 34 36.75±3.889 Length of 3rd metacarpal 33.5 51.5 42.50±12.728 1st phalanx on 3rd metacarpal 12.0 15 13.50±2.121 2nd phalanx on 3rd metacarpal 9.8 12 10.90±1.556 Length of 4th metacarpal 35.0 31 33.00±2.828 1st phalanx on 4th metacarpal 13.0 13 13.00±0.000 2nd phalanx on 4th metacarpal 9.0 9.5 9.25±0.354 Length of 5th metacarpal 33.9 30 31.95±2.758 1st phalanx on 5th metacarpal 9.5 11 10.25±1.061 Wing span 144.0 190 167.00±32.527 Tibia length 12.0 13.5 12.75±1.061 Calcar length 5.0 7 6.00±1.414 Hind foot length 9.0 8 8.50±0.707 Tail length 29.0 38 33.50±6.364 Penis length 7.00 - 7.00±0.00

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Cranial measurements. The breadth of the braincase of this specimen was 7.14 mm and the zygomatic breadth was 7.44 mm. The post-orbital constriction was 4.52 mm. Condylo-canine

and condylo-basal lengths were 10.41 mm and 11.10 mm, respectively. The greatest length of the

skull was 11.18 mm, maxillary toothrow length 5.41 mm, anterior and posterior palatal widths

were 4.88 mm and 6.07 mm, respectively. The mandibular toothrow and the length of mandible

were 5.38 mm and 7.08 mm, respectively (Table 48).

Bacular measurements. The total bacular length of a single H. savii captured from SA1 was

2.67 mm with shaft length of 2.08 mm. The proximal and distal bacular lengths were 0.39 mm

and 0.27 mm, respectively while proximal and distal bacular widths were 0.61 mm and 0.34 mm,

respectively. The height of the baculum was 0.51 mm (Table 49).

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Table 48. Cranial measurements (mm) of Hypsugo savii captured from Margalla Hills National Park (SA1) from June 2009 to May 2011 (n is the number of specimens).

Cranial Parameters n= 1 Breadth of braincase 7.14 Zygomatic breadth 7.44 Postorbital constriction 4.52 Condylo-canine length 10.41 Condylo-basal length 11.10 Greatest length of skull 11.18 Maxillary toothrow 5.41 Anterior palatal width 4.88 Posterior palatal width 6.07 Mandibular toothrow 5.38 Mandible length 7.08

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Table 49. Bacular measurements (mm) of Hypsugo savii captured from SA1from June 2009 to May 2011 (n is the number of specimens).

Bacular Parameters n= 1 Total length of baculum 2.67 Length of shaft 2.08 Length of proximal branch 0.39 Length of distal branch 0.27 Width of proximal branch 0.61 Width of distal branch 0.34 Height of baculum 0.51

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(a) (b) (c) (d) (e)

Figure 19. Dorsal view of the bacula of Pipistrellus ceylonicus (a), P. javanicus (b), P.

pipistrellus (c), P. tenuis (d) Hypsugo savii (e).

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PART IV. ECHOLOCATION CALLS

Bat calls have not been recorded, analyzed and interpreted in Pakistan prior to this study.

Since there has been no reference bat call library, most of the bat calls recorded during this study remained unidentified. All sound files including those that could not be identified are thus copied on the accompanying CD for further analysis.

A total of 100 bat calls were recorded from different netting stations. Most of the calls were recorded when bats were flying except a few emballonurid calls that were recorded at their day time roosts located at Derawar Fort and Mojgarh in Lal Suhanra National Park (SA3). Since both the Egyptian tomb bat (T. perforatus) and naked-rumped tomb bat (T. nudiventris) were co- roosting, the social calls emitted by these species could not be differentiated (Table 50). Social calls in both these species are long multi-harmonic and comprise two to five harmonics. Mean call duration is 19 ms ± 6.4 SD (range = 6 – 25 ms; n = 13). Mean frequency of maximum energy is 23.5 kHz ± 3.1 SD (range 15.4 – 29.8; n = 13). The mean start and end frequency are

36.2 kHz ± 8.9 SD (range 23-53 kHz; n =13) and 15.5 kHz ± 5.3 SD (range 6-24; n = 13). The sonograms of the flight calls of these two species show that both Taphozous spp. emit long multi-harmonic calls with three to four harmonics (Figure 20).

Echolocation calls of three vesper bat species were also identified. These included

Scotophilus heathii, S. kuhlii and Pipistrellus tenuis (Table 51, Figure 21, 22). Three calls of S. heathii (M00028, M00031and M00079) were analyzed. Mean and standard deviations for various call parameters for two pulses of M00028, five of M00031 and seven of M00079 are given in Table 57a. Mean duration for these frequency modulated calls was 7.1 ms ± 1.5 (SD).

Mean Frequency of maximum energy was 40.9 kHz ± 8.0 (SD). Mean start and end call

frequencies were 62.1 kHz

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Table 50. Social calls emitted by T. nudiventris and T. perforatus recorded at Derawar Fort and Mojgarh in Lal Suhanra National Park (SA3).

Call no Duration Fmaxe Start freq End freq Highest Lowest Range No harmonics Call type (ms) (kHz) (kHz) (kHz) (kHz) (kHz) (kHz) (n) m00051 23 15.4 34 13 34 13 21 2 l mh m00052 14 23.2 38 6 59 6 32 5 l mh m00053 27 22.2 44 20 44 20 24 3,4 l mh m00053 14 22.4 27 6 61 6 21 5 l mh m00054 18 24 53 16 81 16 37 5 l mh m00056 20 24.6 42 16 56 16 26 3 l mh m00057 6 29.8 46 24 77 24 22 4 l mh m00058 13 22.8 23 15 57 5 8 5 l mh m00059 18 23.2 37 16 46 6 21 4 l mh m00060 29 25.7 30 23 66 23 7 4 l mh m00061 22 23.6 41 14 78 14 27 5 l mh m00062 18 24.6 28 16 56 15 12 4 l mh m00063 25 24.2 27 16 54 16 11 4 l mh *ms= milli seconds; kHz= kilo hertz; lmh= long multi harmonic

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Figure 20. The sonograms of the flight calls of Taphozous spp.

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Table 51. Call parameters of some vespertilionid bats recorded from some arid subtropical and tropical regions of Pakistan. (a) Scotophilus heathii Call No. Duration FmaxE Start Freq End Freq IPI (ms) (kHz) (kHz) (kHz) (ms) M00028 (n = 2) Mean 5.3 54.9 70.0 53.5 63.7 SD 0.5 0.5 9.9 0.7 4.0 M00031( n= 5) Mean 7.1 32.2 33.0 29.0 115.4 SD 0.5 0.1 0.7 0.0 22.0 M00079 (n= 7) Mean 7.6 43.0 80.7 39.4 71.8 SD 1.8 2.3 16.4 2.0 25.3 Combined (n =14) Mean 7.1 40.9 62.1 37.7 86.2 SD 1.5 8.0 25.6 8.4 31.1 (b) Scotophilus kuhlii M00009 (n = 4) Mean 3.3 54.8 111.3 50.0 72.1 SD 0.1 0.3 5.6 1.6 0.6 M00010( n= 3) Mean 2.8 56.8 106.3 51.7 75.7 SD 0.2 0.6 2.5 0.6 4.8 M00011 (n= 7) Mean 2.5 60.1 114.9 50.4 52.7 SD 0.4 5.1 2.9 0.6 14.9 M00014 (n= 6) Mean 2.4 54.5 89.0 50.5 68.5 SD 0.3 2.2 12.2 1.2 3.1 M00017 (n= 5) Mean 2.0 47.9 83.2 42.5 50.0 SD 0.1 22.4 35.0 20.3 26.1 Combined (n = 25) Mean 2.5 56.9 103.5 50.6 63.7 SD 0.5 3.6 12.3 1.0 12.9 (c) Pipistrellus tenuis M00001 (n= 2) Mean 9.35 33.65 57.45 33.9 163.3 SD 0.6 1.5 3.0 3.8 19.4 M00002 (n= 2) Mean 10.4 34.75 54.5 32.3 78.85 SD 1.1 1.8 7.2 1.6 93.0 Combined (n = 4) Mean 9.9 34.2 56.0 33.1 121.1 SD 1.0 1.5 4.8 2.6 73.4 *(ms= milli seconds; kHz= kilo hertz; IPI= inter pulse interval)

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Amplitude Amplitude 100% 100%

50% 50%

0% 0%

-50% -50%

100 ms 300 ms -100% -100% 20 ms/div 20 ms/div -90 dB -70 dB -50 dB -30 dB -10 dB -90 dB -70 dB -50 dB -30 dB -10 dB Spectrogram, FFT size 512 , Hanning window. Spectrogram, FFT size 512 , Hanning window.

200 kHz 200 kHz

100 kHz 100 kHz

100% Amplitude 100% Amplitude

50% 50%

0% 0%

-50% -50%

460 ms 380 ms -100% -100% 20 ms/div 20 ms/div -90 dB -70 dB -50 dB -30 dB -10 dB -90 dB -70 dB -50 dB -30 dB -10 dB Spectrogram, FFT size 512 , Hanning window. Spectrogram, FFT size 512 , Hanning window.

200 kHz 200 kHz

100 kHz 100 kHz

Figure 21. Representative sonograms of Scotophilus heathii.

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Amplitude 100%

50%

-50%

2187 ms -100% 50 ms/div -90 dB -70 dB -50 dB -30 dB -10 dB Spectrogram, FFT size 512 , Hanning window.

150 kHz

100 kHz

50 kHz

Amplitude 100%

50%

-50%

2387 ms -100% 50 ms/div -90 dB -70 dB -50 dB -30 dB -10 dB Spectrogram, FFT size 512 , Hanning window.

150 kHz

100 kHz

50 kHz

Figure 22. Representative sonograms of Pipistrellus spp.

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± 25.6 (SD) and 37.7 kHz ± 8.4 (SD), respectively where mean inter pulse interval was 86.2 ms

± 31.1 (SD). The calls varied from single harmonic to multi-harmonic.

Twenty five pulses from five calls of S. kuhlii were analyzed (Table 51b). The bats emitted narrow band frequency modulated signals. Mean duration for these calls was 2.5 ms ±

0.5 (SD). Mean Frequency of maximum energy was 56.9 kHz ± 3.6 (SD). Mean start and end call frequencies were 103.5 kHz ± 12.3 (SD) and 50.6 kHz ± 1.0 (SD), respectively where mean inter pulse interval was 63.7 ms ± 12.9 (SD). The calls were single harmonic.

Four pulses from two calls of Pipistrellus tenuis were analyzed (Table 51c). The bats emitted broad band frequency modulated signals. Mean duration for these calls was 9.9 ms ± 1.0

(SD). Mean Frequency of maximum energy was 34.1 kHz ± 1.5 (SD). Mean start and end call frequencies were 56.0 kHz ± 4.8 (SD) and 33.1 kHz ± 2.6 (SD), respectively where mean inter pulse interval was 121.3 ms ± 73.4 (SD). The calls were single harmonic.

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CHAPTER V

DISCUSSION

GENERAL DISCUSSION

Pakistan is represented by ten out of the twenty six mammalian orders (Roberts, 1997)

indicating a diversity comparable to any region of the world with similar climatic conditions

(Mahmood-ul-Hassan et al., 2009). The rate of endemism is however low with only four

endemic mammal species including 13 sub species (CBD, 2009). Small mammals constitute 63%

of the 195 mammal species of the country. They are the most diverse yet the least studied group

of chordates (Roberts, 1997). Since they do not belong to flagship species of the country, they

are neither included on the agenda list of conservation agencies nor in the educational policy of

the country. They are often disliked by the people as dirty creatures responsible for transmitting

diseases to humans and livestock (Mahmood-ul-Hassan et al., 2011). Misbelieves among the

masses and resulting negative public perceptions have been the major cause for bringing many

bat species to the verge of extinction in Pakistan (Sheikh and Molur, 2004).

Bats constitute 28% of the mammal fauna of the country (Roberts, 1997) representing

Indo-Malayan, Palaearctic and Ethiopean species. The Palaearctic fauna is largely found in the

Himalayan and Baluchistan mountainous regions. Many vespertilionid bats e.g. the mountain

nocule Nyctalus montanus, the eastern barbastelle Barbastella leucomelas have probably reached

here from Iran through southern Baluchistan or from the Hindu Kush Mountains and Russian

Uzebeskistan. The Palaearctic bat species that are adapted to more arid conditions and warmer

climate like the Blasius’s horse-shoe bat Rhinolophus blasii, the greater horse-shoe bat

Rhinolophus ferrumequinum and the trident leaf-nosed bat Asellia tridens have invaded the country through southern Baluchistan (Roberts, 1997). Some of these e.g. R. blasii have

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extended their ranges over the past few decades and have been recorded from Lahore district

(Roberts, 1997, present study) which lies at the edge of Indo-Malayan region (Darlington, 1957).

Most of the Indo-Malayan bat fauna has probably failed to cross the Indian desert but those which succeeded to colonize Pakistan have invaded the country either from the south through the

savannah type vegetation of the Rann of Katch or from the north through the Himalayan

foothills. The bi-coloured leaf-nosed bat Hippocideros fulvus and the Indian false vampire bat

Megaderma lyra, the Indian flying fox Pteropus giganteus and the Short-nosed fruit bat

Cynopterus sphinx are examples of Indo-Malayan bat species that colonized via this route and still exist very successfully in Pakistan (Roberts, 1997). The Indian flying fox has greatly

extended its range over the time and colonies of these bats can be located in Peshawar,

Malakand, Dir and Dargai districts (Salim, 2009; Faiz-ur-Rehman, 2010) which are a part of

Palaearctic region of the world (Darlington, 1957). The Ethiopian bat fauna most likely invaded

this region from the west through the Makran coastal belt. The Egyptian fruit bat Rousettus

aegyptiacus, wrinkle-lipped bat Tadarida aegyptiaca and the Egyptian tomb bat Taphozous

perforatus are the example of such bat species (Roberts, 1997). Of these the Egyptian Tomb T.

perforatus bat has greatly extended its range over the past few decades and was captured in the

Pothowar region (present study).

The distribution ranges of almost all the mammals have changed but these changes especially in case of small mammals have not been studied in Pakistan. Increased human populations and resultant habitat changes are two principal factors that have reshaped distribution pattern of almost all the mammals. About a century ago before the introduction of irrigation canal system, four types of ecologically unrelated habitats existed in the study area.

