Systematic Study of Scarabaeidae

Muhammad Abbas

Department of Microbiology Faculty of Biological Sciences Quaid-i-Azam University Islamabad 2015

A thesis submitted in partial fulfillment of the requirements for the Degree of

Doctor of Philosophy

Microbiology

Muhammad Abbas

Department of Microbiology Faculty of Biological Sciences Quaid-i-Azam University Islamabad 2015

TABLE OF CONTENTS

Acknowledgements i Originality statement ii Certificate iii List of abbreviations iv List of figures vi Abstract 1 CHAPTER 1 INTRODUCTION AND REVIEW OF LITERATURE

1.1 Historical geology 2 1.2 Zoogeography 2 1.3 Physiography 2 1.4 Climate 3 1.5 Eco-regions 3 1.6 Taxonomy 4 1.7 Distribution 5 1.8 General appearance 6 1.9 Significance 8 1.10 Scarabaeinae studies in Pakistan 14 1.11. Objectives 15

CHAPTER 2 MATERIALS AND METHODS

2.1. Study area 16 2.2 Collection of scarab beetles 16 2.3 Sampling areas 16 2.3.1. A) Alpine zone 18 2.3.1.1. Alpine meadows 18 2.3.1.2. Sub-alpine scrub and birch forest 18 2.3.1.3. Alpine dry steppe 18 2.3.2 B) Montane temperate forest 19

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2.3.2.1. sub-tropical pine forest 19 2.3.2.2. Himalayan moist temperate forest 19 2.3.2.3. Himalayan dry temperate forest 19 2.3.3. C) Tropical deciduous forest 19 2.4. Collection methods 20 2.5. Identification 21

CHAPTER 3 RESULTS AND DISCUSSIONS

3.1 Subfamily SCARABAEINAE Latreille, 1802 23 3.2 Genus HELIOCOPRIS Hope, 1837 24 3.2.1 Key to the species of HELIOCOPRIS Hope, 1837 24 3.2.2 Heliocopris midas (Fabricius, 1775) 24 3.2.3 Heliocopris midas (Fabricius, 1775) 28 3.3 Genus DIGITONTHOPHAGUS Balthasar, 1959 31 3.3.1 Key to species of Digitonthophagus Balthasar, 1959 31 3.3.2 Digitonthophagus gazella (Fabricius, 1787) 31 3.3.4 Digitonthophagus bonasus (Fabricius, 1798) 35 3.4 Genus ONTHOPHAGUS Latreille 1802 38 3.4.1 Onthophagus (Palaeonthophagu) marginalis (Gebler, 1817) 38 3.4.2 Onthophagus (Palaeonthophagus) nuchicornis (Linnaeus, 1758) 42 3.4.3 Onthophagus (Phanaeomorphus) gagates Hope, 1831 46 3.4.4 Onthophagus (Altonthophagus) kukunorensis Kabakov, 1990 49 3.4.5 Onthophagus (Coloboonthophagus) tragus Fabricius, 1792 52 3.4.6 Onthophagus (s. str.) proletarius Harold, 1875 56 3.5 Genus COPRIS Geoffroy, 1762 59 3.5.1 Key to the species of Genus Copris 59 3.5.2 Copris repertus Walker, 1858 60 3.5.3 Copris sacontala Redtenbacher, 1848 63 3.5.4 Copris punctatus Gillet, 1910 66 3.6 Identification key to the collected genera of tribe Onitini 69 3.6.1 Genus ONITIS Fabricius, 1798 69 3.6.2 Key to species of Genus Onitis 69

3.6.3 Onitis falcatus (Wulfen, 1786) 71 3.6.4 Onitis lama van Lansberge 1875 74 3.6.5 Onitis philemon Fabricius 1801 77 3.6.6 Onitis subopacus Arrow, 1931 80 3.6.7 Onitis virens Lansberge, 1875 84 3.7 Genus CHEIRONITIS Lansberge, 1875 88 3.7.1 Cheironitis arrowi Janssens, 1937 88 3.8 Genus CATHARSIUS Hope, 1837 92 3.8.1 Keys Species of Genus CATHARSIUS Hope, 1837 92 3.8.2 Catharsius pithecius (Fabricius, 1775) 92 3.8.3 Catharsius molossus (Linnaeus, 1758) 96 3.9 Genus METACATHARSIUS Laporte, 1840 100 3.9.1 Metacatharsius inermis (Laporte, 1840) 100 3.10 Key to the collected genera of tribe Scarabaeini 103 3.10.1 Genus GYMNOPLEURUS Illiger 1803 103 3.10.2 Gymnopleurus flagellatus (Fabricius, 1787) 103 3.10.3 Scarabaeus sacer Linnaeus, 1758 107 3.11 List of the species 111

CHAPTER 4 CONCLUSION AND RECOMMENDATIONS

4.1 Conclusion and recommendations 112

CHAPTER 5 REFERENCES

5.1 References 114

5.2 Web References 135

5.3 Publication (International) 135

ACKNOWLEDGEMENTS

All the acclamations and appreciations are for ALMGHTY ALLAH “Who taught Adam names of all things” and “He taught man that which he knew not”.

The most influential persons in the successful completion of practical work and preparation of manuscript is my supervisor, Prof. Dr. Najma Ayub, Department of Microbiology, Quaid- i-Azam, University, Islamabad and Co-supervisor Dr. Syed Azher Hasan, Ex. Chairman, Pakistan Science Foundation, Islamabad. They Provide me guidance & support throughout my studies

I could not have completed this task without the help and guidance of Dr. Fariha Hasan, Chairpersan, Department of Microbiology and Prof. Dr. Wasim Ahmad Dean, faculty of Biological Sciences. Their guidance and encourgment supported me to face every challenge.

I feel pride in expressing my profound and cordial gratitude to Prof. Xingke Yang and Dr. Ming Bai, Institute of Zoology, Chinese Academy of Sciences (IOZ, CAS) Beijing for letting me to do work in their lab and their technical advice, constant help, keen interest for the completion of this task. It was because of their indulgent help and advice that I have been able to accomplish this study.

I am also thankful to the Higher Education Commission for providing me the opportunity to carry out this research and advanced literature from the digital library source.

There are many other people who helped me and I would like to say that without their support it was impossible to achieve this goal. I would like to express special thanks to Bruce Gill, Siqin, Xue, Zaheer, Yang, Wang, Yang Yang, Ishtiaq, Rafaqat Masroor, Mirza Habib, Shabir Amir and Ahsan Feroze I would also like to thank all my colleagues who helped me for the completion of this project.

It is difficult to express in words the support and encouragement of my family for their love, inspiration, well wishes, cooperation and keen interest, which hearten me to achieve success in every sphere of life.

MUHAMMAD ABBAS

ORIGINALITY STATEMEMNT

I hereby declare that this submission is my own work and I have not previously presented any part of this work elsewhere for any other degree

MUHAMMAD ABBAS

CERTIFICATE

This thesis submitted by Muhammad Abbas is accepted in its present form by the Department of Microbiology, Quaid-i-Azam University, and Islamabad, Pakistan; as satisfying the thesis requirements for the degree of Doctor of Philosophy in Microbiology.

Supervisor: ______Professor Dr. Najma Ayub

Co Supervisor: ______Dr. Syed Azhar Hasan Ex-Chairman Pakistan Science Foundation

External Examiner-1: ______Dr. Kishwar Nazir HEC Distinguished Professor Department of Plant Pathology University of Arid Agriculture, Rawalpindi

External Examiner-2: ______Dr. Ghazala Kaukab Associate Professor Department of Biochemistry University of Arid Agriculture, Rawalpindi

Chairperson: ______Dr. Fariha Hasan

Dated: ______

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LIST OF ABRIVIATIONS

ANLW Austria, Wien（Vienna, Amt Der Niederösterreichischen Landsregierung BJMNH China, Beijing, Beijing Museum of Natural History BMNH United Kingdom, London, The Natural History Museum [formerly British Museum of Natural History CABI: Centre for Agricultural Bioscience International Rawalpindi CAS Chinese Academy of Sciences HEC Higher Education Commission ISNB Belgium, Brussels, Institut Royal des Sciences Naturelles de Belgique IZAS China, Beijing, Institute of Zoology, Chinese Academy of Sciences Km kilometer KPK Khyber Pakhtunkhwa LSLC United Kingdom, London, Linnean Society of London, Linnaean Collections MCSN Italy, Genova, Museo Civico di Storia Naturale "Giacomo Doria" MNHN: Muséum National d'Histoire Naturelle, Paris, France NARC National Agriculture Research Council, Islamabad NHML: The Natural History Museum, London, UK NIM: National Insect Museum, Islamabad NMW Austria, Wien, Naturhistorisches Museum Wien PMNH Pakistan, Islamabad, Pakistan Museum of Natural History SNMB Germany, Braunschweig, Staatliches Naturhistorisches Museum Spp species UZIU Sweden, Uppsala, Uppsala University ZIN Russia, St. Petersburg, Russian Academy of Sciences, Zoological Institute ZMUC Denmark, København [= Copenhagen], University of Copenhagen, Zoological Museum ZMUK Germany, Kiel, Universität Kiel, Zoologisches Museum ZMUM Russia, Moscow, Moscow State University ZMUC: Denmark, København [= Copenhagen], University of Copenhagen, Zoological Museum

LIST OF FIGURES

Fig. 1. Dorsal view of scarab beetle (Ratcliffe, 1991) 6

Fig. 2. Ventral view of scarab beetle (Ratcliffe, 1991) 7

Fig. 3. Administrative map of the study area 17

Fig. 4. a) Pitfall trap, b) beetles floating on the water surface in the bucket, 20 c) light trap, and d) hand-picking of beetles with forceps

Fig. 5. a) Fixation of scarab beetles and b) catalog numbering 21

Fig. 6.1. Heliocopris midas 26

Fig. 6.2. Distribution map of Heliocopris midas 27

Fig. 7.1. Heliocopris bucephalus 29

Fig. 7.2. Distribution map of Heliocopris bucephalus 30

Fig. 8.1. Digitonthophagus gazella 33

Fig. 8.2. Distribution map of Digitonthophagus gazella 34

Fig. 9.1. Digitonthophagus bonasus 36

Fig. 9.2. Distribution map of Digitonthophagus bonasus 37

Fig. 10.1. Onthophagus marginalis 40

Fig. 10.2. Distribution map of Onthophagus marginalis 41

Fig. 11.1. Onthophagus nuchicornis 44

Fig. 11.2. Distribution map of Onthophagus nuchicornis 45

Fig. 12.1. Onthophagus gagates 47

Fig. 12.2. Distribution map of Onthophagus gagates 48

Fig. 13.1. Onthophagus kukunorensis 50

Fig. 13.2. Distribution map of Onthophagus kukunorensis 51

Fig. 14.1. Onthophagus tragus 54

Fig. 14.2. Distribution map of Onthophagus tragus 55

Fig. 15.1. Onthophagus proletarius 57

Fig. 15.2. Distribution map of Onthophagus proletarius 58

Fig. 16.1. Copris repertus 61

Fig. 16.2. Distribution map of Copris repertus 62

Fig. 17.1. Copris sacontala 64

Fig. 17.2. Distribution map of Copris sacontala 65

Fig. 18.1. Copris punctatus 67

Fig. 18.2. Distribution map of Copris punctatus 68

Fig. 19.1. Onitis falcatus 72

Fig. 19.2. Distribution map of Onitis falcatus 73

Fig. 20.1. Onitis lama 75

Fig. 20.2. Distribution map of Onitis lama 76

Fig. 21.1. Onitis philemon 78

Fig. 21.2. Distribution map of Onitis philemon 79

Fig. 22.1. Onitis subopacus 82

Fig. 22.2. Distribution of Onitis subopacus 83

Fig. 23.1. Onitis virens 86

Fig. 23.2. Distribution of Onitis virens 87

Fig. 24.1. Cheironitis arrowi 90

Fig. 24.2. Distribution map of Cheironitis arrowi 91

Fig. 25.1. Catharsius pithecius 94

Fig. 25.2. Distribution map of Catharsius pithecius 95

Fig. 26.1. Catharsius molossus 98

Fig. 26.2. Distribution map of Catharsius molossus 99

Fig. 27.1. Metacatharsius inermis 101

Fig. 27.2. Distribution map of Metacatharsius inermis 102

Fig. 28.1. Gymnopleurus flagellatus 105

Fig. 28.2. Distribution map of Gymnopleurus flagellates 106

Fig. 29.1. Scarabaeus sacer 109

Fig. 29.2. Distribution map of Scarabaeus sacer 110

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ABSTRACT

The present studies were carried out to investigate the dung beetles‟ fauna in the selected parts of northern Pakistan including Khyber Pakhtunkhwa province (KPK) and Federally Administered Tribal Areas (FATA). The area is diverse and virgin for its flora and fauna pertaining to different habitats, especially the dung beetles in subfamily scarabaeinae. The study was conducted in the three selected ecological zones including alpine zone, montane temperate forest and tropical deciduous forest. These zones were further divided into sub-zones for comprehensive collection of dung beetles fauna.