These included the (a) flood plains of river Ravi and Chenab, (b) tropical thorn forest, (c) sand

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dune areas and (d) villages (Taber et al., 1967). These habitats supported a variety of macro- and

micro-fauna. With the increase in the human population and rapid colonization following an

intensive canal irrigation system, a lot of habitat changes have taken place. Remnants of tropical

thorn forest or its secondary growth forms largely occur in small patches only. Sand dunes are

rare except near the Thal and Cholistan desert. The hydric habitat has extended in the

meanwhile. A sizeable portion of the irrigated land of the country is affected by alkalinity,

salinity and water logging (Ahmad, 1999). Canal irrigation has benefited those bat species that are favoured by the spread of irrigation and cultivation. In addition, irrigated forest plantations, canal-side, rail road-side and road-side plantations have greatly changed the landscape in which the thorn forest previously predominated (Taber et al., 1967).

Another important habitat of the rural landscape of the study area is the human settlement comprising villages and farmhouses, which are increasing both in number and size in the wake of the fast-expanding human population. Inter-village distances have shortened and it has become easier for certain bat species (e.g. the greater yellow house bat Scotophilus heathii) to populate the entire study area (present study). Ellerman and Morrison-Scot (1951) recorded this species from Sindh but not from Punjab. It was also not included in Siddiqi’s (1961) checklist and has extended its range from east into the study area with irrigation and construction of houses which serve as its roost (present study).

BAT SURVEY AND ABUNDANCE

Habitat loss and degradation are the key factors frightening bat populations globally.

About one fourth of the bat species are considered threatened (Schipper et al., 2008). There is a

need to monitor and conserve bat populations at global scale (Jones et al., 2009). Although some

monitoring programs have been initiated but they are mostly focusing temperate regions and are

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limited in scope, primarily assessing single-species locally (Betke et al., 2008; Hristov et al.,

2010; O’Shea and Bogan, 2003; Walsh et al., 2003). This may be the fact that the bats inhabiting

tropica areas are threatened by deforestation and loss and fragmentation of habitat at an

unprecedented scale (IUCN, 2009; Lane et al., 2006). As the bat diversity is more concentrated

towards tropics and provide many ecological services in respective ecosystems (Jones et al.,

2009), there is a dire need to monitor bat populations inhabiting tropicl areas of the world.

The greatest bat diversity exists in tropical and sub-tropical regions of the world where bats remain active almost throughout the year (Audet and Fenton, 1988; Hickey and Fenton,

1996; Turbill et al., 2001; Chruszcz and Barclay, 2002). Temperature is especially crucial for

small-sized bats that need high energy for their maintenance, thermoregulation and locomotion.

While energy required for thermoregulation increases steeply at temperatures below 30°C

(Lyman, 1970; Geiser, 2003; Speakman and Thomas, 2003; Geiser and Kortner, 2010), the

thermal neutral zone (TNZ) of these (10 g) bats is restricted to the ambient temperatures of around or greater than 30°C (Hock, 1951; Kulzer et al., 1970; Geiser and Brigham, 2000). It is due to this reason that most insectivorous echolocating bats inhabiting temperate-zone enter extended bouts of torpor and hibernate throughout winter (e.g. Park et al., 2000). Furthermore, the insect prey availability is much reduced at low ambient temperatures (Hickey and Fenton,

1996). Thus bats are relatively more abundant and diverse in vegetated terrestrial ecosystems of the world including tropical and temperate habitats (Fenton, 1992; Emmons and Feer, 1997; Hill and Smith, 1984; Vaughan et al., 2000). The present study was also conducted in tropical and arid subtropical regions of Pakistan to document bat fauna of the country.

Bates and Harrison (1997) have made the most comprehensive and up-to-date revision of the Chiroptera of the Indian subcontinent and enlisted 119 species of bats belonging to eight

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families and 37 genera from India, Pakistan, Nepal, Sri Lanka, Maldives, Afghanistan, Tibet, and

Northern Myanmar. Out of these 119 species, 110 are recorded within the Indian limits (Bates

and Harrison, 1997). As far as their status in the region is concerned, one bat species has gone

Extinct, 43 are Threatened, six are Critically Endangered, nine are Endangered and 28 are

Vulnerable. In India, two species of bats viz., the Wroughton’s free-tailed bat (Otomops

wroughtonii) and Salim Ali’s fruit bat (Latidens salimalii) are highly protected and are on

schedule 1 of wildlife (protection) act 1972. Furthermore, persistent efforts by of globally

renowned bat biologists and non-governmental organizations in India have resulted in providing

legal protection to all 13 species of pteropodid bats (Singaravelan et al., 2009; Mahmood-ul-

Hassan et al., 2010). Bats are given no protection by law in Pakistan and many species are

hunted for their body fat to be used by local health practitioners to cure rheumatic pains

(Roberts, 1997).

Our information regarding bat fauna of Pakistan is fragmentary both in terms of number

of species and their distribution (Mahmood-ul-Hassan et al., 2009). Status of South Asian

Chiroptera CAMP Report (Walker and Molur, 2003) has been mostly referred for information and status on Chiroptera (Bats) for Pakistan’s Mammals. There has been no extensive survey to find out the exact number of bats species in the country so far. Roberts (1997) has listed 50 species of bats representing 23 genera and 8 families from Pakistan whereas Walker and Molur

(2003) expect the presence of some additional species from this country. Similarly the taxonomy of many bat species such as those belonging to genus Pipistrellus and Hypsugo, Kerivoula and

Plecotus need validation (Mahmood-ul-Hassan et al., 2009). As for as the survival of bats is concerned, one species of bat is Near Extinction (Triaenops persicus), one species is Endangered

(Nyctalus leisleri), seven are Near Threatened (Rhinolophus blasii, R. ferrumequinum, R.

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Lepidus, Scotoecus pallidus, Scotozous dormer, Barbestella leucomeaals, Otonycterus and

hemprichii) six are Vulnerable (Rousettus aegyptiaucs, Rhinolophus hipposideros, Rhinopoma

musctellum, Eptesicus nasutus, Myotis emarginatus and M. longipes) and three are data deficient

(Eptesicus bottae, E. gobiensis and E. serotinus) (IUCN, 2008). The status of almost 18 bat

species is unknown and is described by their single specimens present either in British Museum

(UK), Harrison Museum (USA) and Punjab University Museum of Zoology (Pakistan) (Roberts,

1997; Mahmood-ul-Hassan et al., 2009).

The original literature on the mammals of this region is of four main kinds: 1) general

accounts of "Indian" mammals, such as those of Jerdon (1867), Murray (1884), Blanford (1888-

1891), Lydekker (1924), Dunbar-Brander (1927), Pocock (1939), Prater (1947), Ahmed (1954),

and Walker (1964), which contain little information on the less conspicuous species, including

rodents and bats; 2) distributional checklists, notably that by Ellerman and Morrison-Scott

(1951) and Siddiqi (1961) - these show no evidence of collections within the present study area;

3) natural history accounts, largely from the past century, such as those by Langley (1860),

Adams (1867), and Douie (1916), which provide some ecological information, as do government

census reports and gazetteers (Anonymous., 1908, 1935, 1961); and 4) modern studies made in

neighboring areas by Hatt (1959) and Petter (1961) for Iraq and Iran, and by Prakash (1939) for

Rajasthan. In addition, Mahmood-ul-Hassan (2009) has reviewed the literature on the bats of

Pakistan.

Roberts (1997) and Bates and Harrison (1997) are the main sources of knowledge on bats from this regions prior to this work. But none of these sources is based on actual field work on bats in this region. Taber et al., (1967) conducted a seven month survey of mammals of Lyallpur region and documented four bat species from this region. These included Pipistrellus ceylonicus

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subcanus, P. kuhlii kuhlii, P. mimus mimus and Scotophilus heathii. Pipistrellus mimus mimus has been reclassified as Pipistrellus tenuis from the same region. Roberts (1997) has listed 14 bat species of representing eight genera and five families from the present study area. These included Rhinolophus blasii, Rhinopoma microphylum, R. hardwickii, Taphozous nudiventris, T. perforatus, Hippociderous fulvus, H. cineraceus, Scotoecus pallidus, Scotophilus heathii, S. kuhlii, Pipistrellus javanicus, P. tenuis, P. pipistrellus and Hypsugo savii. A total of 182 bats belonging to twelve species, seven genera and three families were collected during the present study. These included Rhinolophus blasii (n = 3), Rhinopoma hardwickii (n = 2), Taphozous nudiventris (n = 26), T. perforatus (n = 4), Scotoecus pallidus (n = 2), Scotophilus heathii (n =

53), S. kuhlii (n = 5), Pipistrellus ceylonicus (n = 22), 9 P. javanicus (n = 9), P. pipistrellus (n =

52), 2 P. tenuis (n = 2) and Hypsugo savii (n = 2) (Present study).

Of these species Taphozous nudiventris, T. perforatus, S. heathii, S. kuhlii, Pipistrellus ceylnicus, P. javanicus, P. pipistrellus, P. tenuis and H. savii were captured from Margalla Hills

National Park and adjacent areas (SA1), S. heathii, P. javanicus, P. pipistrellus and H. savii from

Chinji National Park and adjacent areas (SA2), R. hardwickii. T. nudiventris, T. perforatus, S. pallidus, S. heathii and P. pipistrellus from Lal Suhanara National Park and adjacent areas (SA3) and Rhinolophus blasii, S. heathii, S. kuhlii, P. ceylonicus and P. pipistrellus from four districts of Central Punjab viz., Lahore, Gujranwala, Kasur and Toba Tek Singh (SA4).

Despite the same netting effort i.e. 3600 m2 hrs in four SAs, the netting index was not the

same. It was, in decreasing order 1.64 in SA1, 0.90 in SA4, 0.47 in SA3 and 0.42 in SA2. The

netting index for S. heathii at SA1, SA2, SA3 and SA4 was 0.50, 0.19, 0.28 and 0.31,

respectively while for P. pipstrellus it was 053, 0.03, 033 and 0.35, respectively. both these

species were recorded from all the four SAs. Scotophilus kuhlii and P. ceylonicus were recorded

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from SA1 and SA4. Netting indices for both these species were 0.03 and 0.07, and 0.42 and 0.12,

respectively. Pipistrellus Javanicus and H. savii were recorded SA1 and SA2 and netting indices

for both these species were 0.08 and 0.17, and 0.03 each respectively. Pipistrellus tenuis and

Scotoecus pallidus was recorded only from SA1 and SA3 respectively. Netting indices for both these species was 0.06 each. Population estimates for any bat species of the region is not available for comparison.

SPECIES DISTRIBUTION

Family Rhinolophidae

Rhinolophus blasii. This species is distributed in the world from NE South Africa to S. Dem.

Rep. Congo; Ethiopia; Somalia; Morocco; Algeria; Tunisia; Turkey; Yemen; Israel; Jordan;

Syria; Iran; Serbia and Montenegro; Albania; Bulgaria; Romania; Transcaucasia and

Turkmenistan; Afghanistan; Pakistan; Italy; Greece; Cyprus (Simmons, 2005).

In Pakistan, this species is represented by its subspecies R. b. meyeroemi (Corbet and

Hill, 1992). The population status of the species in the Indian subcontinent is unknown therefore it is considered to be a marginal species in the region. (Bates and Harrison, 1997). A single specimen of the species was collected Z. B. Mirza and T. J. Roberts (Roberts, 1997) on 28

December 1968 from Shalimar Gardens, Lahore. The species was included in “Lower risk: near threatened’ in the 1996 IUCN Red List of threatened Animals (Baillie and Groombridge, 1996).

The IUCN / SSC Action Plan (2001) and IUCN 2003 designated it as a species in Lower Risk

(nt) category. It is near threatened in South Asia (Walker and Molur, 2003), near threatened according to IUCN 2007 criteria and least concern according to IUCN 2008 Red List of

Threatened Animals. During the present study this species was recorded from Manawa

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(31º35.647, 074º27.660) in Lahore district. The specimens were collected with the help of a hand

net from an underground tunnel.

Family Rhinopomatidae

Rhinopoma hardwickii. The species is globally distributed from Morocco to Burma, south to

Mauritania, Senegal, Mali, Burkina Faso, Niger, and Kenya; Socotora Isles (Yamen). In Pakistan

this species has been collected from Rohtas in Salt Range, from around Karachi and Landhi in

southern Sindh and Karchat Hills near Hyderabad (Roberts, 1997; Mahmood-ul-Hassan et al.,

2009). The species is rare and locally distributed. This bat uses the same type of diurnal roosts as R. microphyllum. IUCN 2003 and IUCN / SSC Action Plan (2001) – Low Risk (lc). During the present study the species was recorded from Noor Mahal (29º22.695, 071º40.132) in

Bahawalpur district.

Family Emballonuridae

Taphozous nudiventris. This species is globally distributed from Mauritania, Senegal, and

Guinea-Bissau to Djibouti, Egypt, Jordan, NE Turkey, south to Tanzania and east to Burma. In

Pakistan the species is wide spread and common. In Punjab, specimens have been collected from

Multan (Roberts, 1997), Salt Range (Lindsay, 1927; Roberts, 1997) and Bahawalpur (Roberts,

1997). It occurs throughout Sindh from Sukker and Kahairpur in the north to Jaccobabad and

Thatta in the south (Wroughton, 1916; Siddiqui, 1961; Roberts, 1997) however, it has not been collected from any mountainous region of Baluchistan or NWFP (Roberts, 1997). IUCN 2008 –

LC. During the present study the species was recorded from Ratowal (33º28.644, 072º43.638) in

Attock district, from shrine of Abdul Maroof Shah at Mojgarh (29º01.132, 072º08.427) and from

Derwar Fort (28º46.045, 071º20.210) in Bahawalpur districts.

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T. perforatus. This species is distributed from Mauritania and Senegal to Botswana,

Mozambique, Somalia, Djibouti, and Egypt; S Arabia; Jordan; S Iran; Pakistan; NW India. In

Pakistan the species is uncommon and limited in distribution. The Zoological Survey of Pakistan recorded this species at Jatti near Thatta district (Siddiqui, 1961). It has not been recorded from elsewhere in Pakistan (Roberts, 1977). This species is not threatened currently but may be vulnerable to human disturbance of roosts (Bates et al., 1994b). IUCN 2008 – LC. During the present study the species was recorded from Ratowal (33º28.644, 072º43.638) in Attock district, from shrine of Abdul Maroof Shah at Mojgarh (29º01.132, 072º08.427) and from Derwar Fort

(28º46.045, 071º20.210) in Bahawalpur districts.

Family Vespertilionidae

Scotoecus pallidus. This species is endemic to the Indian subcontinent and has a local and restricted distribution in Pakistan. It was first described by Dobson in 1867 from a specimen collected from Mian Mir (Lahore). Further collections made from different regions of northern

Sindh (Kashmore and Mirpur in Jacobabbad, Larkana, Sukker and Dadu Districts) and Punjab

(Muzaffargarh and Sialkot). Its population status is uncertain and deserves further study. IUCN

2008 – LC. During the present study this species was recorded from Bahawalpur Fish Hatchery

(29º23.186, 071º38.148) in Bahawalpur district.