The systematic study revealed the presence of ten genera and twenty-four species from the study area including Digitonthophagus gazelle, D. bonasus, Heliocopris midas, H. bucephalus, Onthophagus (Phanaeomorphus) gagates, O. (Altonthophagus) kukunorensis, O. (Palaeonthophagus) marginalis, O. (Palaeonthophagus) nuchicornis, O. (s. str.) proletarius, O. (Coloboonthophagus) tragus, Onitis falcatus, O. lama, O. philemon, O. subopacus, O. virens, Catharsius molossus, C. pithecius, Metacatharsius inermis, Copris repertus , C. sacontala, C. punctatus, Scarabaeus sacer, Cheironitis arrowi and Gymnopleurus flagellates. Out of these, eights species were newly recorded for Pakistan including Onthophagus gagates, Onthophagus kukunorensis, O. (s. str.) proletarius, He- liocopris bucephalus, Copris punctatus, C. repertus, C. sacontala and Onitis lama. The diagnosis of each genus, identification keys to genera and species, colored photographs and geographical distribution maps for each species along with scale length size were provided.

1. INTRODUCTION AND REVIEW OF LITERATURE

1.1 Historical geology

The Indo-Pakistan subcontinent occupies a peculiar geographical position in Asia. It was originally a part of eastern Gondwana, the southern supercontinent, which fragmented into several plates in the remote past: Indian, Africa, South America, etc. According to the theory of plate tectonics, the plates moved across ancient oceans to their present position (King, 1967: Courtillot, 1983; Sengör, 1985). The Indian plate moved across the In- dian Ocean, to abut against the Asian land mass. In 1947, the subcontinent was parti- tioned politically into two sovereign states, the northwestern and eastern part as Pakistan and rest of the subcontinent as India.

1.2 Zoogeography

Pakistan is the most interesting region biogeographically due to its status as transitional zone between the Palearctic and Oriental regions, and its enormous variety in ecosystems, from desert through forests to alpine meadows and cold deserts. Pakistan covers approximately an area of 796, 095 km2, sandwiched between two main zoogeographical regions, the Palearctic in the west and the Oriental in the east, with also some contribution from Ethiopian region towards the south. It extends between 60º52΄to 75º 22΄ E longitude and 24º to 37΄ latitude. Its southern border touches the coast of Arabian Sea, while northernmost tip lies in the permanent snowfields of Pamir in the greater Himalaya. Towards the east lies India whereas Iran and Afghanistan is located along the western borders (Mas- roor, 2012).

1.3 Physiography

Pakistan comprises of three major geographical units. The first unit is, mountains and plateaus in the north, northwest and west, representing a confluence of three huge mountain ranges, the Karakoram, the Hindu Kush and the Himalayas. Some of the world‟s

highest peaks lie in these ranges. This mountainous rampart has profoundly affected the geology, physiography, hydrology and climate, and has an important role in the present- day composition and distribution of the fauna and flora of the subcontinent. The second unit is the salt range at the foot of the Himalayas. It is one of the most ancient parts of the subcontinent and cradles a record of 600 million years of history of deposits of debris washed down from the Himalayas. It represents the ancient Tethys Sea (Powell, 1979). The third unit is the vast plain that is really the watershed of the mighty River Indus. The Indus and its drainage basin form the dominant physiographic feature of the fertile plains of Punjab and Sindh. The Indus Valley is the most extensively cultivated and densely populated part of Pakistan (Khan, 2006).

1.4 Climate

Pakistan is positioned in the temperate zone. Its arid subtropical climate is typified by cold winter and hot summer. Variations in latitude, altitude and proximity to gigantic mountain ranges all affect Pakistan‟s climate. Four seasons characterize Pakistan including dry winter from December through February, spring period from March through May, hot summers from June through September and rainy season or southwest monsoon from July through August. Higher mountains in the north and west are climatically moist alpine to subalpine with low soothing temperatures in summer, while temperatures in the plain often exceed 54ºC. The northern and southern parts of the country receive 40-140 cm of rain annually, resulting in yearly flooding of the plains. However, the western and southern parts of the country rarely receive more than 10 cm of rain annually and remain desolate and dry (Khan, 2006).

1.5 Eco-regions

Pakistan is a forest poor country; only about 3.8% of its total area is forested. More or less fifteen pronounced ecobiological habitat types have been distinguished in Pakistan (Khan, 2006; Roberts, 1991). These include littoral and mangrove, riverine, swamps and seasonal inundations, sand dune deserts, tropical thorn forests, subtropical semi ever-

green forest, dry temperate semi-evergreen scrub forest, subtropical pine forest, dry mixed deciduous forest, Balochistan hill ranges in southern latitudes and lower slopes of some northern ranges, Baluchistan higher ranges, Himalayan moist temperate forest, Himalayan dry coniferous forest, cold desert and dry alpine zone and Himalayan moist alpine zone.

1.6 Taxonomy

Coleoptera is the largest order in the animal kingdom (Nieto and Alexander, 2010) and insects (Borrer et al., 1984) with more than 350,000 known species. Scarabaeoidea is a very well-defined monophyletic group with a number of distinctive derived features. The family-level classification has varied over the years with some authors recognizing only three families (Lucanidae, Passalidae and Scarabaeidae) and others as many as twenty five. Smith (2006) recognized 12 families in super family Scarabaeoidea.

Scarabaeoides are usually easily recognized by their relatively large-sized spinose tibiae, simple tarsi, stout-body with reinforced hind wings and relatively small antenna. Unique characteristics of adult dung beetles include short legs with lamellate antennae, highly modified pro-thorax, large coxae, dentate tibia and strong intrinsic wing folding mecha- nism and usually c-shaped grubs. These beetles are rounded although some have metallic sheen (Klausnitzer and Krell, 1996).

Family scarabaeidae of superfamily scarabaeodidea is the largest family of order coleoptera that contains 30,000 reported world-wide species (Fincher, 1981). Members of subfamily scarabaeinae feed exclusively on feces of mammals like cows and buffalos (Woodruff, 1973). The members of the subfamily scarabaeinae are popularized as Dung Beetles because the dung of mammals is their main feed. In this way, these highly useful ecological services to both human and livestock by these scavengers in recycling of dung is of utmost importance (Sewak, 2009).

1.7 Distribution

Scarabaeinae are more or less cosmopolitan in distribution, but their abundance and diversity drops off considerably in cooler climates. Scarabaeinae are widely found in both temperate and tropical regions in Australia, America north of Mexico, Canada, Alaska or Greenland, New Zealand, Europe, Japan, Hawaiian Islands and Costa Rica.

1.8. General appearance (Figs. 1 & 2)

A somewhat flattened to convex or strongly convex body. Unevenly curved body sides. No capability to conglobate (rolling into a ball) the body. Dorsal surfaces of body either glabrous or subglabrous, or clothed with distinct hairs, setae or scales. The stiff, erect, dark bristles are not included in vestiture of upper surfaces. Vestiture of dorsal surfaces not including scales or scale-like setae. No deep foveae over the upper surfaces of body. Prothorax, metathorax and-or abdomen lack extrusible glands. Ventral body without hydrofuge surface (s). Number of basal ventrites connate five or six and the number of an- tennomeres 8–9. Mesocoxae are separated by a gap more than shortest diameter of coxal cavity and procoxal cavities internally open. Highly reduced scutellum; or absent or in- visible scutellum and number of abdominal ventrites six.. Visible part of metepisternum short and broad; or moderately elongated exposed portion. Metatibial spurs single and at the middle the mesocoxal cavities are moderately to widely separated.

Fig. 1. Dorsal view of scarab beetle (Ratcliffe 1991)

Fig. 2. Ventral view of scarab beetle (Ratcliffe 1991)

1.9 Significance

The most diverse and the largest order of class insecta is coleoptera which is represented by 166 families (Borrer et al., 1984; Nieto and Alexander, 2010) with more than 350,000 species (Wheeler, 1990). The oldest fossils of beetles, about 265 million years old are from the lower Permian (Ponomarenko, 1995). It includes 12 families, 43 subfamilies, 118 tribes, and 94 subtribes, recognized as valid taxa (Smith, 2006). The family Scara- baeidae includes 27,800 species and the two subfamilies Aphodiinae and Scarabaeinae are represented by approximately 6850 species worldwide and the subfamilies; Orphni- nae, Melolonthinae, Cetoniinae, Valginae, Rutelinae, Trichiinae and Dynastinae include approximately 20,950 species (Ratcliffe and Jameson, 2001)

The family Scarabaeidae (commonly referred to dung beetles) of superfamily scarabaeodidea is the largest family of order coleoptera that has more than 30,000 worldwide reported species (Fincher, 1981). Dung beetles feed exclusively on feces of cows and buffalos (Woodruff, 1973). These coprophagus beetles play a vital role in nature‟s sanita- tion by utilizing cattle‟s dung as food in their larval and adult stages by burying them in- side the soil. This not only provides dung manure to the soil, but also increases its fertility. In absence of native fauna, Bornemissza (1960) introduced the exotic species of dung beetles to improve the pastures in Australia.

It is established that dung beetles also play an active role in controlling the development of many fly pests and enteric parasites of cattle and livestock by rapid dissemination of contaminated faecal matter, thus minimizing the parasitic potential of the dung. On the contrary, the phytophgous beetles popularly known as leaf chafers, cockchafers and rose chafers are the pests of agricultural crop, forests and plants; these scavengers provide useful ecological services to both livestock and humans (Sewak, 2009). Each year, livestock, humans and crops produce proximately 38 billion metric tons of organic wastes worldwide. The calculations do not take into account the benefit of recycling of nutrients, de- creased environmental pollution, decrease in the need for landfills, and the significant reduction in diseases (Pimentel et al., 1997).