Scotophilus heathii. This species is geographically distributed from Afghanistan to S China, including Hainan Isl, south to Sri Lanka, Vietnam, Cambodia, Thailand and Burma. In Pakistan this species is common and widespread throughout the Indus plains. It has been collected from

Kohat (NWFP), Islamabad city, Multan, Lahore and Sialkot districts (Punjab), Kashmoor,

Sakkur, Jacobabad, Mirpur Sakro, Dadu, Landi, Malir, Karachi (Sindh) (Wroughton, 1916;

Lindsay, 1926; Siddiqui, 1960; Taber et al., 1967; Walton, 1974; Roberts, 1997). IUCN 2008 –

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LC. During the present study this species was recorded from all the four SAs. It was recorded

from PMNH (33º43.194, 073º03.631), Rawal Town (33º40.966, 073º07.108), NARC (33º39.892,

073º07.108) in SA1, Dalwal (32º42.725, 072º53.065) in SA2, Bahawalpur Fish Hatchery

(29º23.186, 071º38.148) in SA3 and Kalian Daas (31º12.037, 072º40.487), Govt. College Gojra

(31º09.024, 072º41.077), Rasul Nagar (32º19.424, 073º46.623) and Pattoki (31º02.487,

073º52.536) in SA4.

Scotophilus kuhlii. Geographically this species is distributed from Bangladesh, Pakistan to

Taiwan, south to Sri Lanka, Burma, Cambodia, W Malaysia, Java, Bali, Nusa Tenggara

(Indonesia), southeast to Philippines and Aru Isles (Indonesia). This species is uncommon in

Pakistan and has a very restricted distribution. It is found only in southern Sindh (Roberts, 1997).

IUCN 2008 - LC. During the present study the species was recorded from PMNH (33º43.194,

073º03.631) in SA1 and Badian (31°29.223, 074°24.632) in SA4.

Pipistrellus ceylonicus. Geographically this species is distributed from Pakistan, India, Sri

Lanka, Bangla Desh, Burma, Kwangsi and Hainan (China), Vietnam, Borneo. In Pakistan the species seems to be common in Pakistan. It is likely to be present in warmer southern latitudes in

the Indus plains. It is particularly abundant around Karachi and in Thatta district in Sindh. Taber

et al. (1967) collected it from Lyallpur (Faisalabad) and Khanewal. IUCN 2008 – LC. During the present study the species was recorded from PMNH (33º43.194, 073º03.631) and Rawal Town

(33º40.966, 073º07.108) in SA1 and Manawa (31º35.647, 074º27.660) in SA4.

P. Javanicus. Globally this species is distributed from E Afghanistan, N Pakistan, N, C India, SE

Tibet (China), Burma, Thailand, Vietnam, Through SE Asia to Lesser Sunda Isles and

Philippines; perhaps Australia. No literature is available on the distribution of this species in

Pakistan however a single specimen was collected from Gharial, Murree Hills. IUCN 2008 – LC.

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During the present study the species was captured from PMNH (33º43.194, 073º03.631) and

Rawal Town (33º40.966, 073º07.108) in SA1 and from Dalwal (32º42.725, 072º53.065) and

Ucchali Lake (32º30.329, 072º00.305) in SA2.

P. pipistrellus. This species is distributed from British Isles, S Denmark, W Europe to the Volga

and Caucasus, Morocco; Greece, Turkey, Israel and Lebanon to Afghanistan, Kashmir,

Kazakhstan, Pakistan, Burma, Sinkiang (China), perhaps Korea, Japan and Taiwan in the world.

This species has a restricted range in the Indian subcontinent (Bates and Harrison, 1997) and

seems to be common in Pakistan as there has been no further field studies on bats in Kashmir or

Gilgit (Roberts, 1997) . The Brisitsh Museum has one specimen that was collected from Kashmir

in the beginning of 19th century. Two other specimens were collected from Gilgit by an expedition carried out by University of Maryland in 1965 (Roberts, 1997). IUCN 2008 – LC.

During the present study this species was recorded from NARC (33º39.892, 073º07.108), Rawal

Town (33º40.966, 073º07.108) in SA1, Dalwal (32º42.725, 072º53.065) in SA2, Bahawalpur

Fish Hatchery (29º23.186, 071º38.148) in SA3 and Kalian Daas (31º12.037, 072º40.487), Rasul

Nagar (32º19.424, 073º46.623), Ali Pur Chattha (32º11.272, 074º09.361) and Pattoki (31º02.487,

073º52.536) in SA4.

P. tenuis. Geographically this species is distributed from Afghanistan to the Moluccas; S China,

Laos, Vietnam, Cocos Keeling Isles and Christmas Isle (Indian Ocean). In Pakistan the species has been recorded from Malakand (Roberts, 1997), Chitral (Sinha, 1980), Multan, Chaklala

(Hinton and Thomas, 1926), Chakri (Siddiqui, 1961), Gambat, Sukkur (Siddiqui, 1961), Karachi,

Malir (Walton, 1974). IUCN 2008 – LC. During the present study the species was captured from

Rawal Town (33º40.966, 073º07.108) and NARC (33º39.892, 073º07.108) in SA1.

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Hypsugo savii. This species was globally distributed from France, Portugal, Spain, Italy, S

Switzerland, Austria, E Hungary, Balkan Countries, Morrocco, N Algeria, and the Canary Isles.

(Spain) and Cape Verde Isls through the Crimia and Caucasus, Turkey, Lebanon, Syria, Israel,

Iran, Kazakhistan, Turkemenistan, Uzebekistan, Kyrgyzstan, Tajikistan, Afghistan to N India

and Burma (Wilson and Reeder, 2005). Nothing is known about the distribution and population

status of this bat in the Indian subcontinent (Bates and Harrison, 1997). This bat has not been

shown by Simmons (2005) to be present in Pakistan; Roberts (1997) and Walker and Molur

(2003) have, however shown this species to be present in East Punjab, India where it is

Vulnerable. The distribution and status of this species is unknown in Pakistan and based on

literature survey it can be stated that it may be present in Pakistan. IUCN 2008 - LC. During the present study the species was recorded from PMNH (33º43.194, 073º03.631) in SA1 and from

Ucchali Lake (32º30.329, 072º00.305) in SA2.

The present study indicates that population of the four bat species viz. Rhinopoma microphyllum, Hipposideros fulvus, H. cineraceus and Pipistrellus kuhlii have considerably declined over the past few decades. They were recorded by Roberts (1997) from the study area but neither any roost was located nor any specimen of these species was captured during the whole study period from any of the four SAs. These four species may be considered as

Threatened.

Another group of six bat species viz. Rhinolophus blasii, R. hardwickii, T. nudiventris and Scotoecus pallidus, Scotophilus heathii and Pipistrellus pipistrellus were captured during the present study. These species have also been reported from the same area (Roberts, 1997; Bates and Harrison, 1997; Mahmood-ul-Hassan et al., 2009). Of these S. heathii and P. pipistrellus were captured from all the four SAs while S. pallidus was mist netted from SA3 only; the former

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two were the commonest bats of the study area while netting index for the latter was low. The

other three bat species i.e. R. blasii, R. hardwickii and T. nudiventris were captured from their

day time roosts but their colony sizes were considerably smaller than those reported in literature

(Roberts, 1997; present study). The remaining six bat species, in order of their abundance, P.

ceylonicus, P. javanicus, S. kuhlii, T. perforatus, P. tenuis and Hypsugo savii show range

extension (Roberts, 1997; Bates and Harrison, 1997; Present study).

PART III. MORPHOLOGY

The morphology of the twelve bat species recorded during the present study is discussed

as follows.

Family Emballonuridae

Taphozous nudiventris. All the morphometric measurements viz. head and body length, ear length, forearm length, hind foot length and tail length of the twenty six Taphozous nudiventris

specimens captured during the present study are in accordance with the same measurements

given by Roberts (1997) and Bates and Harrison (1997). Certain cranial measurements of the

Taphozous nudiventris samples captured during the present study fall within the ranges given by

Bates and Harrison (1997). These include breadth of braincase, zygomatic breadth, greatest

length of skull, maxillary toothrow, mandibular toothrow and mandible length while postorbital

constriction of the T. nudiventris samples captured during the present study was smaller than the

measurements by Bates and Harrison (1997) (Table 52).

T. perforatus. The average ear and tail length of the four Taphozous perforatus captured during the present study falls within the range of the measurements of the same parameters given by

Bates and Harrison (1997). However the lower limits of head and body length, forearm length, length of 3rd metacarpal and hind foot length are within the range of the measurements of Bates

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Table 52. Mean external body and cranial measurements of Taphozous nudiventris (I = Roberts (1997), II = Bates and Harrison (1997), III = Present study).

I II III (n= 26) Parameters (mm) Head and Body Length 92 (86-98 ) 96.3(90.0 – 105.0) 86.87(82-96) Ear length 22(19-23) 22.0 (18.0 – 25.0) 14.70(11-19) Forearm 71.0 74.3 (71.0 – 80.0) 71.00(70-74.5) Hind foot 14(11-18) 14.6(11.0 – 18.0) 15.57(14-18.5) Tail 36.5(34-42) 32.6(22.0 – 46.0) 27.57 (19-49) Breadth of braincase - 11.5(10.6 – 12.7) 10.87(10.74-11.00) Zygomatic breadth - 15.9(14.4 – 17.8) 14.15(12.95-15.34) Postorbital constriction - 8.1(7.0 – 8.6) 5.18(5.03-5.33) Condylo-canine length - 23.4(21.6 – 25.6) 19.27(18.54-19.99) Greatest length of skull - 25.8(22.5 – 27.9) 26.16(25.93-26.39) Mandible length - 20.1(18.2 – 21.5) 17.53(16.71-18.34)

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and Harrison (1997). The zygomatic breadth and condylo-canine length of single Taphozous

perforatus captured during the present study were within the range of the same measurements

given by Bates and Harrison (1997). However the breadth of the braincase, greatest length of

skull, mandibular toothrow length and the length of mandible were slightly higher while

maxillary toothrow length was slightly smaller than given by Bates and Harrison (1997) (Table

53).

Family Vespertilionidae

Scotoecus pallidus. Table 54 indicates that the average head and body length, forearm length,

length of 3rd metacarpal, hind foot length and tail length of two S. pallidus captured during the

present study were within the range of measurements given by Bates and Harrison (1997). The

average head and body length was also within the range given by Roberts (1997). The length of

4th metacarpal and the length of 5th metacarpal were slightly smaller than the measurements of

Bates and Harrison (1997). The average postorbital constriction, condylo-canine length and greatest length of skull were within the range of measurements of the same parameters given by

Bates and Harrison (1997) while the average zygomatic breadth, maxillary toothrow length, mandibular toothrow length and the length of mandible were smaller than the measurements given by Bates and Harrison (1997).

Scotophilus heathii. The average body mass, head and body length, forearm length, length of 3rd metacarpal, length of 4th metacarpal, hind foot length and the tail length of thirty nine

Scotophilus heathii captured during the present study were within the range given for the same

species by Bates and Harrison (1997). The average ear length and the length of 5th metacarpal

were slightly smaller than the measurements of Bates and Harrison (1997). The mean breadth of

braincase, postorbital constriction, condylo-canine length, maxillary toothrow length, mandibular

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Table 53. Mean external body and cranial measurements of Taphozous perforatus (I = Roberts (1997), II = Bates and Harrison (1997), III = Present study).

I II III Parameters (mm) Head and Body Length - 74.5(71.0 – 80.0) 84.30 (78.5-92.00) Ear length 18 18.0(14.0 – 20.0) 15.30(13-18) Forearm 62.3 (59-63) 61.3(59.2 – 63.8) 64.30 (61.5-70) 3rd metacarpal - 55.4(53.7 – 57.2) 60.40 (54.5-64.5) 1st phalanx on 3rd metacarpal - 19.0(18.3 – 19.7) 25.80 (24-28) Hind foot 11.5 10.1(8.2 – 12.5) 14.60 (12-17) Tail (21-27) 24.0(20.0 – 28.0) 22.10 (19-25.5) Breadth of braincase - 9.3(9.2 – 9.6) 10.01 Zygomatic breadth - 11.7(11.5 – 12.1) 12.01 Condylo-canine length - 19.0(18.4 – 19.7) 19.58 Greatest length of skull - 20.5(19.9 – 21.5) 22.24 Mandible length - 15.0(14.6 – 15.6) 16.25 *Mean and range for a particular parameter is given when atleat two specimens were recorded.

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Table 54. Mean external body and cranial measurements of Scotoecus pallidus (I = Roberts (1997), II = Bates and Harrison (1997), III = Present study).

Parameters I II III (n= 2) (mm) Head and Body Length 54(50 – 59) 52.8 (50.0 –58.0) 56.50(54-59) Ear length 13 12.8(12.0 – 15.0) 9.50(9-10) Forearm - 36.2(34.1 – 37.3) 35.50(35-36) 3rd metacarpal - 34.6(33.5 – 36.0) 33.75(33.5-34) 4th metacarpal - 34.2(32.8 – 35.4) 32.25(32.0-32.5) 5th metacarpal - 33.7(32. 6 –34.9) 31.75(31.5-32.0) Hind foot 8 8.3 (6.0 – 10.0) 8.50(8-9) Tail 37(31 – 34) 36.9(34.0 – 41.0) 35.50(33-38) Breadth of braincase - 7.7(7.5 – 8.2) 7.12(6.6-7.6) Zygomatic breadth - 10.5(10.5 – 10.5) 9.91(9.7-10.1) Postorbital constriction - 4.3(4.2 – 4.5) 4.37(4.0-4.7) Condylo-canine length - 14.1(13.8 – 14.8) 14.69(14.4-14.9) Greatest length of skull - 15.1(14.5 – 16.1) 15.46(15.1-15.7) Maxillary toothrow - - 4.34(3.0-5.7) Mandibular toothrow - - 4.32(3.3-5.3) Mandible length - 11.4(10.9 – 12.0) 9.64(7.9-11.3)

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toothrow length, and length of mandible of twenty Scotophilus heathii skulls were within the

range of the measurements given by Bates and Harrison (1997). While the average zygomatic

breadth and greatest length of skull were slightly smaller than the ranges of measurements given by Bates and Harrison (1997) (Table 55).

Scotophilus kuhlii. Table 56 indicates that the average head and body length, ear length, forearm

length, length of 3rd metacarpal, length of 4th metacarpal, hind foot length and tail length of the

five Scotophilus kuhlii captured during the present study were within the range of the

measurements given by Bates and Harrison (1997). The head and body length was also within the range given by Roberts (1997). While ear length, hind foot length and tail lengths were

slightly smaller than that given by Roberts (1997). The average zygomatic breadth, postorbital constriction, condylo-canine length, greatest length of skull, maxillary toothrow length, mandibular toothrow length and the length of mandible of five Scotophilus kuhlii captured during the present study were within the ranges given by Bates and Harrison (1997) while the average breadth of braincase was slightly broader than the range.