Carrion consumption is mostly developed in neo-tropical scarabaeinae (Gill, 1991). In 24 hours, dung beetles larvae can consume more than their weight and are considered beneficial because they speed up return of manure into the soil. If dung stays on soil surfaces and dries, 80% of the nitrogen is lost into the atmosphere (Fincher, 1997). No overlap of dung pats, dung excreted by cattle on pasture cover an area of nearly 0.8 square meters per animal per day. Near the dung pats, other cattle‟s avoid grazing because of dislike vegetation growing beside the pats (Anonymous, 1999).

Due to degradation of the dung by dung beetles larvae feeding in the pasture area, the pasture areas increase, otherwise it would cover the earth surface (Fincher, 1981). Each day larvae may consume 175-530 % of their dry body weight in dung (Holter ,1974). The economic value of the ecosystem benefits in the United States was estimated more than 2 billion dollar provided by dung beetles (Anonymous, 1999).

After the deposition within hours, dung beetles transfer all dumped feces of mammals into the soil (Arrow, 1931) and in soil fertility, highly localized differences are important in structuring plants communities in nutrient poor soil (John et al., 2007). In seed dispersal, dung beetle play a vital role (Vulinec, 2000); seed swallowed by frugivorous mammals are often defecated intact and viable (Garber, 1986). However, unless buried, 90 % of seed defecated by rodents or other seed predator may be destroyed. Dung beetle ac- complishes burial of seed completely (Estrada et al., 1991).

Three groups of dung beetles are important. Rollers make burrows a little distant to the dung, they are going to consume and then making small to medium sized lumps of dung rolled into their burrows and the rolled balls are utilized as a source of food or brooding chambers. The tunnelers include species that are known to bury dung where they find. The third group is called dwellers that simply live in manure. They neither roll nor bur- row dung (Hanski and Combefort, 1991). Coprophagous insect larvae mouthparts are specialized for fibrous food or chewing solid such as carrion or dung. Adults have mouthpart which allows feed them only on smaller dung particle and dung liquids

(Halffter and Mathews, 1966). During the feeding, beetles use setael labial and labral pads to concentrate dung particles (Halffter and Edmonds, 1982; Landin, 1960).

Hingston (1923) studied the importance of dung beetles in terrestrial ecosystem; these beetles are nature scavenger by burying of dung into the ground. They also help to main- tain the atmosphere by utilizing and dispersing of the organic waste (Mittal 1993 b) which is very important in ecosystem because of the removal of detritus from the soil surface (Halffter and Matthew, 1966).

According to Halffter and Favila (1993), dung beetles are the best models for the degree of changes in vegetation modified by the animals. In addition, these beetles contribute in nutrient cycling, pest reductions and removing of dung from the soil surface (Fincher 1981; Bornemissza, 1960). For the food resources, these beetles successfully compete with pest flies (Fay and Doube, 1983; Blume et al., 1973; Bornemissza, 1970; Doube and Moola, 1988 and Tyndale-Biscoe,1990).

Dung beetles are mainly coprophagous insect which play vital role in the ecosystem by aiding in soil aeration, the burial of seed, transportation of other organism and the seeds swallowed by mammals were defecated onto the soil surface and destroyed by rodents or other predators of seed (Vulinec, 2000; Rougon and Rougon, 1990), seed protection from predation (Feer, 1999; Estrada & Coates-Estrada, 1991) removing of dung from the soils surface (Tyndale-Biscoe, 1994; Gillard, 1967) and also these beetles help in reduction of population of disease causing organism such as hookworms and flies by challenging for food (fecal) resources and destruction of their larvae and eggs (Hanski, 1991a).

Majority of the dung beetles species are directly affected by changes in mammal popula- tions and rely on mid to large-sized mammal for the food (Estrada & Coates-Estrada 1999). Additionally, different beetle species are specialized in different habitat types such as edge, clearing, forest and tree and crop plantations, they are also extremely disturbance sensitive (Halffter et al. 1992; Halffter and Favila, 1993). Dung beetles are also important as environmental bioindicators (Nichols et al,. 2007, 2008).

Suhail et al. (2002) studied biodiversity and patterns of carabid beetles communities in agro forest area of Faisalabad, Pakistan. Meadows were the most rich and steady areas, while lower values were recorded from maize and sunflower fields. By using bait-free pitfall trapping, 40,694 insect specimens were collected. The dominant group was coleoptera (23.6%) with 10,497 specimens. Carabidae (45.0 %) and Staphylinidae (30.8 %) were the most significant among the beetles. Barberena-Arias and Aide.(2003) studied that the diversity of species and trophic composition of litter insect during plant secondary succession in four chrono-sequences of secondary forests. Each sequence included three sites abandoned for 5, 30 and over 60 yrs ago. A total of 5,919 insects were collected and assigned to 70 adult morpho-species. Two adult morpho-species (coleoptera and diptera) were shared between a 5-year site and a 30-year site. Collembola, Thysanoptera, larvae of Diptera, Coleoptera, Homoptera and Hemiptera were found in all successional stages.

In terms of phylogeny and taxonomy, scarabaeids beetles are one of the most studied group (Iablokov-Khnzorian, 1977; Balthasar, 1963; Browne and Scholtz, 1998; Mona- ghan et al., 2007; Janssens, 1949; Zunino, 1984; Philips et al., 2004), ecological studies and natural history (Hanski and Cambefort, 1991; Halffter and Matthews, 1966) and be- havioral studies (Halffter and Edmond, 1982). The worldwide fauna of dung beetle includes 12 tribes under which 234 genera and 5000 species (Hanski and Cambefort 1991).

Balthasar (1965) produced the first monograph for subfamily scarabaeidae of Oriental and Paleractic regions. Halffter & Mathew (1966) described dung beetle of subfamily scarabaeinae and discussed their natural history. Nikolajev (1973) described two new species and new genus of scarabaeidae beetle from Palaearctic region. Nikolajev and Ka- bakov (1977) identified four new species under family scarabaeidae from Afghanistan.

Kabakov (1982) contributed to the fauna of Lamellicorn beetle under genus Onthophagus from middle Asia, Afghanistan and Iran. Jessop (1985) provided identification guide to Eurysternine dung beetle (coleopteran: scarabaeidae). Gill (1986) introduced new species of Copris muller from south India. Krikken and Huijbregts (1987) identified five new

large-eyed species of Onthophagus from Sundaland and produced a key for identification. Storey and Weir (1990) reported a new species, Onthophagus latreille from Austral- ia.

Browne and Scholtz (1995) studies articulation and base of hind wings of Scarabaeoidea based on examination of over 250 genera from 13 Scarabaeoidea families. Faragella (2000) worked on three major species of dung beetles in three ecological systems of western Saudia Arabia. Ractcliffe and Jameson (2000) produced key for families and sub families of Scarabaeoidea of New World. Kon et al. (2000) reported one new species of genus Onthophagus from Sawara Borneo.

Ziani and Gudenzi (2001) explored Onthophagus species in Syria. Harpootlian (2001) studied scarab beetles of South Carolina. Clarissa and Simmons (2003) observed the fertilization success and genital morphology in Onthophagus taurus. Edmonds and Mellow (2003) produced biological and taxonomic notes on a rare Phanaeine, Phanaeus alvarengai.

Bert and Kohlmann (2006) classified subfamily Scarabaeoidea on the basis of historic analysis of the classification and Vaz-de-melo and halffter (2006) reported new genus of dung beetle along with two new species from Chile while Carpento et al. (2007) conducted a survey of scarab beetles from northern Pakistan (unpublished data). (Keith and Ka- ren. 2008) redescribe Mexican duringbeetle species Liatongus monstrosus and observed type of Attavicinus which is a new genus.

Deschodt and Scholtz (2008) studied dung beetles of size, less than 5mm under tribe Ca- thonini of temperate forests in South Africa and explored one new species and Gordon and Barbero 2008) made a study of dung beetle fauna of family scarabaeinae occurring in Laikipia district of Kenya and found 79 species which were diagnosed, keyed and prefer- ences were also discussed.

Sewak (2009) prepared diagnostic keys for the species, genera and tribe of subfamily Coprinae of India. Enrico et. al. (2009) did the systematic and phylogenetic studies of new genus of dung beetle Eodrepanus. It comprises only nine species. Tarasov and Ka- bakov (2010) identified two genus Onthophagus from Indochina. Silve et al. (2010) reported dung beetle community at two sites in Brazil forest and plain areas of the country.

Suhail et al. (2002) studied the biodiversity of coccinellids and their role as bioindicators in an agroecosystem. Collection was made by hand picking and hand netting from both of the crop and forest area. They noted that the coccinellids were very vulnerable to natural enemies and anthropological influences. Shannon diversity index was used to count the species diversity. It was concluded that aphid outbreaks in this area caused the population fluctuation of coccinellids beetles.

Parkash (2002) described various factors causing decrease in faunal diversity in Raja- sthan. He noted that due to human interaction and various ecological changes in the native ecosystems, a few animal species have been vanished from Rajasthan. This isolated and inhospitable desert was inhabited by fair number of mammals, birds, reptiles and insects. At several spots due to water logging and seepage from the canal, marshy condi- tions were prevailing. The assessment of the impact of canal irrigation and land use pattern on insect and rodents were studied. He finally discussed various strategies for the conservation and management of faunal diversity.

It is not easy to record, describe or even count the number of extant species. Life forms are so diverse that many years of specialized training are required to be able to recognize and describe them and even then, such training will provide knowledge of just one small group of living organisms (Agarwal et al., 1996). The simplest measurement of species diversity is a species count. Simple species counts remain the most popular approach to evaluate species diversity and to compare habitats or species assemblages (Humphries et al., 1996).

1.10 Scarabaeinae studies in Pakistan

The present studies were carried out in the northern parts of Khyber Pakhtunkhwa (KPK) and Federally Administrated Tribal Areas (FATA) which are predominantly rugged, mountainous and not easily accessible. Since last 10 years these areas were badly affected due to many reasons. The earthquake in 2005 brought devastation in northern Pakistan where official death toll was more than hundred thousand and millions of people were rendered homeless. Unfortunately since 2007, the war on terror started and more than 3.5 millions of people from these and surrounding areas have become refugees. In 2010, because of flash floods, approximately one-fifth of Pakistan's total land area was underwa- ter specially Khyber Pakhtunkhwa (KPK) province infrastructure which was already damaged. As a result of these natural calamities and unavoidable circumstances, this rendered un-conducive environment to carry out field trips and collect data.

Despite their immense potential and diversity, the Scarabaeinae fauna of northern part of Pakistan have not been investigated in depth so far. The first comprehensive account of Scarabaeinae beetles of Pakistan (the then part of Indian subcontinent as British India) was published by Arrow (1931) in The Fauna of British India, wherein he reported 34 species from Pakistani region. Since then, several taxonomic works on the systematic of dung beetles have been published. Siddiqui and Kamaluddin (2011) provided the first checklist for the distribution of the species in genus Onthophagus Latreille, 1802 including some new records from Pakistan. Likewise, description of some new species and distributional notes on Scarabaeinae from Iran has also been published (Kabakov, 2006; Zi- ani, 2006; Kabakov, 2008; Mowlavi et al., 2008; Ziani and Gharakhloo, 2010). Recently, a series of publications on non-coprophagous scarab beetles from Pakistan emerged (Keith and Sabatinelli, 2010; Keith and Saltin, 2012; Ratcliff and Ahmed, 2010). The catalogue of Palearctic Coleoptera (Löbl and Smetana, 2006) provides a list to 173 species in 8 tribes and 23 genera for subfamily Scarabaeinae in the region. In Pakistan, the family Scarabaeinae is represented by 70 species in 19 genera.

However, all these aforementioned works did not represent collections from northern parts of Pakistan, which in turn is ecologically and biologically very diverse region. Ar- row (1931) reported 11 species of dung beetles in subfamily Scarabaeinae from some parts of northern Pakistan. Arrow‟s work (1931) was though a serious attempt to describe well this poorly known group of beetles yet his work remains outdated in the context of present systematic changes. Making the situation even worse, Arrow was unable to pro- cure and include in his list the specimens from most parts of Pakistan, especially the climatically hostile and rugged mountainous regions of northern parts. This all situation ne- cessitated to work afresh on the current systematic of subfamily Scarabaeinae while pro- curing specimens from every possible locality of northern Pakistan.

1.11. Objectives The knowledge of various aspects of subfamily scarabaeinae will provide foundation for future studies; therefore, the present study is aimed at knowing about-  Collection of scarabaeinae fauna from selected parts of northern Pakistan  Distribution and diagnosis for collected specimens  Preparation of taxonomic identification keys

The hypothesis framed for the present study is that the northern parts of Pakistan (our present study area) harbors unknown and unexplored diversity of dung beetles belonging to subfamily scarabaeinae. The study area falls at the junction of both Palearctic and Ori- ental regions and therefore is expected to harbor potentially new records or even new descriptions of dung beetles.

2. MATERIALS AND METHODS 2.1. Study area The present study area of northern Pakistan comprises of selected parts of Khyber Pakh- tunkhwa (KPK) province and Federally Administered Tribal Areas (FATA) (Fig. 3). Its southern tip is only 250 meters above mean sea level, while to the north, the elevation rises quickly, reaching the high mountains of the Hindu-kush and Karakorum within a few hundred kilometers. On average, these mountains are 3,000 meters high; Tirich Mir in Chitral district is the highest at 7708 meter (www.ais-portal.com). The study area is bounded by Punjab and Gilgit-Baltistan in the east and south by Balochistan, in the west by Afghanistan and the China in the north. The narrow Wakhan corridor in Afghanistan separates the Khyber Pakhtunkhwa from the rest of central Asia in the north.

Approximately, three quarters of this area is under the administration of the provincial government and rest is part of the Federally Administered Tribal Areas. The area under the administrative control of the Khyber Pakhtunkhwa province is divided into seven civ- il divisions, which are further subdivided into 25 districts (www.iucn.pk). The Federally administered tribal area (FATA) is divided into seven agencies.

2.2. Collection of scarab beetles

The material for the present study was collected from different localities of the study area. In addition to, already preserved specimen in Pakistan Museum of Natural History (PMNH), Centre for Agricultural Bioscience International Rawalpindi (CABI) and National Insect Museum (NIM) National Agriculture Research Council (NARC) Islamabad were examined for the comprehensive study. The study area was further divided into three ecological regions each. 2.3. Sampling areas The present study was conducted in northern Pakistan in the three selected ecological zones including Alpine Zone, Montane Temperate Forest and Tropical Deciduous Forest.

Fig. 3. Administrative map of the study area

These zones were further divided into sub zones. Description of these zones and sub- zones are given below.

2.3.1. A) Alpine zone

Alpine zone was typified by continuous grass fields and dotted all over with tumbled boulders. Alpine zone was subdivided into alpine meadows, sub-alpine scrub and birch forest and alpine dry steppe.

2.3.1.1. Alpine meadows

Alpine meadows comprised of Northern Hazara District, Chitral, Swat, Kohistan and all the regions where mountains extended above the coniferous forest tree line.

2.3.1.2. Sub-alpine scrub and birch forest

This area comprised of upper slopes throughout higher mountain range of the Himalayas, including the north-eastern corner of Hazara District, Swat and Kohistan.

2.3.1.3. Alpine dry steppe

It was typified by side valleys of lower Chitral, Kohistan, western border of Waziristan, some parts of Safed Koh, Malakand, Swat and Dir.

2.3.2 B) Montane temperate forest

Montane temperate forest, which is the only real “tall tree” forest in Paki- stan included fallowing sub-regions.

2.3.2.1. Sub-tropical pine forest

A fairly narrow zone ranging between 3000ft and 6500ft is typified by lower Kaghan Valley around Kuwai, Batrassi pass (Hazara) and lower Swat around Marghazar and Buner.

2.3.2.2. Himalayan moist temperate forest

It is predominantly a coniferous forest with high rainfall during monsoon season and with glades of mixed deciduous broad-leaved species. This area included parts of eastern Swat bordering Indus Kohistan, lower Ka- ghan, Shogran, Murree hills and Kohistan.

2.3.2.3. Himalayan dry temperate forest

It consist of inner and northernly ranges of the Himalayas confined to the more sheltered lower slopes including Jabba valley in Swat, Dir, Chitral, Inner valleys of Hazara, Indus Kohistan and Swat Kohistan.

2.3.3. C) Tropical deciduous forest

Jhelum valley, Rawalpindi foothills, Margalla hills, Kahota, Lethrar and Noorpur Shahan forms tropical deciduous forest.

2.4. Collection methods

Specimens were collected through hand picking, light and pitfall traps (Fig 4). The collected specimens were manually stored in vials containing 70% ethanol solution. All the specimens were transferred to insect repository of Pakistan Museum of Natural History (PMNH) Islamabad for their systematic studies. The specimens were then transfixed with entomological pins, passing through the right side of the thorax on stretching board as depicted in Fig- ure 5. Body, legs and antennae were straightened. In order to retain their permanent posture, specimens were left to dry on stretching board for at least 2-3 days, depending on temperature. All the dried specimens were kept in wooden collection boxes and care was taken to protect the specimens from pest attack by placing naphthalene balls in the boxes. All the specimens were systematically arranged.

a d

Fig. 4. a) Pitfall trap b) beetles floating on the water surface in the bucket c) light trap and d) hand-picking of beetles with forceps

Each specimen was labeled with catalogue number, place of capture, date of collection and collector‟s name as shown in Fig. 5 (a) and (b).

Fig. 5 (a) Fig. 5 (b)

2.5. Identification

The identification of the specimens was done in Pakistan Museum of Natural History (PMNH) Islamabad with the help of microscope and in comparison with reference material. Sources of literature include Arrow (1931), Balthasar (1963), Biswas (1978a, b), Biswas and Chatterjee (1985), Gorden and Oppenheimer (1975) and Sewak (1985, 1986, 1991, 2004a, b, 2005, 2006a, b, c). Several morphological characteristics such as antennal segmentation, head shape, horn, ventral sclerites of the abdomen, elytra, claws of tarsi, body shape, middle and posterior tibiae, pro, meso and meta-tibia and pygidium, were used for species identification (Barari, 2001; Borror et al., 1989; Booth et al., 1990.). Some of the specimens were identified by comparing specimens with collection in the Institute of Zoology (IOZ), Chinese Academy of Sciences (CAS), Centre for Agricultural Bioscience International Rawalpindi CABI and (NIM) National Insect Museum, National Agriculture Research Council Islamabad.

For each species complete synonymies, geographical distributions, diagnosis and photographs are provided. The identification keys have been constructed for all the species. The taxonomic characters of 1460 dung beetles were examined for their species level taxonomy. Kyowa Optical microscope (Model SDZ-P) was used to analyze various parts of the body. Distribution maps were prepared in Esri ArcGIS 2013.

3. RESULTS AND DISCUSSION Taxonomic Account Order COLEOPTERA Linnaeus, 1758 Suborder POLYPHAGA Emery, 1886 Family SCARABAEIDAE Latreille, 1802 Subfamily SCARABAEINAE Latreille, 1802

The present work is produced as a result of extensive field collection in the northern parts of Pakistan and approximately 4,250 scarabaeinae specimens were procured. The laboratory examination under the microscope using Kyowa Optical microscope (Model SDZ-P) Japan, examining important morphological characters as well morphometry reveals the presence of 24 species of scarabaeinae in 10 genera, out of which eight species were reported for the first time from Pakistan. Identification keys to genera and species were prepared and a brief account of the distribution and diagnosis of taxonomically important characters for each species with is also provided. Locality records of each species are presented in the form of distribution maps.

3.1 Subfamily SCARABAEINAE Latreille, 1802

Body slightly flattened to moderately convex; or strongly convex. Antennae lamellate with 8 or 9 segments. Body incapable of conglobation (rolling into a ball). Dorsal surfaces of body glabrous or sub glabrous; or clothed with distinct setae, hairs or scales. Dorsal surfaces of body without deep foveae. Pro and metathorax and-or abdomen lack extrusible glands. Ventral side of the body without hydrofuge surface. Number of basal ventrites connate, 5 or 6. mes-ocoxae isolated by more than shortest diameter of coxal cavity. mesocoxal cavities at centre moderately to widely separated. Internally open procoxal cavities and highly reduced scutellum, or not visible or absent. Visible portion of metepisternum short and broad, or moderately elongate. metatibial spurs single. Elytra exposing pygidium, six visible fused abdominal sternites. Sometimes anterior tarsi absent either in female or in both sexes.

3.2 Genus HELIOCOPRIS Hope, 1837

Large head with carina or a cephalic horn. Body large broad and massive. Clypeus is curved or somewhat excised in the middle and eyes are not very deeply divided. Anten- nae 9 jointed and prothorax shows a complete basal groove. The front coxae are prominent, the middle coxae are long nearly parallel and not very widely separated. Hind coxae have ridges at the inner end within the coxal cavities and the abdomen is completely covered by the elytra. The two species of the genus Heliocopris have been found in the studied area

3.2.1 Key to the species of HELIOCOPRIS Hope, 1837

1. Medial gibbosity on the pronotum less tapering toward the head end, compara- tively shorter, its anterior margin broadly truncate and emarginate. Median division of the cephalic-shield protuberance not emarginate in the middle ………………………………..……………...Heliocopris midas (Fabricius, 1775)

2. Gibbosity on the pronotum occupies almost the entire width, distinctly surrounded in ridge-like toward the head end and simply bent. The pronotum is composed of more or less transverse, smooth and ridged. Head with a horn and pronotum with well developed gibbosity. Elytra very smooth, shiny and with very delicate striae. ………………………………………………. H. bucephalus (Fabricius, 1775)

3.2.2 Heliocopris midas (Fabricius, 1775) (Fig. 6.1 & 6.2) Scarabaeus midas Fabricius, 1775, Syst. Ent., 1: 21 (BMNH). Heliocopris midas: Bates, 1868, Col. Hefte, 4: 87.

Type Repository

Unknown

Diagnosis

Length: 24 mm to 30 mm. The pronotum is less tapering toward the head end and relatively shorter its anterior margin more broadly truncate and also more broadly emarginated but the emargination hardly narrower than the median emargination of the cephalic shield. Median division of the cephalic-shield protuberance is not emarginated in the middle.

Material Examined

1♂2♀, Mansehra, 03.VIII.2006; 4♂ 5♀, Buner, 11.VIII.2007; 1♂ 2♀, Dir 27.VII.2012; 2♂ 1♀, Peshawar, 12.VII.2012.

Worldwide Distribution

Southern Asia: India and China.