Pipistrellus ceylonicus. Average head and body length, of the twenty two Pipistrellus ceylonicus

captured during the present study was greater than the specimens measured by Roberts (1997); however it was within the range of the measurements taken by Bates and Harrison (1997). The average hind foot length was also within the range given by Bates and Harrison (1997). The

upper limit of the ear length, forearm length and length of the tail was within the ranges given by

Bates and Harrison (1997). The average zygomatic breadth and postorbital constriction of the

five Pipistrellus ceylonicus captured during the present study was within the range of the

measurements given by Bates and Harrison (1997). The average breadth of the braincase,

condylo-canine length, maxillary toothrow length, posterior palatal width, mandibular toothrow

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Table 55. Mean external body and cranial measurements of Scotophilus heathii (I = Roberts (1997), II = Bates and Harrison (1997), III = Present study).

Parameters I II III (n= 53) (mm) Body mass (36-39g) (36-39g) 35.6(24.6-46)g Head and Body Length 83(72 – 92) 82.5(67.0 – 93.0) 79.46(71-94) Ear length 16(14 – 17) 12.7(9-16) Forearm - 60.7(55.4 – 65.8) 58.70(52.5-64) 3rd metacarpal - 59.4(53.7 – 68.4) 55.04(52-58) 4th metacarpal - 58.2(54.0 – 63.9) 52.09(50.5-58) 5th metacarpal - 54.6(50.3 – 59.9) 50.13(47-53.5) Hind foot 12(11 – 13) 12.0(9.0 – 15.0) 12.8(10.5-14.5) Tail 55(51 – 60) 59.1(43.0 – 71.0) 55.0(45.5-68) Breadth of braincase 10.2(9.7 – 10.8) 9.99(9.2-12.4) Zygomatic breadth 15.6(14.5 – 16.9) 14.31(12.4-15.8) Postorbital constriction 5.5(5.2 – 5.9) 5.38(4.5-6.3) Condylo-canine length 20.2(19.0 – 21.3) 19.01(17.0-20.6) Greatest length of skull 23.4(21.7 – 25.2) 21.39(18.0-23.0) Maxillary toothrow - 7.82(7.1-9.4) Mandibular toothrow - 8.46(7.7-9.1) Mandible length 16.3(14.8 – 18.0) 16.08(12.9-17.2)

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Table 56. Mean external body and cranial measurements of Scotophilus kuhlii (I = Roberts (1997), II = Bates and Harrison (1997), III = Present study).

Parameters I II III (n= 5) (mm) Head and Body Length 76(64 – 87) 69.8(60.0 – 78.0) 72.10(65-86) Ear length 17 13.5(9.0 – 17.0) 12.10(9-16) Forearm - 49.0(44.0 – 56.4) 49.40(44.5-52) 3rd metacarpal - 48.8(44.4 – 58.8) 47.30(42-54.5) 4th metacarpal - 48.3(43.7 – 57.2) 44.90(39.5-47.5) 5th metacarpal - 45.0(42.1 – 53.9) 41.10(35.5-43.5) Hind foot 11 10.0(8.0 – 13.0) 10.60(10-11.5) Tail 52(47 – 58) 47.5(40.0 – 65.0) 42.40(38-46) Breadth of braincase - 8.9(8.8 – 9.4) 9.59(8.7-12.4) Zygomatic breadth - 13.0(12.4 – 13.7) 13.43(13.0-14.9) Postorbital constriction - 4.7 (4.4 – 5.1) 4.73(4.1-5.1) Condylo-canine length - 17.3(16.3 – 18.0) 17.62(16.8-18.0) Greatest length of skull - 19.6(18.7 – 20.4) 18.98(18.0-19.7) Maxillary toothrow - - 6.27(5.4-6.6) Mandibular toothrow - - 7.53(6.5-8.7) Mandible length - 13.7(12.9 – 14.4) 14.41(13.2-16.3)

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length and the length of mandible were slightly smaller than the ranges mentioned by Bates and

Harrison (1997) (Table 57).

P. javanicus. The average head and body length, ear length, forearm length, length of the 3rd metacarpal, length of the 4th metacarpal length of 5th metacarpal, hind foot length and the length

of the tail of ten Pipistrellus javanicus captured during the present study were within the range of

the measurements given by Bates and Harrison (1997). The average breadth of braincase,

zygomatic breadth, postorbital constriction, maxillary toothrow length were slightly larger while

condylo-canine length, greatest skull length, mandibulat toothrow length and the length of the

mandible were within the ranges given by Bates and Harrison (1997) (Table 58).

P. pipistrellus. The mean values for all the parameters compared with available literature were

slightly lower than the measurements mentioned by Bates and Harrison (1997). However the

upper limits for head and body length, ear length, forearm length and tail length were in

accordance with the measurements given by Bates and Harrison (1997). The mean values for all

the parameters compared with available literature were slightly lower than the measurements

mentioned by Bates and Harrison (1997). However the upper limits for head and body length, ear

length, forearm length and tail length were within the range of measurements given by Bates and

Harrison (1997). The average breadth of braincase was within the range of measurements (Bates

and Harrison, 1997). The zygomatic breadth and the postorbital constriction were slightly larger

while all the other cranial parameters were smaller than Bates and Harrison (1997) (Table 59).

P. tenuis. The mean head and body length, ear length, forearm length, length of 3rd metacarpal,

length of 4th metacarpal, calcar length and tail length of the two Pipistrellus tenuis captured during the present study were within the range of measurements given by Bates and Harrison

(1997). The breadth of the braincase, postorbital constriction, condylo-canine length and

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Table 57. Mean external body and cranial measurements of Pipistrellus ceylonicus (I = Roberts (1997), II = Bates and Harrison (1997), III = Present study).

Parameters I II III (n= 22) (mm) Head and Body Length 51(48- 56) 53.5(45.0 – 64.0) 63.60(54-77) Ear length 12 12.2(9.5 – 14.0) 8.06(5.5-12) Forearm 35 37.2(33.0 – 42.0) 29.92(28-38) 3rd metacarpal - 35.8(33.0 – 39.5) 25.85(21-32.5) 4th metacarpal - 35.1(32.6 – 38.8) 25.49(21-30) 5th metacarpal - 33.6(30.7 – 36.7) 25.52(20-30.4) Wing span - 251.1(227 – 262) 192(189-208) Hind foot 9 8.3 (6.0 – 11.0) 6.34(5.5-9) Tail 38 (36-44) 38.2(30.0 – 45.0) 25.68(22-31) Breadth of braincase - 7.3(6.8 – 7.8) 6.55(5.5-9.4) Zygomatic breadth - 9.8(9.2 – 11.0) 9.62(5.8-15.2) Postorbital constriction - 4.0(3.7 – 4.3) 4.04(3.5-5.4) Condylo-canine length - 13.7(13.1 – 14.3) 10.08(9.5-10.5) Greatest length of skull - 15.0(14.4 –15.8) 10.76(10.4-11) Maxillary toothrow - - 5.01(3.5-9.4) Posterior palatal width - - 5.88(4.5-9.5) Mandibular toothrow - - 4.64(3.6-8.3) Mandible length - 11.2(10.6 – 12.0) 9.28(6.9-16.0)

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Table 58. Mean external body and cranial measurements of Pipistellus javanicus (I = Bates and Harrison (1997), II = Present study).

Parameters I II (n= 9) (mm) Head and Body Length 47.1(40.0 – 55.0) 52.00(48.0-57.0) Ear length 11.8(5.0 – 15.0) 10.20(7.6-16.5) Forearm 33.2(30.0 – 36.0) 35.13(32.5-37.5) 3rd metacarpal 33.0(25.9 – 34.8) 31.38(28.5-35.0) 4th metacarpal 32.6(29.9 – 34.7) 31.06(26.35.5) 5th metacarpal 31.4(29.0 – 33.4) 30.31(26-34.5) Hind foot 6.0(3.0 – 8.0) 7.89(7-9.5) Tail 33.9(26.0 – 40.0) 30.38(19-35) Breadth of braincase 6.6(6.3 – 7.1) 7.31(5.8-9.6) Zygomatic breadth 8.5(8.2 – 9.0) 10.23(7.0-13.9) Postorbital constriction 3.7(3.3 – 4.3) 4.58(3.6-5.6) Condylo-canine length 12.4(11.9 – 13.1) 12.08(9.8-20.3) Greatest length of skull 13.6(13.0 – 14.6) 14.09(11.3-21.1) Maxillary toothrow - 5.35(3.3-7.3) Mandibular toothrow - 5.26(4.8-6.1) Mandible length 9.9(9.3 – 10.7 ) 10.29(7.3-11.2)

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Table 59. Mean external body and cranial measurements of Pipistellus pipistrellus (I = Roberts (1997), II = Bates and Harrison (1997), III = Present study).

Parameters I II III (mm) Head and Body Length (42- 45) 44.0(40.0 – 48.0) 39.33 (32.0-44.5) Ear length (11 - 12) 11.1(10.5 – 12.0) 8.81(6.0-11.0) Forearm ‐ 31.0(30.0 – 31.6) 28.23(26.0-31.0) 3rd metacarpal ‐ 29.9(29.5 –31.0) 25.27(23.5-28.5) 4th metacarpal - 29.6(28.7 – 30.8) 24.81(22.0-27.5) 5th metacarpal - 28.9(28.4 – 29.8) 24.63(23.0-27.5) Hind foot 7 6.1(6.0 – 7.0) 6.31(5.0-8.0) Tail (33 - 34) 32.9(29.0 – 35.0) 25.86(21.0-31.6) Breadth of braincase - 6.1(5.9 – 6.3) 6.01(5.5-6.9) Zygomatic breadth - 7.6(7.2 – 7.9) 8.56(6.3-12.4) Postorbital constriction - 3.3(3.2 – 3.5) 3.55 (3.2-3.9) Condylo-canine length - 10.9(10.4 – 11.3) 10.02(8.5-10.5) Greatest length of skull - 12.1(11.9 – 12.5) 11.04(10.5-11.6) Maxillary toothrow - ‐ 3.56(3.2-3.9) Posterior palatal width - ‐ 4.80(4.5-5.2) Mandibular toothrow - ‐ 3.43 (2.8-4.0) Mandible length - 8.4(7.9 – 8.7) 7.87 (6.4-9.9)

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mandible length of the single Pipistrellus tenuis were within the measurements ranges given by

Bates and Harrison (1997) (Table 60).

Hypsugo savii. The mean head and body length, forearm length, length of the 4th metacarpal and

tail length of the two Hypsugo savii captured during the present study were within the

measurement ranges of Bates and Harrison (1997) whereas the ear length and wing span were

slightly smaller while length of 3rd metacarpal, length of 5th metacarpal, and hind foot length

were slightly larger than the measurements given by Bates and Harrison (1997). The breadth of

braincase, postorbital constriction, maxillary toothrow length, mandibular toothrow length and

the length of the mandible of single Hypsugo savii captured during the present study were

slightly larger while zygomatic breadth, condylo-canine length, and greatest length of skull were slightly smaller than the measurements given by Bates and Harrison (1997) (Table 61).

1997; Benda and Tsytsulina, 2000). More than 160 Palaearctic vespertilionids viz. Plecotus,

Barbastella, and Myotis (Selysius subgenus) which are morphologically very similar have been

differentiated on the basis of this attribute (Thomas, 1915; Chaine, 1926; Topal, 1958; Hill and

Harrison, 1987; Strelkov, 1989a; 1989b; Yoshiyuk, 1989; Smirov, 2000; Strelkov, 1989a,

1989b; Benda and Tsytsulina, 2000; Kruskop and Lavrenchenko, 2000; Tsytsulina, 2001).

Taphozous nudiventris magnus from T.n. kachhensis are also differentiated through this distinctive feature (Asan and Albayrak, 2007). Baculum length in various bat species is positively associated with relative male body mass but this association has not been proved substantially through phylogenetic testing (Hosken et al., 2001). Age peculiarities of baculum

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Table 60. Mean external body and cranial measurements of Pipistellus tenuis (I = Bates and Harrison (1997), II = Present study).

Parameters I II (n = 2) (mm) Head and Body Length 39.1(33.0 – 45.0) 35.00(33.0-37.0) Ear length 9.7(5.0 – 11.0) 7.50(7.0-8.0) Forearm 27.7(25.0 – 30.2) 28.00(27.5-28.5) 3rd metacarpal 26.7(23.9 – 29.7) 25.25(25.0-25.5) 4th metacarpal 26.4(23.7 – 29.2) 23.75(23.0-24.5) 5th metacarpal 25.9(23.5 – 28.5) 23.75(23.0-24.5) Calcar 5.3(3.0 – 7.0) 6.00(5.0-7.0) Tail 28.9(20.0 – 35.0) 22.25(20.0-24.5) Breadth of braincase 6.0(5.6 – 6.3) 6.30 Zygomatic breadth 7.4(7.3 – 7.6) 7.80 Postorbital constriction 3.3(2.9 – 3.7) 3.96 Condylo-canine length 10.2(9.3 – 10.7) 9.42 Greatest length of skull 11.5(10.7 – 12.1) 10.19 Maxillary toothrow - 3.78 Posterior palatal width - 5.23 Mandibular toothrow - 4.27 Mandible length 7.9(7.2 – 8.3) 7.82 *Mean and range for a particular parameter is given when atleat two specimens were recorded.

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Table 61. Mean external body and cranial measurements of Hypsugo savii (I = Bates and Harrison (1997), II = Present study).

Parameters I II(n=2) (mm) Head and Body Length 52.0(47.0 – 60.0) 55.50(42.0-69.0) Ear length 12.1(10.0 – 14.0) 9.50(9.0-10.0) Forearm 31.0(32.1 – 38.0) 36.75(34.0-39.5) 3rd metacarpal 31.9(30.4 – 33.2) 34.50(33.5-35.5) 4th metacarpal 31.5(30.2 – 34.0) 33.00(31.0-35.0) 5th metacarpal 30.5(29.1 –31.3) 31.95(30.0-33.9) Wing span 235.5(226.0 –251.0) 217.00(190.0-244.0) Hind foot 7.1(6.4 – 8.0) 8.50(8.0-9.0) Tail 33.0(30.0 – 35.0) 33.50(29.0-38.0) Breadth of braincase 6.7(6.6 – 6.8) 7.14 Zygomatic breadth 8.7(8.5 – 9.1) 7.44 Postorbital constriction 3.6(3.5 – 3.7) 4.52 Condylo-canine length 12.8(12.4 – 13.3) 10.41 Greatest length of skull 14.0(13.6 – 14.4) 11.10 Maxillary toothrow - 5.41 Mandibular toothrow - 5.38 Mandible length 9.8(9.6 – 10.3 mm) 11.02 *Mean and range for a particular parameter is given when atleat two specimens were recorded.