Remarks

This species is previously recorded by Hashmi and Tashfeen (1992) from southern Paki- stan. The present record extends the distribution of this species to northernmost Pakistan. This species is plentiful in the coprophagus environment at lower elevations. The specimens were usually collected away from the dung.

Figure 6.1: Heliocopris midas

Figure 6.2: Distribution map of Heliocopris midas

3.2.3 Heliocopris bucephalus (Fabricius, 1775) (Figs. 7.1 & 7.2) (New country record) Scarabaeus bucephalus Fabricius, 1775, Syst. Ent., 1: 24 (BMNH). Scarabaeus cristatus DeGeer, 1778, Mem. Hist. Ins., 7: 636, t.47, f.5♀,. Heliocopris tmolus Fischer von Waldheim,1822, Entomogr. Russ., 1: 13, f.2♀,.

Type Repository

Unknown

Diagnosis

Length: 29 mm to 30 mm. Large pronotum covers the head with a strong clypeus under- neath. The color is reddish-brown to black. Clypeus covers the biting mouth parts. A pair of compound eyes and lamellate antennae. The front wings are hard and there is a cover sheath of membranous hind wings. Females have no horn. The males have three horns one on the clypeus and two on the pronotum. The elytra very shiny and smooth with very lightly impressed delicate striate. The pygidium covered with delicate scattered punctures.

Material Examined

2♂1♀, Abbottabad, 11.VII.2007; 3♂4♀, Kohat,11.VIII.2010; 4♂, Kurram Agency, 11.VII.2012; 2♂3♀, Besham, 11.VIII.2012; 2♂1♀, Peshawar, 11.VIII.2013.

Worldwide Distribution

Indian, Southeast Asia and Southern China.

Remarks

This species constitutes a new country record for Pakistan. This species is commonly found in the foothill environments, pastures and forest edges in loamy soil and well- drained sandy clay.

Figure 7.1: Heliocopris bucephalus

Figure 7.2: Distribution map of Heliocopris bucephalus

3.3 Genus DIGITONTHOPHAGUS Balthasar, 1959

Protibia of males slender, long and more or less curved with a long, usually digitiform process in the inner angle of anterior edge. Outer teeth usually short and standing at right angles. Outer apical edge of metatibiae trilobate in most cases and anterior edge of basal antennal segment frequently serrate. The upper side usually black or very dark brown

3.3.1 Key to species of Digitonthophagus Balthasar, 1959

1. Semicircular head with the front margin. Forehead is finely punctured and divided from the clypeus by a strong curved carina. Elytra are finely striate and unpunctured. Pygidium bears an angulate basal carina and a few scattered punctures………………………………...Digitonthophagus gazella (Fabricius, 1787)

1”. Semicircular head with the front margin. Clypeus is lightly granulates and is separated from the forehead which is more sparsely granulated by a strong curved carina. Elytra are finely striate with extremely few and minute punctures. Pygidium bears an angulate basal carina and similarly light and minute punctures………………………..…………….…..……D. bonasus (Fabricius, 1798)

3.3.2 Digitonthophagus gazella (Fabricius, 1787) (Figs. 8.1 & 8.2) Scarabaeus gazella Fabricius, 1787, Mant. Ins., 2: 377. Scarabaeus dorcas Olivier, 1789, Ent., 1(3): 121. Copris antilope Fabricius, 1798, Ent. Syst. Suppl.,: 32. Copris metallicus Fabricius, 1798, Ent. Syst. Suppl.,:28. Onthophagus intermedius Reiche, 1840, Revue Zool., 3:243. Digitonthophagus gazella: Balthasar, 1959, Sbornik Ent. Odd. Nár. Mus. Praze, 33: 464.

Type Repository Zoological Museum, University of Copenhagen

Diagnosis

Length: 6 mm to 11 mm. Head is semicircular with the front margin. Forehead is finely punctured and divided from the clypeus by a strong curved carina. Clypeus in male is slightly rugulose and shining. Clypeus in female is slightly rugulose and thick. The vertex bears a straight carina. Pronotum bears thin granules in its median part. Elytra are finely striate and unpunctured. Pygidium bears an angulate basal carina and a few scattered punctures. The metasternum bears only a few punctures.

Material Examined

2♂4♀, Buner, 22.VIII.2007; 3♂, Mansehra, 13.VII.2012; 4♀, Abbottabad, 9.VIII.20012; 2♀, Peshawar, 10.VII.2013.

Worldwide Distribution

India, Baroda, Mauretania, Belgium, Sri Lanka, W. Africa, Kenya, Japan, Texas (introduced), Malawi, Senegal, Australian (introduced), Madagascar, Arabia, Central and South America (introduced).

Remarks

D. gazella is found in cow, human and horse dung. This species is distinct on the basis of horn present on clypeus and having no tubercle placed in middle. Uncommon to fairly common in cattle pastures in lowlands and less abundant on lowland grasslands and beaches. Siddiqui and Kamaluddin (2012) reported this species from buffalo and cow dung. Collected specimens showed resemblance with published description given by Ar- row (1931).

Figure 8.1: Digitonthophagus gazella

Figure 8.2: Distribution map of Digitonthophagus gazella

3.3.3 Digitonthophagus bonasus (Fabricius, 1798) (Figs. 9.1 & 9.2) Scarabaeus bonasus Fabricius, 1775

Type Repository

Zoological Museum, University of Copenhagen

Diagnosis

Length: 9 mm to14 mm. Head is semicircular in shape with the front margin. The clypeus is lightly granulates and is separated from the forehead which is more sparsely granulated by a strong curved carina. A short acute erect horn in the middle of the forehead. The vertex bears a pair of horns. Pronotum is very smooth in front and behind with a few scattered granules in its middle part. There is a slight median groove on each side. Elytra are finely striate with extremely few and minute punctures. The middle and hind femora are short and broad. The tibiae externally strongly toothed. Pygidium bears an angulate basal carina and similarly light and minute punctures.

Material Examined

4♂5♀, Kanju, 12.VII.2007; 2♂1♀, Swat, 10.VII.2008; 5♂, Mansehra, 13.VII.2012; 4♀, Swabi, 26.VII.20013.

Worldwide Distribution

India, Sri Lanka, Myanmar, Thailand, South Afghanistan, Tonkin (Vietnam), Cambodia and Belgium.

Remarks

Previously recorded by Arrow (1931) from Pakistan. Specimens were similar to published description provided by Arrow (1931). Species is distinct on the basis of horn present on clypeus and having tubercle placed in middle with slight basal tooth at inner edge. It is similar to Digitonthophagus gazella with visual external morphology (Syed et al., 1981; Siddiqui et al., 1974).

Figure 9.1: Digitonthophagus bonasus

Figure 9.2: Distribution map of Digitonthophagus bonasus

3.4 Genus ONTHOPHAGUS Latreille 1802

Clypeus is fused with the ocular lobes and variable in shape. Short body, compact, smooth or covered hairs or setae. The lateral margin of the prothorax is angulate in the middle and rounded at the base. The legs are stout, femora thick and front tibia with four but occasionally three external teeth. The middle and hind tibiae dilate greatly from base to end. The elytra cover the abdomen and have a single lateral carina and seven striae.

3.4.1 Onthophagus (Palaeonthophagus) marginalis (Gebler, 1817) (Figs. 8.1 & 8.2) Copris marginalis Gebler, 1817, Mém.Soc. Nat. Moscou, 5: 315 (MNHN). Onthophagus marmoratus Ménétriés, 1832, Cat.Rais.,: 176. Onthophagus circumscriptus Faldermann, 1835, Nouv.Mem. Moscou, 4: 241. Onthophagus marginalis: Reitter, 1892, Verh.Naturforsch. Ver. Brünn,: 68.

Type Repository

Leningrad Academy of sciences

Diagnosis

Length: 7 mm to 9 mm. Head is smooth with a few punctures in the middle. The clypeus unevenly punctured and the pronotum smooth in front. The anterior part of the pronotum is angularly excavated in front. The front tibia is slightly longer. Front tibiae are very short and broad. Elytra are fine and distinct but not very closely punctured. Pygidium is distinct but not very closely punctured. Mesometasternal suture is straight and entire front part of the metasternum is slightly depressed.

Material Examined

2♀, Kurram Agency, 12.VIII.2006; 1♂ 3♀, Upper Dir, 08.IX.2007; 2♂1♀, Chitral 08.IX.2008; 2♂, Malakand, 23.VIII.2009; 1♂2♀, Battal, 28.IX.20011; 3♂ 2♀, Shandur, 20.VIII.2012

Worldwide Distribution

Greece, Indonesia, North China, Afghanistan, Kashmir, Tibet, North India, Syria, Rus- sia, Iran, Tajikistan, Tanzania, Turkmenistan, Kyrgyzstan, Kazakhstan, Turkey, Bulgaria, Azerbaijan , Armenia, Poland, Nepal, Belgium and Israel.

Remarks

This is a high altitude scarabaeinae beetle, often encountered just in and below the alpine zone. The description of specimens meet the distinguishing characters for that reported in literature.

Figure 10.1: Onthophagus marginalis

Figure 10.2: Distribution map of Onthophagus marginalis

3.4.2 Onthophagus (Palaeonthophagus) nuchicornis (Linnaeus, 1758) (Figs. 11.1 & 11.2) Scarabaeus nuchicornis Linnaeus, 1758, Syst. Nat., 10(1): 347 (Type specimens transferred from UZIU to LSLC). Copris acornis Geoffroy, 1785, Ent. Paris,: 14. Scarabaeus xiphias Fabricius, 1792, Ent. Syst., 1(1): 59. Scarabaeus trituberculatus Schrank, 1798, Fauna Boica, 1: 397. Scarabaeus dillwyni Stephens, 1830, Illustr. Brit. Ent. Mandib., 3: 174. Onthophagus nuchicornis: Mulsant, 1842, Col. France, Lamell.,: 114. Onthophagus indistinctus Mulsant, 1842, Col. France, Lamell., : 115. Onthophagus rubripes Mulsant, 1842, Col. France, Lamell., : 115. Onthophagus nuchicornis ab. immaculatus Mulsant, 1842, Col. France, Lamell.,: 116. Onthophagus rhinoceros Melsheimer, 1844, Prov. Acad. Philad., 2: 134. Onthophagus alpinus Kolenati, 1846, Melet. Ent., 5: 5.

Type Repository

United Kingdom

Diagnosis

Length: 6 mm to 9 mm. The head with clypeus shining very lightly punctuate. Clypeus and gena evenly articulate and nearly semicircular. Clypeus and frons separated by a dif- ference in height the frons slightly higher and the pronotum less convex. The elytral striae are less shallow and punctate. The ventral surfaces of thorax more or less punctuate and most of the punctures bearing long, reddish setae. Abdomen largely impunctate except few scattered basal punctures on the last three segments. The pygidium roughly shallow punctuate and each puncture bearing a short tan setae.

Material Examined

2♂ 1♀, Kurram Agency, 08.IX.2007; 2♀, Shundoor, 15.IX.2008; 3♂, Swat, 27.VIII.2010; 2♂ 1♀, Malakand, 20.VIII.2012; 1♂, Buner, 02.IX.2013.

Worldwide Distribution

Mongolia, Turkmenistan, Estonia, Kazakhstan, Turkey, Albania, Armenia, Moldova, Bosnia and Herzegovina, Spain, Bulgaria, Belarus, Croatia, Ukraine, Yugoslavia, Poland, Czech Republic, Slovakia, Lithuania, Latvia, Georgia, Finland, Denmark, France, Ger- many, Luxembourg, Austria, Belgium, Britain, Greece, Russia, Hungary, Ireland, Italy, Netherland, Norway, Sweden, Switzerland and North America (introduced).