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shape have been studied in detail in Rodentia and Carnivora (e.g., Friley, 1949; Callery, 1951;

Heidt, 1970; Tarasov, 1974, 1984; Aksenova, 1980) but studied only sparingly in Chiroptera

(Maeda, 1978). Correlation between increasing condylobasal length, forearm length and baculum

length was found in Nyctalus aviator (Maeda, 1978) Murina silvatica, Vespertilio superans,

Myotis macrodactylus, and Rhinolophus cornutus (Yoshiyuk, 1989) but V. superans aged from

10 to 202 days showed no such relationship (Yoon et al., 1990). The comparison of the shape and size of bacula in Myotis myotis and Myotis blythi depicted that it is not the size but shape of

the baculum that distinguishes these two species (Albayrak and Asan, 2001).

The baculum of a single Rhinolophus blasii was 3.5 mm long, with a 2.5 mm long shaft.

The proximal and distal bacular lengths were 1.0 mm and 0.00 mm while proximal and distal

bacular widths were 1.0 mm and 0.5 mm, respectively. The bacular height was 1.1 mm. The

baculum of Rhinopoma hardwickii was 1.1 mm long with a shaft which was 1.0 mm long. The

proximal and distal bacular lengths were 0.1 mm and 0.00 mm while proximal and distal

breadths of baculum were 0.3 mm and 0.2 mm. The baculum was of 0.4 mm high.

Mean total baculum length of the two T. nudiventris was 0.58 mm ± 0.017 (SD) with

shaft length of 0.54 mm ± 0.069 (SD). The proximal and distal lengths of the baculum were 0.04

mm ± 0.052 (SD) and 0.0 mm, respectively. The proximal and distal bacular widths were 0.25

mm ± 0.035 (SD) and 0.20 mm ± 0.000 (SD), respectively. The bacula were 0.10 mm ± 0.000

(SD) high.

The total length of the baculum of T. perforatus was measured as 0.69 mm with shaft

length of 0.61 mm. The proximal and distal bacular lengths were 0.07 mm and 0.00 mm,

respectively. The proximal and distal breadths of baculum were 0.27 mm and 0.22 mm

respectively. The bacular height was 0.07 mm.

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Mean total bacular length of ten Scotophilus heathii was measured 1.76 mm ± 0.150 (SD)

with shaft length of 1.51 mm ± 0.232 (SD). The proximal and distal baular lengths were 0.19

mm ± 0.078 (SD) and 0.04 mm ± 0.034 (SD), respectively while proximal and distal bacular

breadths were 0.95 mm ± 0.120 (SD) and 0.46 mm ± 0.058 (SD), respectively. The bacular

height was 0.29 mm ± 0.074 (SD).

The total length of the baculum of single S. kuhlii was 1.74 mm with a shaft length of

1.52 mm. The proximal and distal bacular lengths were 0.07 mm and 0.15 mm, respectively. The proximal and distal bacular widths were 1.05 mm and 0.49 mm, respectively while the baculum was 0.49 mm in height.

The total length of a single Scotoecus pallidus captured from SA3 was 5.0 mm with shaft length of 4.1 mm. The proximal and distal bacular lengths were 0.6 mm and 0.3 mm, respectively. The proximal and distal breadths of the baculum were 1.0 mm and 0.5 mm, respectively. The baculum height was 0.5 mm.

Mean total length of the bacula of four Pipistrellus ceylonicus was 3.66 mm ± 1.190

(SD), with the shaft length of 2.93 mm ± 1.125 (SD). The proximal and distal bacular lengths

were 0.39 mm ± 0.2 06 (SD) and 0.28 mm ± 0.015 (SD), respectively while the proximal and

distal widths were 0.75 mm ± 0.183 (SD) and 0.37 mm ± 0.035 (SD), respectively. The bacular

height was 0.60 mm ± 0.188 (SD).

The mean total length of the bacula of P. javanicus was 3.57 mm ± 0.860 (SD) with a

shaft length of 2.77 mm ± 0.833 (SD). The proximal and distal bacular lengths were 0.47 mm ±

0.218 (SD) and 0.29 mm ± 0.023 (SD), respectively. The mean proximal and distal bacular

widths were 0.73 mm ± 0.064 (SD) and 0.42 mm ± 0.020 (SD), respectively. The baculum height

was 0.54 mm ± 0.040 (SD).

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The mean total length of the eleven bacula of P. pistrellus was 3.19 mm ± 0.421 (SD) while the shaft of the bacula was 2.35 mm ± 0.245 mm long. The proximal and distal bacular lengths were 0.59 mm ± 0.218 (SD) and 0.26 ± 0.075 (SD), respectively. The proximal and distal bacular widths were 0.76 mm ± 0.055 (SD) and 0.38 mm ± 0.059 (SD). The height of the baculum was 0.60 mm ± 0.156 mm.

The total bacular length of a single P. tenuis was 2.79 mm with shaft length of 2.08 mm.

The proximal and distal bacular lengths were 0.39 mm and 0.27 mm, respectively while proximal and distal bacular widths were 0.61 mm and 0.34 mm, respectively. The height of the baculum was 0.51 mm.

The total bacular length of a single H. savii captured from SA1 was 2.67 mm with shaft length of 2.08 mm. The proximal and distal bacular lengths were 0.39 mm and 0.27 mm, respectively while proximal and distal bacular widths were 0.61 mm and 0.34 mm, respectively.

The height of the baculum was 0.51 mm. No data for comparison of bacular measurements of bats of the region is available.

ECHOLOCATION

Insectivorous bats use various strategies to search and catch insects from air, within dense vegetation, from ground and foliage. Bats use ultrasonic echo-location calls (e.g. Findley, 1993;

Altringham, 1996) that enable them capture prey in total darkness. This adaptation makes them most suitable for exploiting nocturnal food resources (e.g. Moeller, 1985; Findley, 1993;

Vaughan et al., 1997b; Feng et al., 2000; Zhang et al., 2000; Rydell et al., 2002; Fukui et al.,

2004; Jones et al., 2006).

Table 62 compares call parameters of T. p. senegalensis to other studies (Dietz, 2005;

Benda et al., 2008) with almost similar Fmax, Fmin and FmaxE. The PD and IPI are however

164 different. Details of the bat echolocation calls vary considerably within the same species with respect to the region and habitat (Barclay et al., 1999). Recordings from Israel, Nile Valley and

United Arab Emirates also show that calls are narrowband with two prominent harmonics, the lower harmonic end frequency of the lower frequency ranges between 27 to 29 kHz (Deitz, 2005;

Davis, 2007). The difference in the call structure of this specimen may either be due to difference in subspecies or due to recording conditions.

Identification of bats by their ultrasonic calls helps us to infer their diversity through passive bat surveys. Reference calls of bats in the study area were not available and a bat reference call library is urgently needed for confirming the occurrence of bat species as established by Herr and Klomp (1995) at Australia. Furthermore, identification of some species by call is problematic, so an investigation of the variability of echolocation calls is also necessary to establish the validity and applicability of bat identification from their calls.

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Table 62. Descriptive parameters of echolocation calls of the Egyptian tomb bat (T. perforatus) from Pakistan. Explanation: n – number of individual calls analyzed (in parentheses number of call sequences from which calls were obtained); Fmax – start frequency; Fmin – end frequency; FmaxE – frequency with maximum energy (peak frequency); PD – pulse duration; IPI – inter-pulse interval; bold upper lines – mean±SD, lower lines – range.

Mean±SD Range Variables Present Dietz, Benda et al., Ulanovsky study 2005 2008 et al., 2010

N 8(1) - 8(1) 16

Fmax (kHz) 31.4±0.7 34.0±1.0 27.54±0.27 30-32 - 32.6–35.4

Fmin (kHz) 25.6±0.5 28.3±0.6 - 25-26 27 – 29 27.3–28.9

FmaxE (kHz) 30.0±0.4 - 30.1±0.8 - 29.4-30.5 28.9–31.0

PD (ms) 14.2±2.7 - 8.0±1.6 13.1±1.3 14.2-19.0 6.0–10.4

IPI (ms) 69.3±27.8 - 153.7±28.9 386±63 42.7-122.2 127.0–216.0

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SUMMARY

The present study was conducted from June 2009 to May 2011 in those arid subtropical and tropical regions of Pakistan which included less pronounced monsoon influenced areas of the Salt Range, the Upper Indus Plains and the sand dune areas typified by the Cholistan. Bat

surveys were conducted in two protected areas i.e. the Margallah Hills National Park (SA1) and

the Chinji National Park (SA2) that were located in the arid subtropical region and in another,

the Lal Suhanara National Park (SA3), situated in the tropical sand dune region of the Upper

Indus Plains. In addition, bat samples were also collected from Gujranwala, Lahore, Tob Tek

Singh and Kasur districts (SA4). These sub-areas were selected to maximize the chances of capture of as many bat species inhabiting arid-subtropical and tropical habitats of Pakistan as possible.

A total of 182 bats belonging to twelve species were recorded. These included R. blasii

(Family Rhinolophidae), R. hardwickii (Family Rhinopomatidae), Taphozous nudiventris and T. perforatus (Family Emballonuridae), Scotoecus pallidus, Scotophilus heathii, S. kuhlii,

Pipistrellus ceylonicus, P. javanicus, P. pipistrellus, P. tenuis and Hypsugo savii

(Vespertilionidae). Rhinolphous blasii was captured only from SA1, R. hardwickii and S. pallidus from SA3 and P. tenuis from SA1. Taphozous nudiventris and T. perforatus were captured from SA1 and SA3, S. kuhlii and P. ceylonicus from SA1 and SA4, H. savii from SA1

and SA2 and P. javanicus from SA1 and SA2. Scotophilus heathii and P. pipistrellus were

recroded throughout the study area.

Maximum bat activity was recorded in spring (n = 65) that was followed by summer ( n =

61), autumn (n = 32) and winter (n = 24). Rhinolophus blasii and S. pallidus were recorded only

during winter, R. hardwickii and P. tenuis during autumn, while S. kuhlii was recorded only

167

during summer. Taphozous nudiventris and T. perforatus were captured during summer and

autumn. Pipistrellus pipistrellus was recorded during autumn, spring and winter while S. heathii was captured throughout the year.

Although the netting effort was the same, the number of bats captured from the SAs was different. A total of 72 bats were recorded from SA1, 52 from SA4, 43 from Lal SA3 and 15 from SA2. The dominance was highest for SA2 and lowest for SA1. Both Shannon and Simpson indices show that the diversity was the highest at SA1 followed by SA3, SA4 and SA2. Evenness was found to be highest at SA4 followed by SA3, SA2 and SA1.

The mean head and body length of three Rhinolophus blasii was 39.33 mm ± 0.577 (SD) forearm length was 40.17 mm ± 1.155 (SD) and the tail length was 19.23 mm ± 1.940 (SD). The greatest skull length of a single R. blasii was 17.22 mm and mandible length was 11.80 mm. The baculum of a single R. blasii sample was 3.5 mm long.

The mean head and body length of two Rhinopoma hardwickii 66.00 mm ± 5.657 (SD).

The mean forearm length was 54.00 mm ± 0.0 (SD). The tail length was 59.00 mm ± 2.828 (SD).

The greatest skull length was 19.68 mm ± 0.108 (SD), and the length of mandible was 11.28 mm

± 1.652. The baculum of single R. hardwickii was 1.1 mm long.

The mean head and body length of twenty six Taphozous nudiventris was 86.87 mm ±

5.556 (SD) and the tail length was 27.57 mm ± 12.187 (SD). The greatest skull length was 26.16 mm ± 0.323 (SD) and the length of mandible was 17.53 mm ± 1.149 (SD). The mean total baculum length of the two specimens was 0.58 mm ± 0.017 (SD).

The head and body length of four T. perforatus was measured as 84.30 mm ± 5.450 (SD) long. The forearm was 64.30 mm ± 3.457 (SD) long and the length of tail was 22.10 mm ± 2.702

(SD). The greatest length of skull was 22.24 mm and the length of mandible was recorded as

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16.25 mm. The total length of a single T. perforatus was measured as 0.69 mm. The head and

body length of fifty three Scotophilus heathii was 79.46 mm ± 6.941 (SD). The mean forearm

length was 58.69 mm ± 2.929 (SD) and the tail length was 55.00 mm ± 7.360 (SD). The greatest

length of skull was 21.39 mm ± 1.378 (SD) and the length of mandible was recorded as 16.08

mm ± 0.882 (SD). Mean total bacular length of ten S. heathii was measured 1.76 mm ± 0.150

(SD).

The mean head and body length of five specimens of S. kuhlii was 72.10 mm ± 8.096

(SD). The forearm was 49.40 mm ± 3.03 (SD) long and the length of tail was 42.40 mm ± 4.04

(SD). The greatest length of the skull was 18.98 mm ± 0.613 (SD) and the mandible length was

14.41 mm ± 1.173 (SD). The total length of the baculum of a single S. kuhlii was 1.74 mm.

The head and body length of two Scotoecus pallidus was 56.50 mm ± 3.536 (SD). The forearm was 35.50 mm ± 0.707 (SD) long and the length of the tail was 35.50 mm ± 3.536 (SD).

The greatest length of skull was 15.46 mm ± 0.449 (SD) and mandible length was measured 9.64 mm ± 2.425 (SD). The total length of the baculum of a single S. pallidus captured from SA3 was

5.0 mm.

The mean head and body length of twenty two Pipistrellus ceylonicus was 63.60 mm ±

7.486 (SD). The length of forearm was 29.92 mm ± 2.492 (SD) and tail length was 25.68 mm ±

3.442 (SD). The greatest length of the skull was 10.76 mm ± 0.257 (SD) and the length of

mandible was 9.28 mm ± 3.956 (SD), respectively. Mean total length of the bacula of four P.

ceylonicus was 3.66 mm ± 1.190 (SD).

Mean head and body length of the ten P. javanicus was 52.00 mm ± 2.712 (SD). The

forearm was 35.13 mm ± 1.996 (SD) long and the length of the tail was 30.38 mm ± 5.236 (SD).

169

The greatest skull length was 13.01 mm ± 4.546 (SD) and the length of mandible was 10.29 mm

± 1.679 (SD). The mean total length of the four bacula was 3.57 mm ± 0.860 (SD).

The head and length of fifty two P. pipistrellus was 39.33 mm ± 2.690 (SD). The forearm was 28.23 mm ± 1.264 (SD) long and the length of the tail was 25.86 mm ± 3.396 (SD). The

greatest length of skull was 11.04 mm ± 0.342 (SD) and the length of mandible was 7.87 mm ±

0.802 (SD). The mean total length of the eleven bacula of P. pistrellus was 3.19 mm ± 0.421

(SD).