Remarks

This coprophagus species is also found at the higher altitudes in sympatry with O. marginalis. More cold hardy than O. taurus. The diagnostic features for the specimens in hand suit best the characters set reported in literature. Good dung burial and shredding noted. I have encountered this species in sandy/heavy loams soil.

Figure 11.1: Onthophagus nuchicornis

Figure 11.2: Distribution map of Onthophagus nuchicornis

3.4.3 Onthophagus (Phanaeomorphus) gagates Hope, 1831 (Figs. 12.1 & 12.2) (New country record) Onthophagus gagates Hope, 1831, Zool. Misc. : 22 (BMNH). Onthophagus angulatus Redtenbacher, 1844, Hügel‟s Kaschmir, 4(2): 522.

Type Repository British Museum.

Diagnosis

Length: 8 mm to11 mm. The clypeus bent at the forehead and vertexes are closely punctured. There is a fine curved anterior carina and a similar posterior one. The pronotum is sparsely punctured and the pronotum is convex and has only a slight tubercle on each side. Elytra are deeply striate, the striae weakly punctured and the intervals lightly punctured. Pygidium is rather strongly punctured at the apex and less strongly at the base. The metasternum a little depressed in front.

Material Examined

2♂, Allai, 08.V.2006; 1♂, Buner, 02.IV.2007; 2♀, Malakand, 20.VIII.2011; 2♀, Chitral, 15.IV.2012; 1♀, Upper Dir, 27.VIII.2013.

Worldwide Distribution Taiwan, Tibet, Laos, India, China, North Vietnam, Nepal and Assam.

Remarks

The studied specimens clearly exhibit characters of O. gagates. O. gagates is new addition to the country record for Pakistan. Previously recorded from Chhattisgarh, Assam, Manipur and Madhya Pradesh of India.

Figure 12.1: Onthophagus gagates

Figure 12.2: Distribution map of Onthophagus gagates

3.4.4 Onthophagus (Altonthophagus) kukunorensis Kabakov, 1990 (Figs. 13.1 &13.2) (New country record) Onthophagus (Altonthophagus) kukunorensis Kabakov, 1990, Trudy Zool. Ins., 211: 33 Onthophagus ( Colobonthophagus) tragus (Fabricius, 1792) Onthophagus hypertropis Olsoufieff, Ann. Muse. Zool. Acad. Imp., 1900, p. 5: 269. Onthophagus mandarinus Harold Berliner Entom. 1859, 2: 224. Caccobius tsunoellus Matsumara, Insecta Mutsumurana, 1938, p. 12: 55.

Type Repository

Denmark, København [= Copenhagen], University of Copenhagen, Zoological Museum

Diagnosis

Length: 5 mm to 10 mm. The head is broad, the eyes (seen from above) are fairly large and the clypeal margin is bilobed in the middle. Pronotum is distinctly, but not very strongly nor very closely puncture. Elytra are strongly striate, striate contain fairly numerous and distinct punctures and the intervals are slightly convex and minutely punctured. Front tibia with four short external teeth. The terminal spur is very short, flat and broad. A minute tooth appears between the two horns. Terminal tooth of the front tibia is long and the spur elongate and strongly curved. Pygidium is dense and finely punctured.

Material Examined

2♂ 1♀, Chitral 14.IX.2007; 3♂ 2♀, Swat 14.IX.2009; 3♂ 2♀, Kurram agency 02.VIII.2010; 4♂ 5♀, Mansehra 13.IX.2012.

Worldwide Distribution

India, Bangladesh, Thailand, Taiwan, China, Belgium, Myanmar, Malaysia, Java, Sula- wesi, North Vietnam and Nepal.

Remarks

This species is another new addition to the scarab fauna of Pakistan and represents the western-most distribution. This species is usually found at higher altitudes in alpine forests as well as at the alpine pastures.

Figure 13.1: Onthophagus kukunorensis

Figure 13.2: Distribution map of Onthophagus kukunorensis

3.4.5 Onthophagus (Coloboonthophagus) tragus Fabricius, 1792 (Figs. 14.1 & 14.2) Scarabaeus tragus Fabricius 1792, Ent. Syst., 1: 56. Onthophagus tragus, Arrow 1931, Fauna of British India including Ceylon and Myanmar (Coleoptera : Lamellicornia : Coprinae), 3 : 303·304. 50 Rec. zool. Surv. India. Occ. Pa- per No. 296 Onthophagus (Colobonthophagus) tragus 1963, Balthasar. Monographie der Scarabae- idae und Aphodiidae der Palaearktischen und Orientalischen Region (Coleoptera : Lamel- licomia). 2 : 1-627, 226 figs., 16 pls..

Type Repository

Unknown

Diagnosis

Length: 7 mm to 9 mm.. Head is broad and has large eyes. The sides of the head are bluntly angular and the clypeal margin bilobed in middle. The pronotum is distinctly but not very closely nor very strongly punctured. Front angles are rounded lateral margins straight in front. Elytra strongly striate the striate contain fairly numerous and distinct punctures the intervals are slightly convex and very sparsely and minutely punctured except at the sides where the punctures are larger. Pygidium is thick and rather lightly and thinly punctured. The metasternal shield is distinctly punctured at the sides and almost smooth in the middle and the sides of the metasternum strongly but not closely punctured.

Material Examined

1♂ 2♀, Peshawar, 14.IX.2006; 2♂ 3♀, Swat, 14.IX.2007; 4♂, Malakand, 02.VIII.2010; 3♂ 4♀, Islamabad, 13.IX.2012.

Worldwide Distribution

India, Myanmar, Bangladesh, Malaysia, Indonesia, Sulawesi, Laos, Thailand, Taiwan, South China, North Vietnam, Nepal and Tonkin (Vietnam).

Remarks

This species is common at the foothill habitats. The studied description of specimens matches those in literature.

Figure 14.1: Onthophagus tragus 54

Figure 14.2: Distribution map of Onthophagus tragus

3.4.6 Onthophagus (s. str.) proletarius Harold, 1875 (Figs. 15.1 &15.2) (New country record) Onthophagus proletarius Harold, 1875, Col. Hefte, 13: 89 (MNHN). Onthophagus mendicus Gillet, 1924, Ann. Soc. Sc. Brux., 44: 67 (ISNB).

Type Repository

M. Rene Oberthur‟s collection.

Diagnosis

Length: 5 mm to 7 mm. The head is not broad sides are lightly rounded and the pronotum is rather closely and evenly punctured. The front angle are moderately sharp. Well punctured and shining forehead separated from the clypeus by a short curved carina and middle of the front margin of the pronotum is vertical. Clypeus in male is shining shaped into a rounded lobe and unevenly punctured. Clypeus in female is transversely rugulose with the front margin rounded except in the middle where it is straight. Pronotum is a little hollowed smooth and shining just behind the head. The elytra are finely striate with the intervals flat and minutely punctured. The pygidium is shining and strongly punctured.

Material Examined

1♂ 2♀, Allai, 14.IX.2006; 2♂ 3♀, Besham, 14.IX.2007; 4♂, Chitral, 02.VIII.2010; 3♂4♀, Upper swat, 13.IX.2012; 2♂ 3♀, Dir, 14.IX.2007.

Worldwide Distribution

Myanmar, Malay Peninsula, Indo-China and S. China,Taiwan, Vietnam, Laos, Malaysia, Burma.

Remarks

These specimens constitute the first ever records of this species from Pakistan and ex- tending the range of this species to it western-most distribution. The species is found in the foothills as well as Himalayan dry-temperate forest.

Figure 15.1: Onthophagus proletarius

Figure 15.2: Distribution map of Onthophagus proletarius

3.5 Genus COPRIS Geoffroy, 1762

Body compact, convex or slightly depressed. Head is broad more or less semicircular and the clypeus is notched in the middle at the front margin. Prothorax is broad and transverse and its front margin is furnished with a membranous fringe and the hind margin with a deep median groove. Middle coxae are long and nearly parallel. Tarsi are short and the basal joint about twice as long as the second. Scutellum is wanting and the elytra are not very short having eight striae and a single lateral carina.

3.5.1 Key to the species of Genus Copris

1. Lateral teeth of the pronotum long and pointed. Clypeus distinctly emarginate. Cephalic ridge straight ….………..…..………………………………………….. 2

1” The centre gibbosity on the pronotum edged on both sides by a sharp small tubercle with punctures. Meta femora with sparse punctures, terminal spur of protibia simple and the apex itself rounded………….…… Copris repertus walker, 1858

2. The medial gibbosity deeper more roundish-emarginate at the base, the tubercles are taller and sharper ………………………. C. sacontala L. Redtenbacher, 1844

2” The punctures on the anterior wall of the medial gibbosity more dense the anterior wall itself approximately twice as wide as high. The small medial tubercles at the upper margin of the gibbosity very small. The elytra more densely punctuate …………………...... C. punctatus Gillet, 1910

3.5.2 Copris repertus Walker, 1858 (Figs. 16.1 & 16.2) (New country record) Copris repertus Walker, 1858, Ann. Mag. Nat. Hist., 2(3): 208 (BMNH). Copris sodalist Walker, 1858, Ann. Mag. Nat. Hist., 2(3): 208. –Waterhouse, 1891, Ann. Mag. Nat. Hist., 7(6): 512. (Syn.) Catharsius claudius Harold, 1877, Ann. Mus. Civ. Genova, 10: 48 (MCSN). –Lansberge, 1886, Tijdschr. V. Entom., 29: 11 (Syn.).

Type Repository

British Museum

Diagnosis

Length: 14 mm to 19 mm. Head is nearly semicircular. In male, the head bears slightly compressed erect horn, feebly curved with a minute tooth behind near the base on each side. Clypeus rugulose deeply notched in the middle. In female, the head bears a short, transverse, elevated carina with the angles. The Pronotum is shining with a strong longitudinal groove. In males, pronotum is almost smooth except in the front angles. The front angles truncate, sides are strongly rounded. Elytra are lightly striate except at the base and apex, the striate minutely punctured. Metasternal shield is smooth and the sides of the metasternum are strongly punctured. The front tibia is large and with four external teeth, the front femur is rather strongly punctured and the front tibial spur is nearly straight.

Material Examined

1♂ Peshawar, 16.VIII.2006; 2♂2♀, Dir, 04.VIII.2007; 3♂ 1♀, Abbottabad, 12.VIII.2012; 2♀, Buner, 26.VII.20013.

Worldwide Distribution Myanmar and India

Remarks

The species has been reported from Pakistan at foothill areas. The specimens were collected at night under the street lights and from houses, grasslands and pastures. The species is found in sandy to heavy loamy soils. The species is known for its excellent dung burial.

Figure 16.1: Copris repertus

Figure 16.2: Distribution map of Copris repertus

3.5.3 Copris sacontala Redtenbacher, 1848 (Figs. 17.1 & 17.2) (New country record) Corpis sacontala Redt. Hugel‟s Kashmir, iv, 2, 1848, p. 519. (NMW).