Only two specimens of P. tenuis were captured from SA1. The head and body length of

these specimens was 35.00 mm ± 2.828 (SD). The forearm length was 28.00±0.707 while the

length of the tail was 22.25 mm ± 3.182 (SD). The greatest length of the skull was 10.19 mm.

and the mandible length was 7.82 mm. The total bacular length was 2.79.

The head and body length of the two Hypsugo savii was 55.50 mm ± 19.092 (SD). The

forearm was 36.75 mm ± 3.889 (SD) long while the length of the tail was 33.50 mm ±6.364

(SD). The greatest length of the skull was 11.18 mm and the length of mandible was 7.08 mm.

The total bacular length of a single H. savii was 2.67 mm.

The echolocation calls of bats of Pakistan have never been recorded and thus the

accuracy of species identification on the basis of their calls remains a bit doubtful.

170

FUTURE RECOMMENDATIONS

1. Bat Surveys. Since bats are the least studied groups of mammals in Pakistan, extensive

bat surveys should be conducted to document as many bat species from the country as

possible. There are many bat species which have not been recorded after they were

originally described from the country for example Rhinopoma microphyllum (Brunnich

1782); Hipposideros fulvus Gray, 1838; H. cineraceus Blyth, 1853 and Pipistrellus kuhlii

(Kuhl, 1819). More bat surveys involving greater field efforts may confirm their presence

or complete absence from the habitats from where they were previously recorded.

2. Distribution Range and Species Specific Habitat Analysis. The Egyptian tomb bat

Taphozous perforatus has been captured from Attock for the first time. Capture of the

second county record from this area raises two scientific questions.

a. Whether the species inhabited this area but could not be recorded by earlier

surveyors? Or

b. Has the species extended its range from south to north?

Thus serious scientific studies should be conducted to redefine distribution of all the bat

species using global positioning system.

3. Reconfirmation of the Bat Taxonomy in Pakistan. Species form fundamental building

blocks of biological diversity as they provide goods and services to sustain life on earth.

Use of novel techniques in biology has made this era a new age of discoveries and new

species are being discovered each day with an unprecedented rate across broad taxonomic

and geographic spectra. Many new species of mammals, a group that was once

considered to be fully explored, have recently been described throughout the world.

These new mammalian species are described either from a previously poorly known

171

geographical area or they have arisen as a result of using molecular genetic techniques.

The later discoveries were made in such areas where geographic range of a well-known

mammal species was actually the combined ranges of two or more cryptic species—ones

not easily recognized by morphological features. Pakistan qualifies both the above

mentioned conditions. It is not only poorly surveyed part of the world but also owing to

its unique geographic position on the globe, many of the cryptic small mammal species

need to be reconfirmed taxonomically. The same species of Plecotus auritus and P.

austriacus can never exist in Pakistan and UK due to isolation of the two populations for

thousands of years and the answer to this question can only be found if molecular

genetics of the bats found in Pakistan is studied.

4. Bat Call Library. Bats have regional accents and the bat call for a single species

recorded from one region of the world do not match exactly to the calls of the same

species in other region of the world. Bats are so specialized that their calls vary with

respect to age and habitat. So a call library for the bats of Pakistan should be developed.

This will also save unnecessary killing of the bats for their taxonomic identification in the

country where specialized bat taxonomists are not available at the moment.

5. Bat conservation education for school children. Bats are generally considered

loathsome and are disliked by public in Pakistan. They are often associated with

witchcrafts and are thought to be responsible for transmitting diseases and entering in

human ears. It is highly difficult to shift the existing paradigm and convince people about

positive role of bats in ecosystem. It will take a lot of coordinated efforts to change the

existing mindset and turn them from “bat-foes” to “bat-friends”. The only way to

overcome this problem is to educate school children so that they should start admiring the

172

role of bats at tender age and should become stewards for bat conservation in the country.

So awareness raising campaigns for school children should continue as one of the

important next step.

173

LITERATURE CITED

Adams, A.L. (1867). Wanderings of a naturalist in India. Edmonston and Douglas, Edinburgh,

333 pp.

Ahmad, F. (1999). Ecological restoration in Cholistan. Journal Geographic, 2: 34-38.

Ahmad, Z. (2007). Country Report on Plant Genetic Resources for Food and Agriculture.

Pakistan Agricultural Research Council. 86 pp.

Ahmed, H. (1954). Wildlife in Sind. Pakistan Journal of Forestry, 4: 33-36.

Akbar, G., T.N. Khan and M. Arshad. (1996). Cholistan desert, Pakistan. Rangelands, 18: 124-

128.

Akbar, K.F., (1988). Phytosociological Studies of the Quaid-i-Azam University Campus,

lslamabad. M. Phil. Thesis.Department of Biological Sciences, Quaid-I-Azam

University, Islamabad.

Aksenova, T.G. (1980). Comparative morphological analysis of baculum construction in voles

from the tribe Microtini (Rodentia, Cricetidae). Communication I. Trudy

Zoologicheskogo Instituta Rossiskoy Akademii Nauk, 99: 62-77.

Albayrak, I. and N. Asan. (2001). The structure of baculum in Myotis myotis and Myotis blythi

(Chiroptera: Vespertilionidae) from Turkey. Turkish Journal of Zoology, 25: 229-

233.

Altringham, J. D. (1996). Bats: biology and behaviour. Oxford: Oxford University Press.

Anonymous. (1908). The imperial gazetteer of India. Clarendon Press, Oxford.

Anonymous. (1935). Punjab district gazetteer. Pt. B, Lyallpur district Government Printing,

Punjab, Lahore.

174

Anonymous. (1961). District census report, Lyallpur. Manager of Publications (West Pakistan),

Karachi.

Anonymous. (2010). Bats Conservation Times. Bats Conservation International. 8: 1 pp.

Arshad, M., G. Akbar and S. Rashid. (2003). Wealth of medicinal plants of Cholistan desert,

Pakistan: Conservational strategies. Hamdarad Medicus, 105: 25-34.

Asan, N, and I. Albayrak. (2007). Some Taxonomic features of Taphozous nudiventris

Cretzchmar, 1830 vel 1831 from Turkey (Chiroptera: Emballonuridae). Turkish

Journal of Zoology, 31: 165-170.

Ashraf, M.A., M.J. Maah1, I. Yusoff and K. Mehmood. (2010). Effects of Polluted Water

Irrigation on Environment and Health of People in Jamber, District Kasur, Pakistan.

International Journal of Basic and Applied Sciences, 10: 37-57.

Audet, D. and M.B. Fenton. (1988). Heterothermy and the use of torpor by the bat Eptesicus

fuscus (Chiroptera: Vespertilionidae): a field study. Physiological Zoology, 61:

197–204.

Awan, M.R., B.H. Niazi and Z. Khattak. (1992). Impact of soil on vegetation in Islamabad and

Rawalpindi Areas. Pakistan Journal of Agricultural Research, 13: 368-372.

Baig, M.S., M. Akram and M.A. Hassan. (1980). Possibilities for range development in

Cholistan desert as reflected by its physiography and soils. Pakistan Journal of

Forestry, 30: 61-71.

Baillie, J. and B. Groombridge. (1996). 1996 IUCN red list of threatened animals. IUCN, Gland,

Switzerland. 378 pp.

Barlow, K.E. and G. Jones. (1997a). Function of pipistrelle social calls: field data and a playback

Experimental Animal Behavior, 53: 991-999.

175

Barlow, K.E. and G. Jones. (1997b). Differences in songflight calls and social calls between two

phonic types of the vespertilionid bat Pipistrellus pipistrellus. Journal of Zoology

(London), 241: 315-324.

Baryshnikov, G.F. and A.V. Abramov. (1997). Structure of baculum (os penis) in Mustelidae

(Mammalia, Carnovora). Communication I. Zoologicheskii Zhurnal, 76: 1399-

1410.

Bass, M.S., M. Finer, C.N. Jenkins, H. Kreft and D.F. Cisneros-Heredia (2010). Global

conservation significance of Ecuador’s Yasuní National Park. PLoS ONE 5, e8767.

Bates, P. J. J. and D. L. Harrison. (1997). Bats of the Indian Subcontinent. Harrison Zoological

Museum. UK. 258 pp.

Bates, P., D. Thong and S. Bumrungsri. (2005). Voucher specimen preparation: bats. Harrison

Institute, England. Part of the Darwin Initiative Project: Taxonomic initiative for

Southeast Asian bat studies (Vietnam, Thailand, Cambodia and Lao PDR), 12 pp.

Bates, P.J.J., D.L. Harrison and M. Muni. (1994b). The bats of Western India. Part 3: Journal of

the Bombay Natural History Society, 91: 224-240.

Benda, P., and K.A. Tsytsulina. (2000). Taxonomic revision of Myotis mystacinus group

(Mammalia: Chiroptera) in the western Palearctic. Acta Societatis Zoologicae

Bohemicae, 64: 331-398.

Beolens, Michael Watkins & Michael Grayson (2009). "Vives". The Eponym Dictionary of

Mammals. Johns Hopkins University Press. pp. 432–433.

Betke, M., D.E. Hirsh, N.C. Makris, G.F. McCracken, and M. Procopio. (2008). Thermal

imaging reveals significantly smaller Brazilian free-tailed bat colonies than

previously estimated. Journal of Mammalogy 89, 18–24.

176

BirdLife International. (2011). Important Bird Areas factsheet: Ucchali Wetland Complex.

http://www.birdlife.org

Blanford, W.T. (1888). The fauna of British India, including Ceylon and Burma. Mammalia (2

vols.), Taylor and Francis, London.

Callery, H. (1951). Development of the os genital in the golden hamster, Mesocricetus (Cricetus)

auritus. Journal of Mammalogy, 32: 204-207.

Chaine, J. (1926). L’os penien, etude descriptive et comparative. Actes de la Societe Linneenne

de Bordeaux, 78: 5-195.

Champion, H.G., S.K. Seth and G.M. Khattak. (1965). Forest Types of Pakistan. Pakistan Forest

Institute, Peshawar.

Chaudhry, A. A, A.S. Malik and I. Ahmad. (1992). A survey of breeding birds in Lahore.

Proceedings Pakistan Congress of Zoology, 12: 383-394.

Chruszcz, B.J., R.M.R. Barclay. (2002). Thermoregulatory ecology of a solitary bat, Myotis

evotis, roosting in rock crevices. Functional Ecology, 16: 18–26.

Convention on Biological Diversity. (2009). Fourth National Report. Government of Pakistan.

Ministry of Environment, Islamabad. 97 pp.

Corbet, G.B. & Hill, J.E. (1992). A World List of Mammalian Species, Third edition. Natural

History Museum Publications & Oxford University Press, London and Oxford.: v-

viii, 1-243.

Daniel, B.A. (2009). Bat taxonomy and echolocation workshop for researchers at M.K.U. Small

Mammal Mail: Bi-annual Newsletter of CCINSA and RISCINSA 1(2).

Darlington, P.J. (1957). Zoogeography: The geographical distribution of animals. John Wiley &

Sons. New York, 675 pp.

177

Davidson-Watts, I., S. Walls and G. Jones. (2006). Differential habitat selection by Pipistrellus

pipistrellus and Pipistrellus pygmaeus identifies distinct conservation needs for

cryptic species of echolocating bats. Biological Conservation, 133: 118-127.

Davis, L. 2007. An Introduction to bats of United Arab Emirates. Echoes Ecology Ltd. UK.

www.echosecology.co.uk. pp. 24.

DeBlase, A.F. (1980). The bats of Iran: Systematics, distribution, ecology. Field Museum of

Natural History (Chicago, Ill).

Dietz, C. (2005). Illustrated identification key to bats of Egypt. Electronic Publication. Version

1.0. Germany.

Dietz, C. and O.V. Helversen. (2004). Identification key to the bats of Europe. 72 pp.

Electronical publication, version 1.0. Available at www.uni-tuebingen. de/tierphys/

Kontakt/ mitarbeiter_seiten/ dietz.htm.

Dildier, R. (1954). Etude systematique de l’os penis des mammiferes. Mammalia, 18: 237-256.

Dobson, G.E. (1876). Monograph of the Asiatic Chiroptera, and catalogue of the species of bats

in the collection of the Indian Museum, Calcutta Trustees of the Indian Museum.

London. 228 pp.

Douie, J.D. (1916). The Punjab, North-west Frontier Province and Kashmir. Cambridge

University Press, 373 pp.

Dunbar-Brander, A.A. (1927). Wild animals in central India. Edward Arnold and Co., London,

296 pp.

Ellerman, J.R. and T.C.S. Morrison-Scott. (1951). Checklist of Palearctic and Indian Mammals

1758 to 1946. Bristish Museum (Natural History), London. 810 pp.

178

Emmons, L. H. and F. Feer. (1997). Neotropical rainforest mammals: a field guide. 2nd edition.

The University of Chicago Press, Chicago.

Faiz-ur-Rehman. (2010). Inter population variations in the Indian flying fox (Pteropus

giganteus) of NWFP and Punjab-Pakistan. M. Phil. Thesis Department of Wildlife

and Ecology, University of Veterinary and Animal Sciences, Lahore, Pakistan.

Feng, J., S.Y. Zhang, Z.X. Li, L.X. Sheng and L.X. Wang. (2000). Echolocation calls of greater

horseshoe bat at different behavior. Acta Zoologica Sinica, 46: 230–232.

Fenton, M. (1996). Science and the conservation of bats. Journal of Mammalogy, 78/1: 1-14.

Fenton, M.B. (1992). The foraging behavior and ecology of animal-eating bats. Canadian Journal

of Zoology, 68: 411-422.

Findley, J.S. (1993). Bats: a community perspective. Cambridge University Press, Cambridge.

Freeman, P.W. (1988). Frugivorous and animalivorous bats (Microchiroptera): Dental and

cranial adaptations. Biological Journal of the Linnean Society, 33:249–272.

Friley, C.E. (1949). Age determination, by use of the baculum, in the river otter, Lutra

canadensis Schreber. Journal of Mammalogy, 30: 102-110.

Fujita, M. S. and M. D. Tuttle. (1991). Flying foxes (Chiroptera: Pteropodidea) - Threatened

animals of key ecological and economic importance. Conservation Biology, 4: 455-

463.

Fukui, D., N. Agetsuma and D.A. Hill. (2004). Acoustic identification of eight species of bat

(Mammalia: Chiroptera) inhabiting forests of southern Hokkaido, Japan: potential

for conservation monitoring. Zoological Science, 21: 947–955.

179

Geiser, F. (2003). Thermal biology and energetics of carnivorous marsupials. pp 234-249 in

Predators with Pouches: the Biology of Carnivorous Marsupials, edited by M.

Jones, C. Dickman and M. Archer. CSIRO Publishers, Melbourne, Australia.