Type Repository Vienna Museum

Diagnosis

Length: 20 to 24 mm. Head reasonably broad and short except between eyes and feebly serrated in the middle of the front margin. The sides of pronotum strongly curved and the front angle truncate. Elytron finely and absolutely striates and with minute punctures. Py- gidium is fairly and strongly covered with ill-defined punctures and the metasternal shield is smooth in front with a deep transverse impression and strongly punctured. The metasternum sides are thickly punctate and clothed with hair. Front-tibia is broad and sustains three outward teeth and the hind and middle tibiae are digitate at the edge.

Material Examined

3♂ 4♀, Murree Hills, 10.V.2009; 1♀, Chitral, 10.VII.2011; 2♂ 1♀, Swat, 16.VII.2012; 1♂, Kurram Agency, 09.VIII.2013.

Worldwide Distribution

India and Sri Lanka.

Remarks

This species represents the new country record for Pakistan. The species is found in the Himalayan dry temperate forests and inner valleys in the mountainous environments in

sandy to heavy loamy soils. Found mostly in grasslands and pastures in cooler regime and is very good tolerant to frost.

Figure 17.1: Copris sacontala

Figure 17.2: Distribution map of Copris sacontala

3.5.4 Copris punctatus Gillet, 1910 (Figs. 18.1 & 18.2) (New country record) Copris punctatus Gillet, Notes Leyd. Mus. Xxxii, 1910, p. 4.

Type Repository

Unknown

Diagnosis

Length: 16 mm to 19 mm. Head is strongly punctured with the outline semicircular. Pro- notum is strongly but unevenly punctured and sides are strongly and somewhat irregularly rounded front angle truncate and the hind angle nearly obsolete. Front angles are not hollowed beneath. Elytra are intensely striate with closely and not very finely punctured. Pygidim is somewhat strongly and diligently punctured and the metasternal shield has a deep median groove is very deeply hollowed on each side with a few punctures in the hollows..

Material Examined

3♂ 4♀, Swat, 12.VII.2007; 2♂ 1♀, Allai, 07.VII.2012; 1♂ 3♀, Chitral, 13.VIII.2013; 2♂ 4♀, Kurram Agency, 12.VIII.2012.

Worldwide Distribution

Malay Peninsula and Borneo.

Remarks

This species represents the new country record for Pakistan and extends the distribution all across India to Pakistan. The species is distributed in the in the Himalayan moist-

temperate forests in the mountainous environments. This species is mostly found in the vicinities of human habitations in the rocky habitats.

Figure 18.1: Copris punctatus

Figure 18.2: Distribution map of Copris punctatus

3.6 Identification key to the collected genera of tribe Onitini

1. Small scutellum; base of pronotum not margined; absent protarsi on both sexes; prosternum without any process in both sexes...... Onitis Fabricius

1” Scutellum rather large; base of pronotum margined; in males protarsi absent and present in females; prosternum forming a strong bifid process behind procoxae in middle in male and in females such process absent; parameres twice as long as phallobase ...... Cheironitis van Lansberge

3.6.1 Genus ONITIS Fabricius, 1798

Relatively not broad head and un-processed or excavated prothorax. Body oblong and have stout legs and without front tarsi. Antennae 9 segmented with club relatively small; mouthparts especially maxillae with their membranous lobe broad, well pronounced and often fully visible in dorsal view; simply punctate pronotum or entirely smooth with a pair of well marked small impressions in middle of base; scutellum is visible but minute; elytra with 10 striae. The front-tibiae have four external teeth. Front legs are more or less elongate. The elytra completely cover the abdomen. Tibia without articulated terminal spurs, slender and curved towards the end The five species of the genus Onitis have been recorded in the northern part of Pakistan.

3.6.2 Key to species of Genus Onitis Fabricius, 1798

1. Pronotum very lightly punctured ……………….………………………….……2

1” Pronotum clearly and deeply punctured …..…………………………………..…..3

2. Pronotum feebly punctuate, punctures shallow and vague, Pygidium smooth, glabrous, Fronto clypeal carina interrupted or with tubercle at middle. Metasternum not excavated at middle.………...... …. 0nitis falcatus (Wulfen 1786)

2” Black; pronotum very lightly punctured ………...….O. lama van Lansberge, 1875

3. Metasternum flat, not grooved . . . . . ……………………………………………..4

3” Metasternum longitudinally grooved in front, Ventral surface entirely metallic. Upper Side rather shining…………….………….….O. philemon Fabricius, 1801

4. Fronto clypeal carina broadly interrupted; protibia with terminal external tooth projecting in front. Frontal carina with a wide medial gap, base of pronotum not bordered between foveae………………………….... O. subopacus Arrow, 1931

4”. Frontal carina with only a very short medial gap, fronto clypeal carina narrowly interrupted, protibia with terminal external tooth tapering in front .…………………………………………………………O. virens Lansberge, 1875

3.6.3 Onitis falcatus (Wulfen, 1786) (Figs. 19.1 & 19.2) Scarabaeus falcatus Wulfen, 1786, Descr. Capens. Ins., : 14, t.2, f.17 Onitis sphinx Herbst, 1789, Nat. Kafer, 2: 186, t.13, f.8 (nec Fabricius, 1775). Onitis hymaleyicus Redtenbacher, 1844, Hügel‟s Kaschmir, 4(2): 518 (ANLW). Onitis kiuchii Masumoto, 1996, Ent. Rev. Jap., 50: 88. –Hanboonsong et Masumoto, 2000, Elytra, 28(1):

Type Repository

Vienna Museum.

Diagnosis

Length: 15 mm to 22 mm. Posterior part of the head is irregularly granular. Clypeus is oval moderately strongly and lightly rugulose. Clypeo-frontal carina is widely interrupted in the middle and there is a short frontal tubercle behind it and in front a short oblique clypeal carina. The base of pronotum strongly rounded but not distinctly lobed. The pronotum is lightly and rather sparingly punctured without a well-marked median groove or line. The pygidium is thick unpunctured and smooth. The elytra finely striate with 1, 3 and 5th interval a little raised and distinctly narrower than the 2nd and 4th and metasternum sides are finely and hairy and also punctured densely

Material Examined

2♂ 3♀, Buner, 09.IX.2007; 4♂ 1♀, Mansehra, 12.VII.2008; 3♂ 1♀, Abbottabad, 14.VIII.2010; 3♂ 2♀, Besham , 25.VIII.2012; 1♂, Swat, 12.VII.2012; 3♂ 3♀, Malakand, 06.VIII.2013.

Worldwide Distribution

India, Myanmar, China, Taiwan, Philippines, Laos, Cochinchina and Thailand

Remarks

Previously recorded by Arrow (1931) from Pakistan. The species is widely distributed in the Himalayan Region semi arid zone but so far not recorded from the trans Himalaya and costal areas. This forest dwelling species attracts towards light and feeds on fresh and old cattle dung mainly of buffalos and cows and rarely found in accumulated dung.

Figure 19.1: Onitis falcatus

Figure 19.2: Distribution map of Onitis falcatus

3.6.4 Onitis lama van Lansberge 1875 (Figs. 20.1 & 20.2) (New country record) Onitis lama, Lansberge 1875, Ann. Soc. Ent., Belgium, 18 : 123. Onitis lama, Arrow 1931, Fauna British India Ceylon and Myanmar:3 : 389-390, PI. XI, Fig. 11 & 12. Onitis lama. Balthasar1963 Monographie der Scarabaeidae und Aphodiidae der Palae- arlctischen und Orientalischen Region (Coleoptera: Lamellicornia) 2 : 44-45.

Type Repository

Unknown

Diagnosis

Length: 18 mm to 22 mm. The head have a strongly elevated, slightly acute frontal carina with a short straight transverse carina and trisinuate vertex carina. Front angles of protho- rai blunt sides strongly rounded, base almost completely rounded. Front femur in males have a very sharp oblique tooth beyond middle on lower edge and hind femur with a tooth near end of lower edge. Broad front tibia armed with four external teeth

Material Examined

5♀, Buner, 18.VII.2006; 4♂ Peshawar, 20.V.2008; 3♂, 4♀, Islamabad, 12.VI.2009; 4♂ 1♀, Swat, 14.VIII.2012; 1♂ 3♀, Abbottabad, 02.VII.2013.

Worldwide Distribution

India

Remarks

This species constitute the first ever record of this beetle from Pakistan. Previously reported from Gujrat, Haryana, Punjab, Rajasthan, Uttarkhand and Uttar Pradesh in India (Arrow, 1931).

Figure 20.1: Onitis lama

Figure 20.2: Distribution map of Onitis lama

3.6.5 Onitis philemon Fabricius 1801 (Fig. 21.1 & 21.2) Onitis philemon Fabricus 1801, Syst Eleuth., 1: 30 (ZMUC). Onitis philemon var. sphinx Illiger, 1804, Mag. Ins., 3: 149 (SNMB). Onitis distinctus Lansberge, 1875, Ann. Soc. Ent. Belg., 18: 138 (ISNB). –Balthasar, 1963, Monogr. Scarab. Aphod. Palaearkt. Orient. Reg., 2: 40 (Syn.).

Type Repository

Brussels Museum.

Diagnosis

Length: 13.5 mm to 19 mm . Head is rugulose with the ocular lobes. Irregularly punctured pronotum and without clearly marked median lines, in front the lateral margins is straight and in middle strongly rounded. The pygidium is very feebly and sparsely punctured and elytra are moderately strongly striate and sparsely punctured. In males, clypeus is granulate and in females closely transversely rugose. Middle femur has a sharp tooth near the end of the lower edge. Hind trochanter is a little toothed beneath.

Material Examined

3♂ 4♀, Mansehra, 01.VII.2006; 2♂ 5♀, Kohat, 21.VIII.2007; 2♂ 3♀, Peshawar, 11.VI.2010; 5♂ 4♀, Chitral, 20.VII.2012; 2♂ 4♀, Abbottabad, 01.VII.2012; 4♂ 5♀, Nowshera, 18.VIII.2013

Worldwide Distribution

China, Taiwan, Bangal and India

Remarks

Widely distributed species found in loam to heavy loam, pastures and a high rainfall species. It is the most abundant and uniformly distributed species of the genus in the forests. It attracts towards light and feeds on carnivore and herbivore dung of human, cow and horse. Active throughout the year, prefers fresh or old dung and more common in dung heap accumulated for manure and collected along with O. subopacus.

Figure 21.1: Onitis philemon

Figure 21.2: Distribution map of Onitis philemon

3.6.6 Onitis subopacus Arrow, 1931 (Figs. 22.1 & 22.2)

Onitis philemon Lansberge, 1875, Ann. Soc. Ent. Belg., 18: 133 (BMNH (nec Fabricius 1801)). Arrow, 1931, Fauna Brit. India, Lamell., 3: 395 (Junior homonym of Onitis philemon Fabricius, 1801). Onitis subopacus Arrow, 1931, Fauna Brit. India, Lamell., 3: 395 (BMNH) (n. n. for Onitis philemon Lansberge, 1875 preoccupied).

Type Repository

Unknown

Diagnosis

Length: 15 mm to 20 mm. Prothorax and head is reasonably shining and relatively dull elytra. A short transverse clypeal carina is located in front of the pronotum and interrupted carina is somewhat closely but feebly punctured. The pygidium is opaque and unpunctured. The elytra exceptionally striate and the 1, 3 and 5th interval are commonly slightly convex. Middle femur bears a rounded lobe near the middle of the posterior edge and a sharp tooth towards the end and middle tibia is slender at the base and strongly and abruptly dilated. Trochanter of the hind leg is sharply toothed.