Geiser, F. and G. Kortner. (2010). Hibernation and daily torpor in Australian mammals.

Australian Zoologist, 35: 204-215.

Geiser, F. and R.M. Brigham. (2000). Torpor, thermal biology, and energetics in Australian

long-eared bats (Nyctophilus). Journal of Comparative Physiology, 170: 153–162.

Gopalakrishna, A. and K. B. Karim. (1972). Arrangement of the foetal membranes and the

occurrence of haemodichorial placenta in the vespertilionid bat Pipistrelle

Pipistrellus mimus mimus. Current Science, 41: 144-146.

Gumal, M. T. and P. A. Racey. (1999). Legal protection of bats in Sarawak, Malaysia. Species.

25: 31-32.

Hagler Bailly Pakistan. (2007). Environmental Baseline Study of Margala and Margala North

Blocks. Final Report. 146 pp.

Harland H.J.D. (1970). Geological Survey water-supply. Paper, 269: 1473 pp.

Harris, A. H. (1974). Myotis yumanensis in interior southwestern North America with comments

on Myotis lucifugus. Journal of Mammalogy, 55: 589–607.

Harris, J. A. (1970). Bat-guano cave environment. Science. 169: 1342-1343.

Hatt, R.T. (1959). The mammals of Iraq. Miscellaneous Publications Museum of Zoology,

University of Michigan, 106: 1-113.

Heidt, G.A. (1970). The least weasel Mustela nivalis Linnaeus. Developmental biology in

comparison with other North American Mustela. Publications of the museum of the

Michigan State University, Biological Series, 4: 227-282.

180

Herr, A. and N.I. Klomp. (1995). A call exchange network for bat ecologists and fauna

managers: the South-eastern Australian bat call library. The Australasian Bat

Society Newsletter, 5: 11.

Hickey, B.C. and M.B. Fenton. (1996). Behavioural and thermoregulatory responses of female

hoary bats, Lasiurus cinereus (Chiroptera: Vespetilionidae), to variations in prey

availability. Ecoscience 3, 414–422.

Hijazi, S. (1984). A Phytosociolocical Study of Margalla Hills National Park, Islamabad. M.

Phil. Thesis. Quaid-i- Azam University, Islamabad.

Hill, J. and J. Smith. (1984). Bats: A Natural History. Austin: University of Texas Press.

Hill, J. E. and Smith J. D. (1985). Bats, a natural history. Texas University Press, Austin.

Hill, J.E. and D.L. Harrison. (1987). The baculum in Vespertilionidae (Chiropotera,

Vespertilionidae) with a systematic review, a synopsis of Pipistrellus and Eptesicus,

and the descriptions of a new genus and subgenus. Bulletin of the British Museum

(Natural History), Zoology, 52: 225-305.

Hinton, M.A.C. and O. Thomas. (1926). Report No. 42. Kashmir and Punjab. Bombay Natural

History Society’s Mammal Survey of India, Burma and Ceylon. Journal of Bombay

Natural History Society, 31: 606-614.

Hock, R.J. (1951). The metabolic rates and body temperatures of bats. Biology Bulletin, 101:

289–299.

Hosken, D.J., K.E. Jones, K. Chipperfield and A. Dixon. (2001). Is the bat os penis sexually

selected? Behavioral Ecology and Sociobiology, 50: 450-460.

181

Hristov, N.I., M. Betke, D.E. Hirsh, A. Bagchi, and T.H. Kunz. (2010). Seasonal variation in

colony size of Brazilian free-tailed bats at Carlsbad Caverns using thermal infrared

imaging. Journal of Mammalogy, 91: 183–192.

http://en.wikipedia.org/wiki/Kasur

http://www.fallingrain.com/world/PK/04/Gujranwala4.html

http://www.forumforfree.com/forums/index.php?mforum=forumpk&showtopic=3641

http://www.wildlifeofpakistan.com/national_parks.html

Hulbert, S.H. (1984). Pseudoreplication and the design of ecological field experiments.

Ecological Monograph, 54: 187-211.

Hussain. A. (2008). Impact of SCARP ponds on the birds and flora of Cholistan desert. Ph.D.

thesis Department of Zoology and Fisheries, University of Agriculture, Faisalabad.

283 pp.

Hutcheon, J.M. and J.A.W. Kirsch. (2006). A moveable face: deconstructing the microchiroptera

and new classification of extant bats. Acta Chiropterologica, 8: 1- 10.

Hutson, A.M., S.P. Mickleburgh and P.A. Racey. (2001). Microchiropteran bats: global status

survey and conservation action plan. IUCN/SSC Chiroptera Specialist Group.

IUCN, Gland, Switzerland and Cambridge, UK. x + 258 pp.

IUCN. (2008). IUCN Red List of Threatened Species. Version 2.8.1. www.iucnredlist.org.

IUCN. (2009). IUCN Red List of Threatened Species. Version 2009.1. www.iucnredlist.org.

Jacobs, D.S., G.N. Eick, M.C. Schoeman and C.A. Matthee. (2006). Cryptic species in an

insectivorous bat, Scotophilus dinganii. Journal of Mammalogy, 87(1):161–170.

Jerdon, T.C. (1867). The mammals of India. Thomason College Press, Roorke, XV+ 319 pp.

182

Jones, G. and E.C. Teeling. (2006). The evolution of echolocation in bats. Trends in Ecology and

Evolution,): 149-156.

Jones, G., and S.M. van Parijs. (1993). Bimodal echolocation in pipistrelle bats: are cryptic

species present? Proceedings of the Royal Society of London, 251: 119–125.

Jones, G., D.S. Jacobs, T.H. Kunz, M.R. Wilig and P.A. Racey. (2009). Carpe noctem: the

importance of bats as bioindicators. Endangered Species Research, 8: 93-115.

Jones, G., M. Morton, P.M. Hughes and R.M. Budden. (1993). Echolocation, flight morphology

and foraging strategies of some West African hipposiderid bats. Journal of Zoology

(London), 230: 385-400.

Jones, K., A. Purvis, and J. Gittleman. (2003). Biological correlates of extinction risk in bats.

American Naturalist, 161: 601–614.

Kalka, M.B., A.R. Smith and E.K.V. Kalko. (2008). Bats limit arthropods and herbivory in a

tropical forest. Science, 320: 71-73..

Kalko, E.K.V. (1997). Diversity in tropical bats. In: Ulrich H (ed.) Tropical Biodiversity and

Systematics, Proceedings of the International Symposium on Biodiversity and

Systematic in Tropical Ecosystems. Zoologisches Forschungsinstitut und Museum

Alexander Koenig, Bonn. pp: 13-43.

Kelm, D.H., K.R. Wiesner, O. von Helversen. (2008). Effects of artificial roosts for frugivorous

bats on seed dispersal in a Neotropical forest pasture mosaic. Conservation Biology,

22: 733–741.

Khan, A.U. (1994). History of decline and present status of natural tropical Thorn forest in

Punjab. Biological Conservation 67: 205–210.

183

Khattak, Z.D., and S. Ahmed. (1990). Phytosociological studies of the vegetation on the north

and south facing slopes of the Margalla Hills. Journal of Science and Technology,

University of Peshawar, 14: 129–132.

Koopman, K. F. (1993). Order Chiroptera (137 -241). In D. E. Wilson and D. M. Reeder (eds.)

Mammal Species of the World: A taxonomic and geographic reference. 2nd ed.

Smithsonian Institution Press, Washington, D.C., 137-241.

Kruskop, S.V. and L.A. Lavrenchenkov. (2000). A new species of long-eared bat

(Vespertilionidae, Chiroptera) from Ethiopia. Myotis, 38: 5-17.

Kulzer, E., Nelsen, J.E., McKean, J.L. and M. Ohres. (1970). Untersuchungen .uber die

Temperaturregulation australischer Flederm.ause (Microchiroptera). Z. Vergl.

Physiol. 69, 426 451.

Lane, D.J.W., T. Kingston, B.P.Y.H. Lee. (2006). Dramatic decline in bat species richness in

Singapore, with implications for Southeast Asia. Biological Conservation 131, 584–

593.

Lane, G. and S. Diver. (2000) Organic Greenhouse and Vegetable Production. Attra. 800: 346-

349.

Langley, E.A. (1860). Narrative of a residence at the court of Meer Ali Moorad, with wild sports

in the valley of the Indus. Hurst and Blackett, London, (2 vols.).

Lidicker, W.Z., Jr. (1968). A phylogeny of New Guinea rodent genera based on phallic

morphology. Journal of Mammalogy, 49: 609-643.

Lidicker, W.Z., Jr. and A. Yang. (1986). Morphology of the penis in the taiga vole (Microtus

xanthognathus). Journal of Mammalogy, 67: 497-502.

184

Lindsay, H.M. (1926). (i) Report No. 43: Nelliampathy palteau and Palni Hills (591-597); (ii)

Report No. 44: Kangra andChamba (597-606); (iii) Report No. 45: The Punjab Salt

Range and Murree (606-614); Bombay Natural History Society’s Mammal Survey

of India. Journal of the Bombay Natural History Society, 31: 591-614.

Lindsay, H.M. (1927). Kangra and Chamba, Bombay Natural History Society’s Mammal

Survey of India. Journal of the Bombay Natural History Society, 31: 597–607.

Lobova, T.A., C.K Geiselman and S.A. Mori. (2009). Seed Dispersal by Bats in the Neotropics.

New York Botanical Garden Press, New York.

Lydekker, R. (1924). The game animals of India, Burma, Malaya and Tibet. Rowland Ward,

Ltd., London, 412 pp.

Lyman, C.P. (1970). Thermoregulation and metabolism in bats. In: Wimsatt, W.A. (Ed.),

Biology of Bats. Academic Press, New York, pp. 301–330.

MacArdle, B.H. (1994). Multivariate analysis: a graduate student’s guide. Auckland: Department

of Statistics, University of Auckland.

Maeda, K. (1978). Baculum of the Japanese large noctule, Nyctalus lasiopterus aviator Thomas,

1911. Acta Anatomica Nipponica, 53: 447-453.

Maeda, K. (1982). Studies on the classification of Miniopterus in Eurasia, Australia and

Melanesia. Honyurui kagaku (Mammalian Science) Suppl. 1. Mammal Research

Association, Japan. pp: 1–176.

Mahmood, H. (2000). Some selected urban avian biodiversity of Lahore with special emphasis

on the species of concern. M.Sc. Thesis submitted to the Kinnaird College for

Women Lahore.

185

Mahmood-ul-Hassan M. and P.O. Nameer. (2006). Diversity, role and threats to the survival of

bats in Pakistan. Journal of Animal and Plant Sciences. 16: 38-42.

Mahmood-ul-Hassan M., M. G. Jones and C. Dietz. (2009). The Bats of Pakistan, the least

known creature. VDM. Verlag. Dr. Muller, Germany. 168 pp.

Mahmood-ul-Hassan, M., Faiz-ur-Rehman and M. Salim. (2011). Public perceptions about the

fruit bats in two horticulturally important districts of Pakistan. The Journal of

Animal and Plant Sciences, 21: 135-141.

Mahmood-ul-Hassan, M., T.L. Gulraiz, S.A. Rana and A. Javid. (2010). The diet of Indian

flying-foxes (Pteropus giganteus) in urban habitats of Pakistan. Acta

Chiropterologica, 12: 341–347.

Masood, N. (2004). Ecological Linkages of some sedentary bird species in selected urban

habitats of Lahore. Thesis submitted to the Kinnaird College for Women Lahore.

Masud, R.M. (1979). Master plan for Margallah Hills National Park, Islamabad, Pakistan.

National Council for Conservation of Wildlife, Islamabad. 48 pp.

Meyer, C.F.J., L.M.S. Aguiar, L.F. Aguirre, J. Baumgarten, F.M. Clarke, J. Cosson, S.E.

Villegas, J. Fahr, D. Faria, N. Furey, M. Henry, R. Hodgkison, R.K.B. Jenkins,

K.G. Jung, T. Kingston, T.H. Kunz, M.C.M. Gonzalez, I. Moya, J. Pons, P.A.

Racey, K. Rex, E. M. Sampaio, K.E. Stoner, C.C. Voigt, D. von Staden, C.D. Weise

and E.K.V. Kalko. (2010). Long-term monitoring of tropical bats for anthropogenic

impact assessment: Gauging the statistical power to detect population change.

Biological Conservation, 43: 2797-2807.

186

Mickleburgh, S. P., A. M. Hutson and P.A. Racey. (1992). Old world fruit bats. An action plan

for their conservation. IUCN/SSC Chiroptera Specialist Group. IUCN, Gland,

Switzerland. pp: 1-16.

Moeller, H. (1985). Cladus Vertebrata. In R. Siewing (ed). Lehrbuch der Zoologie. Band 2,

Systematik, Fischer, Stuttgart: 362-554.

Mohammad, S. (2009). Habitat, abundance, morphology and general biology of Indian flying fox

(Pteropus giganteus) at Dir, Malakand, Mardan and Charsadda districts of NWFP–

Pakistan. M. Phil. Thesis Department of Wildlife and Ecology, University of

Veterinary and Animal Sciences, Lahore, Pakistan.

Moreno, C. E. and G. Halffter. (2001). Spatial and temporal analysis of α, β and γ diversities of

bats in a fragmented landscape. Biodiversity Conservation 10:367–382.

Murray, J.A. (1884). The vertebrate zoology of Sind. Richardson and Co., London, 424 pp.

Naeem, M., K. Khan, S. Rehman and J. Iqbal. (2007). Environmental assessment of ground

water quality of Lahore area, Punjab, Pakistan. Journal of Applied Sciences, 7: 41-

46.

Nagorsen, D.W. and R.M. Brigham. (1993). The Bats of British Columbia. UBC Press,

Vancouver, British Columbia, 164 pp.

Nawaz, A., Z.B. Mirza, V. Zakaria, M. Rafiq, M. Shah, R. Malik, N.K. Khan and M. Younas.

(2007). Ecological Baseline of Margalla Hills National Park. A report published by

Capital Development Authority (CDA) and Himalayan Wildlife Foundation,

Islamabad. Document Ref: D7BL1MHP. 129 pp.

O’Farrell, M.J. (1997). Use of echolocation calls for the identification of free-flying bats.

Transactions of the Western Section Wildlife Society, 33: 1-8.

187

O’Shea, T.J. and M.A. Bogan. (2003). Monitoring Trends in Bat Populations of the United States

and Territories: Problems and Prospects. US Geological Survey, Biological

Resources Division, Information and Technology Report, USGS/BRD/ITR–2003-

0003.

Park, K.J., G. Jones and R.D. Ransome. (2000). Torpor, arousal and activity of hibernating

Greater Horseshoe Bats (Rhinolophus ferrumequinum). Functional Ecology, 14:

580–588.

Parker, R.N. (1924). A forest flora of the Punjab, with Hazard and Delhi. Government Printing,

Punjab, Lahore, XXX + 501 pp.

Parsons, S. and G. Jones. (2000). Acoustic identification of twelve species of echolocating bat by

discriminant function analysis and artificial neural networks. Journal of

Experimental Biology, 203: 2641-2656.

Pereswiet-Soltan, A. (2007). Relation between climate and bat fauna in Europe. Travaux du

Museum National d’Histoire Naturelle, Grigore Antipa. L: 505-515.

Petter, F. (1961). Repartition geographique et ecologie des rongeurs desertiques (du Dahara

occidental a l Iran oriental). Mammalia, 25: 1-219

Pocock, R.I. (1939). The fauna of British India including Ceylon and Burma. Mammals (2 vols.,

Primates and Carnivora only), Taylor and Francis, London.

Pottie, S.A., D.J.W. Lane, T. Kingston and B.P.Y.H. Lee. (2005). The microchiropteran bat

fauna of Singapore. Acta Chiropterologica, 7: 237–247.

Prakash, B. (1939). Animal remains from Harappa. Memorial Museum of Archaeology Survey

of India, 51: 1-62.

188

Prater, S.H. (1947). The book of Indian animals (mammals). Bombay Nat. Hist. Soc., Bombay,

263 pp.

Purohit, A. and K.B. Vyas. (2006). Review of sex ratio in several bat species inhabiting the Great

Indian Desert. Vespertilio, (9–10): 233–235.

Purohit, A. and K.R. Senacha. (2004). Distribution of bats in and around Jaisalmer of the Great

Indian Desert, India. Vespertilio, 8: 99-104.

Qadir A, R.N. Malik and S.Z. Hussain. (2008). Spatio-temporal variations in water quality of

Nullah Aik-tributary of the river Chenab, Pakistan. Environmental Monitoring and

Assessment, 140: 43-59.

Rabeder, G. (1976). Die Carnivoran (Mammalia) aus dem Altpleistozan von Deutsch-Altenburg

2. Mitbeitrage zur Palaontologie von Osterreich, 1: 5-119.

Ramirez-Pulido, J., J. Arroyo-Cabrales and A. Castro-Campillo. (2005). Estado actual y relación

nomenclatural de los mamíferos de México. Acta Zoologica Mexicana, 21: 21–82.

Roberts, T.J. (1977). The mammals of Pakistan. Ernest Benn Limited, 361 pp.

Roberts, T.J. (1997). Mammals of Pakistan. Revised Ed. Oxford Univ. Press. Oxford.

Russo, D. and G. Jones. (1999). The social calls of Kuhl’s pipistrelles Pipistrellus kohli (Kuhl,

1819): structure and variation (Chiroptera: Vespertilionidae) Journal of Zoology

(London), 249: 476-481.

Russo, D. and G. Jones. (2000). The two cryptic species of Pipistrellus pipistrellus (Chiroptera:

Vespertilionidae) occur in Italy: evidence from echolocation and social calls.

Mammalia, 64: 187-197.

189

Russo, D. and G. Jones. (2002). Identification of twenty-two bat species (Mammalia: Chiroptera)

from Italy by analysis of time-expanded recordings of echolocation calls. Journal of

Zoology (London), 258: 91-103.

Russo, I.R.M., C.T. Chimimba and P. Bloomer. (2006). Mitochondrial DNA differentiation

between two species of Aethomys (Rodentia: Muridae) from southern Africa.

Journal of Mammalogy, 87: 545–553.

Rydell, J., H.T. Arita, M. Santos and J. Granados. (2002). Acoustic identification of

insectivorous bats (order Chiroptera) of Yucatan, Mexico. Journal of Zoology, 257:

27–36.

Said, R.M. (1956). Working Plan for the forests of the Jhelum, Mianwali, and Shahpur Forest

divisions from 1952-53 to 1981-82. Vol-1. Government printing, Lahore.

Schipper, J., J.S. Chanson, F. Chiozza, N.A. Cox, M. Hoffmann, and V. Katariya. (2008). The

status of the World’s land and marine mammals: diversity, threat, and knowledge.

Science, 322: 225–230.

Sergey, P. (2001). Adsorption of Cadmium onto Hematite: Temperature Dependence. Journal of

Colloidal and Interface Science, 234: 1-8.

Sheikh, K.M. and S. Molur. (2004). Status and Red List of Pakistan’s Mammals. Based on the

Conservation Assessment and Management Plan. IUCN Pakistan 312 pp.

Sheikh, M.I. (1987). Forests and Forestry in Pakistan. Pakistan Forest Institute, Peshawar,

Pakistan.

Sheikh, M.I. (1993). Trees of Pakistan. Pictorial Printers (Pvt.) Ltd., Islamabad.

Shinwari, M.I. and M.A. Khan. (2000). Folk use of medicinal herbs of Margalla Hills National

Park, Islamabad. Journal of Ethnopharmacology, 69: 45–56

190

Shinwari, Z.K., B.A. Khan, and A.A. Khan. (1996). Proceedings of the First Training Workshop

on Ethnobotany and its application to conservation. National Herbarium: PASA

National Agricultural Research Centre, Islamabad, Pakistan, September 16–21,

1996.

Siddiqi, M.S. (1961). Checklist of mammals of Pakistan with particular reference to the

mammalian collection in the British Museum (Natural History), London. Biologia,

7: 93-225.

Simmons, N.B. (2005a). Chiroptera. In: The Rise of Placental Mammals, K. D. Rose and J. D.

Archibald, eds., pp. 159–174, Johns Hopkins University Press, Baltimore.

Simmons, N.B. (2005b). Order Chiroptera. In: Mammal Species of the World: A Taxonomic and

Geographic Reference, D.E. Wilson and D.M. Reeder, eds., Smithsonian Institution

Press, Washington, DC.

Simpson, G.G., A. Roe and R.C. Lewontin. (1960). Quantitative zoology. Revised ed. Harcourt,

Brace and World, Inc. New York, 440 pp.

Simpson, G.R. (1996). Prevalence of Kaposi’s sarcoma-associated herpesvirus infection

measured by antibodies to recombinant capsid protein and latent

immunofluorescence antigen. Lancet, 348, 1133–1138.

Singaravelan, N., G. Marimuthu and P.A. Racey. (2009). Do fruit bats deserve to be listed as

vermin in the Indian Wildlife (Protection) and amended acts? A critical review.

Oryx 43: 608–613.

Sinha, Y.P. (1980). The bats of Rajasthan: taxonomy and zoogeography. Records of Zoological

Survey of India, 76: 7-63.

191

Sinha, Y.P. (1986). The bats of Bihar: Taxonomy and Field Ecology. Records of the Zoological

Survey of India. Occasional Papers, 77: 1-60.

Smirnov, D.G. (2000). Variation in the baculum of bats (Chiroptera, Vespertilionidae) from the

middle Volga Basin and the adjacent territories. Plecotus, 3: 20-34.

Speakman, J.R. and D.W. Thomas. (2003). Physiological ecology and energetics of bats. in:

Kunz, T.H., Fenton, M.B. (Eds.), Ecology of Bats II. Academic Press, New York, in

press.

Srinivasulu, C., P.A. Racey and S. Mistry. (2010). A key to the bats (Mammalia: Chiroptera) of

South Asia. Journal of Threatened Taxa, 2: 1001-1076.

Strelkov, P.P. (1989a). New data on the structure of baculum in Palaearctic bats I. The genera

Myotis, Plecotus and Barbastella. pp. 87-94 in European bat research 1987 (V.I.

Hanak, I. Horacek, and J. Gaisler, Eds.). Charles University Press, Praha, 420 pp.

Strelkov, P.P. (1989b). New data on the structure of baculum in Palaearctic bats II. Genus

Eptesicus pp: 95-100 in European bat research 1987 (V.I. Hanak, I. Horacek, and J.

Gaisler, eds.). Charles University Press, Praha, 420 pp.

Sun, K., J. Feng, L. Jin, Y. Liu and Y. Jiang. (2008). Identification of sympatric bat species by

the echolocation calls. Frontiers of Biology, 3: 227–231.

Taber, R.D., A.N. Sheri and M.S. Ahmad. (1967). Mammals of the Lyallpur region, West

Pakistan. Journal of Mammaolgy, 48: 392-407.

Tarasov, S.A. (1974). Age peculiarities of baculum shape in some mammals. Zoologicheskii

Zhurnal, 53: 1109-1111.

Tarasov, S.A. (1984). Postnatal development of baculum in some voles (Rodentia, Cricetidae).

Zoologicheskii Zhurnal, 53: 1109-1111.

192

Tauseef-ur-Rehman, M.N. Khan, M.S. Sajid, R.Z. Abbas, M. Arshad, Z. Iqbal and A. Iqbal.

(2011). Epidemiology of Eimeria and associated risk factors in cattle of district

Toba Tek Singh, Pakistan. Parasitology Research, 108:1171–1177.

Tauseef-Ur-Rehman, M.N. Khan, M.S. Sajid, R.Z. Abbas, M. Arshad, Z. Iqbal and A. Iqbal.

(2011). Epidemiology of Eimeria and associated risk factors in cattle of district

Toba Tek Singh, Pakistan. Parasitological Research, 108: 1171–1177.

Thomas, O. (1915). The penis bone, or ‘baculum’ as a guide to the classification of certain

squirrels. Annals and species. Annali del Museo Civico di Storia Naturale de

Genova (Ser. 2), 9: 86-88.

Tian, L., B. Liang, K. Maeda, W. Metzner and S. Zhang. (2004). Molecular studies on the

classification of Miniopterus schreibersii (Chiroptera: Vespertilionidae) inferred

from mitochondrial cytochrome b sequences. Folia Zoology, 53: 303–311.

Topal, G. (1958). Morphological studies on the os penis of bats in the Carpathian basin. Annales

Historico-Naturales Musei Nationalis Hungarici (S.N. 9), 50: 331-342.

Tsytsulina, K. (2001). Myotis ikonnikovi (Chiroptera, Vespertilionidae) and its relationships with

similar species. Acta Chiropterologica, 3: 11-19.

Turbill, C., K.G. Ortner, and F. Geiser. (2001). Patterns of torpor use by free-ranging Australian

forest bats. Abstracts 12th International Bat Research Conference. Faculty of

Science and Technology, Universiti Kebangsaan Malaysia, Selangor, Malaysia, pp.

56–57.

Ullah, R., R.N. Malik and A. Qadir. (2009). Assessment of groundwater contamination in an

industrial city, Sialkot, Pakistan. African Journal of Environmental Science and

Technology, 3: 429-446.

193

Vaughan, N., G. Jones and S. Harris. (1996). Effects of sewage effluent on the activity of bats

(Chiroptera: Vespertilionidae) foraging along rivers. Biological Conservation, 78:

337–343.

Vaughan, N., G. Jones and S. Harris. (1997). Identification of British bat species by multivariate

analysis of echolocation parameters. Bioacoustics, 7: 189-207.

Vaughan, N., G. Jones and S. Harris. (1997b). Identification of British bat species by

multivariate analysis of echolocation call parameters. Bioacoustics, 7: 189–207.

th Vaughan, T., J. Ryan, and N. Czaplewski. (2000). Mammalogy. 4 Edition. Toronto: Brooks

Cole.

Verts, B.J. and L.N. Carraway. (1998). Land mammals of Oregon. University of California

Press, Berkeley, California, 668 pp.

von Helversen, O. and Y. Winter. (2003). Glossophagine bats and their flowers: costs and

benefits for plants and pollinators. In: Kunz, T.H., Fenton, M.B. (Eds.), Bat

Ecology. University of Chicago Press, Chicago, 346–397 pp.

Walker, E.P. (1964). Mammals of the World. Vol. 2. Johns Hopkins University Press, Baltimore,

MD, USA.

Walker, S. and S. Molur. (2003). Summary of the status of South Asian Chiroptera. Extracted

from C.A.M.P. 2002. Report. Zoo Outreach Organization. CBSG South Asia and

Wild, Coimbatore, India.

Walsh, A.L., C.M.C. Catto, T.M. Hutson, S. Langton and P.A. Racey. (2003). The United

Kingdom Bat Monitoring Programme: Turning conservation goals into tangible

results. In: O’Shea, T.J., Bogan, M.A. (Eds.), Monitoring Trends in Bat Populations

of the United States and Territories: Problems and Prospects. US Geological

194

Survey, Biological Resources Division, Information and Technology Report,

USGS/BRD/ITR–2003-0003, pp. 103–118.

Walton, D.W. (1974). New records of bats (Chiroptera) from Pakistan. Journal of the

Mammalogical Society of Japan, 6: 43-50.

Weller, T.J., S.A. Scott, T.J. Rodhouse, P.C. Ormsbee and J.M. Zinck. (2007). Field

identification of the cryptic vespertilionid bats, Myotis lucifugus and M.

yumanensis. Acta Chiropterologica, 9(1): 133–147.

Williams-Guillén, K., I. Perfecto and J. Vandermeer. (2008). Bats limit insects in a Neotropical

agroforestry system. Science, 320: 71 pp.

Wilson, D.E. and D.M. Reeder. (1993). Mammals species of the world, a taxonomic and

geographic reference. 2nd edition. Smithsonian Institution Press, Washington, D.C.

Wilson, D.E. and D.M. Reeder. (2005) Mammal Species of the World: A Taxonomic and

Geographic Reference. Third Edition. The Johns Hopkins University Press,

Baltimore.

World Gazetteer. (2010). Pakistan: largest cities and towns and statistics of their population.

http://world-gazetteer.com/wg.php?x=&men=gcis&lng=en&des=wg&geo=-

172&srt=npan&col=abcdefghinoq&msz=1500&pt=c&va=&srt=pnan

Wroughton, R.C. (1916). Report No 20: Chindwin River. Bombay Natural History Society’s

Mammal Survey of India, Burma and Ceylon. Journal of the Bombay Natural

History Society, 24:291–316.

WWF – Pakistan. (2008). The Study of Ecology and Ecological Linkages of the Lahore Canal

Bank from Mustafaabad Bridge to Thokar Niaz Baig. Report by World Wide Fund

for Nature-Pakistan. 39 pp.

195

Yoon, M.H., K. Ando and T.A. Uchida. (1990). Taxonomic validity of scientific names in

Japanese Vespertilio species by ontogenetic evidence of the penile pseudobaculum.

Journal of Mammalogical Society of Japan, 14: 119-128.

Yoshiyuk, M. (1989). A systematic study of the Japanese Chiroptera. National Science Museum,

Tokyo, 242 pp.

Zhang, S.Y., H.H. Zhao, J. Feng, L.X. Sheng, H. Wang and L.X. Wang. (2000). Relationship

between echolocation frequency and body size in two species of Hipposideros.

Chinese Science Bulletin, 45: 740–743.

196