Material Examined

2♂ 5♀, Abbottabad, 3♀, 11.V.2005; 1♂5♀, Murree, 13.VI.2006; 5♀, Chitral, 21.VII.2007; 2♂ 1♀, Besham, 14.VII.2008; 2♂ 3♀, Islamabad, 02.V.2009; 7♂ 5♀, Now- shera, 03.VII.2010; 4♂ 5♀, Peshawar, 23.VIII.2012; 2♂, Swat, 10.VI.2012; 4♂, Kohat, 03.VII.2013; 2♂ 3♀, Dir, 13.IV.2014.

Worldwide Distribution

India, Myanmar, Thailand, Malay Peninsula, China, Afghanistan and Nepal

Remarks

It was reported by Balthasar (1963b) and Syed et al. (1981) from the foothills areas of Pakistan. Arrow (1931) recorded this species from Myanmar, Srilanka and India. The species is distinct on the basis of middle tibia elongate and curved along with double or single tooth near the base beneath. This species shows wide distribution in the country. Good tolerance for shade. Good cow dung burial. It is also widely distributed in the forests. It attracts towards light and feeds only on herbivore dung of cattle, fresh or old, appears to be more common in dung heap accumulated for manure and prefers open grasslands and cultivated field. It prefers fresh and old dung ranging from 1-14 days and there- after its population declines.

Figure 22.1: Onitis subopacus

Figure 22.2: Distribution map of Onitis subopacus

3.6.7 Onitis virens Lansberge, 1875 (Figs. 23.1 & 23.2) Onilis virens Lansberge 1875, Ann. Soc. Ent.• Belgium, 18 : 135. (ISNB) Onitis virens. Arrow 1931, Fauna British India Ceylon and Myanmar (Coleoptera: La- mellicornia: Coprinae)3 : 396-397, PI. XII, Fig. 20& 21. Onitis virens. Balthasar 1963, Monog raphie der Scarabaeidae und Aphodiidae der Pal- aearktischen und Orientalischen Region ( Coleoptera : Lamellicomia), 2 : 40.

Type Repository

Belgium, Brussels

Diagnosis

Length: 18 mm to 22 mm. Oval, moderately convex and smooth in shape. Clypeus ellip- tical, closely transversely rugulose, separated from frons by an intervallic carina. Just be- fore the interrupted carina have short transverse clypeal carina and just behind it a conical tubercle. Prothorax fully punctured with a median line in front, narrow median groove behind, on both side of middle of basal margins has deep longitudinal pit and elytra distinctly punctured intervals flat and striate. The pygidium flat and punctured and the metasternal shield smooth. Front tibia elongate slandered curved with a long spine at extremity but short broad.

Material Examined

4♂ Kohat, 26.VII.2005; 1♂ 5♀, Mansehra 29.VI.2006; 2♂ 3♀, Abbottabad 20.V.2009; 2♂ 3♀, Nowshera 12.V.2010; 2♂ 4♀, Islamabad 02.VII.2012; 3♂4♀, Mardan 03.VII.2012; 4♂ 5♀, Peshawar 13.VIII.2013.

Worldwide Distribution

Bangladesh, India, China, Myanmar, Belgium, Tonkin (Vietnam), North Vietnam and Laos. Remarks

Arrow (1931) and Syed et. al. (1981) reported that specimens from foothills areas of Pa- kistan. Identified specimens were distinct on the basis of front tibia elongate having blunt spine and middle femur bears strong rounded lobe at middle trochanter of hind leg sharply toothed. Present in dung at open pastures. It attracts towards light and feed only on herbivore dung of cattle, lives in sandy muddy soil, rare on rocks and not common in accumulated dung heap kept for manure but does not prefer human dung . It prefers dung of 3-7 days old dung but may also remain in the dung up to 14 days but occasionally. It shows close resemblance to O. subopacus and can be differentiated from the latter on examining the characters of protibiae. In this species, 4th tooth of protibia blunt and joined with 3rd tooth and lower edge of protibia at end serrated while in O. subopacus protibiae tridentate with a terminal blunt process and lower edge of it with a minute tooth near base beneath.

Figure 23.1: Onitis virens

Figure 23.2: Distribution map of Onitis virens 87

3.7 Genus CHEIRONITIS Lansberge, 1875

Oblong and rather depressed. In males prolegs elongate, femur with one or more sharp teeth at front edge, protibiae quadridentate with its extremity produced, tarsi and spur absent. In females prolegs stout, tibiae broad, quadridentate, not produced at end, tarsi very short, spur articulated. Meso and meta tibiae strongly dilated at extremities. Meso and meta tarsal segments progressively diminish in size first segment more than twice as long as 2nd. Anterior margin of clypeus reflexed and excised in middle. Pronotum transverse, base margined with a slight impression on both side in middle. The Scutellum relatively less smaller and severely pointed and elytra covering abdomen, smooth or hairy at extremity with simple narrow epipleura. Procoxae very large and prominent; abdomen with a continuous carina around sides and pygidium. This genus is Afrotropical, Oriental and Palaearctic.

3.7.1 Cheironitis arrowi Janssens, 1937 (Figs. 24.1 & 24.2) Cheironitis arrowi, Bezdek and Krell, 2006, 3: 158. Chironitis arrowi Janssens, 1937, 11: 153, 159 - Balthasar, 1935, 115: 86 - Balthasar, 1963, II: 14, pl. 1 Chironitis indicus, Arrow (non Lansberge, 1875), 1931, III: 402.

Type Repository

Unknown

Diagnosis

Length: 9.2 mm to 10 mm . Oblong and rather depressed shape. Anterior margin of clypeus strongly feebly bilobed, reflexed, separated from frons by a short, transverse, clypeo- frontal carina with another carina behind it. Frons with some short erect setae. Pronotum broader than elytra unevenly rugose with very irregular punctures, coarsely and partly confluent in middle, finely and sparsely at lateral sides. Elytra very lightly striate; 2nd

and 4th intervals broad rather flat and 3rd and 5th narrow raised and rather sharply cari- nate. Pygidium lightly and sparsely punctate. Metasternum longitudinally grooved, ante- riorly clothed with erect hairs, very minutely and sparingly punctuate; lateral sides of metasternum and abdomen clothed with fine setae.

Material examined

2♂, Malakand, 26.II.2005; 3♀, Mansehra, 29.VIII.2006; 2♂3♀, Swat, 20.VII.2010; 1♂2♀, 2♂1♀, Islamabad 02.IX.2012; 3♂, 4♀, Mardan 27.0I.2012; 4♂5♀, Murree, 13.VIII.2013.

Remarks

Sexually dimorphic species. Clypeus shining and is finely and sparsely punctuate. The specimens were found from fresh cow and buffalo dung in the mountainous environments. Specimen was previously recorded from Pakistan, India, Afghanistan and Arabia (Balthasar 1963).

Figure 24.1: Cheironitis arrowi

Figure 24.2: Distribution map of Cheironitis arrowi

3.8 Genus CATHARSIUS Hope, 1837

Head is semicircular and broad and the body is broad, compact and very convex in shape. Fully margined prothorax at the base and long parallel middle coxae. Hind tibia shows one and The middle tibia shows two terminal spur. The hind and middle tibiae are completely widened from the base to extremity. Elytra with seven striate completely covering the abdomen.

3.8.1 Keys Species of Genus CATHARSIUS Hope, 1837

1. Head without smooth area adjoining each eye. Female pronotum with transverse ridges, but the transverse ridge without teeth; Head very transverse; pronotum of male with two tubercles. Cephalic horn of the large males long, thin, and distinctly erect, and the apical half occasionally bent somewhat forward. Pronotum of the female simple, and as in the small males, granulate all over. Black and shining ..………………………….………………..Catharsius pithecius (Fabricius, 1775)

1” Head with slight smooth area adjoining each eyes. Clypeus closely transversely rugulose. In front of eyes, an actually small, but striking, smooth and shining area …………...... C. molossus (Linnaeus, 1758)

3.8.2 Catharsius pithecius (Fabricius, 1775) (Figs. 25.1 & 25.2) Scarabaeus pithecius Fabricius, 1775, Syst. Ent., 1: 21 (BMNH). Scarabaeus nanus Fabricius, 1792, Ent. Syst., 1: 42 (ZMUC). –Gillet, 1911, Col. Cat. Pars. 38, Scarabaeidae: Coprinae I, : 69 (Syn.). Scarabaeus sabaeus Fabricius, 1781, Spec. Ins., 1: 23 (BMNH). Copris sinensis Hope, 1842, Proc. Ent. Soc. Lond.: 60 (BMNH). –Gillet, 1911, Col. Cat. Pars. 38, Scarabaeidae: Coprinae I, : 69 (Syn.). Catharsius pithecius: Gillet, 1911, Col. Cat . Pars. 38, Scarabaeidae: Coprinae I, : 69.

Type Repository

British museum.

Diagnosis

Length: 14 mm to 22 mm.. The head is nearly semi-circular and in male head is lightly sculptured and in female head is more deeply sculptured. The clypeus very feebly expur- gated in middle of the front margin. Elytra extremely but clearly striate with hardly punctures in the striate and the pygidium finely and not very closely punctured and the metasternal shield angular in front, very smooth and shining and both side of the meta sternum closely and rugosely punctured.

Material Examined

4♂ 1♀, Islamabad, 02.VII.2005; 2♂ 5♀, Peshawar, 13.VII.2007; 3♂ 2♀, abbottabad, 18.VII.2008; 1♀, Nowshera, 20.IX.1980; 2♂, Buner, 16.VIII.2009; 5♂2♀, Kohat, 28.VII.2012.

Worldwide

India, Sri Lanka, china and Taiwan.

Remarks

This specie was found in loams, pastures and forest edges at higher elevations and also abundant in cattle dung.

Figure 25.1: Catharsius pithecius

Figure 25.2: Distribution map of Catharsius pithecius

3.8.3 Catharsius molossus (Linnaeus, 1758) (Figs. 26.1 & 26.2) Scarabaeus molossus Linnaeus, 1758, Syst.Nat. Ed., 10: 347 (LSLC). Scarabaeus abbreviatus Herbst, 1789, Nat.Kafer, 2: 53, t.8, f.10.-Gillet, 1911, Col. Cat.Pars. 38 Scarabaeidae: Coprinae I,: 69 (Syn.). Scarabaeus berbiceus Herbst, 1789, Nat. Kafer, 2: 227, t.16, f.1.2.-Gillet, 1911, Col. Cat. Pars. 38. Scarabaeidae: Coprinae I,: 69 (Syn.). Catharsius janus Olivier, 1789, Ent. 1 (3): 101, t.26, f.227.-Gillet, 1911, Col. Cat. Pars. 38 Scarabaeidae: Coprinae I,: 69 (Syn.). Copris ursus Fabricius, 1801, Syst.Eleuth., 1: 43.-Castelnau, 1840, Hist. Nat. Col., 2: 77 (syn.). Catharsius molossus: Harold, 1877, Ann.Mus. Civ. Genova, 10: 44. Catharsius davidis Deyrolle et Fairmaire, 1878, Ann.Soc. Ent. Fr., 8 (5): 95. (New Syn.) Catharsius timorensis Lansberge, 1879, Com.Séa. Soc. Ent. Belg., 22: 148.-Balthasar, 1963, Monogr. Scarab. Aphod. Palaearkt. Orient. Reg., 1: 307 (Syn.). Catharsius dayacus Lansberge, 1886, Tijdschr.Entom., 29: 6.-Balthasar, 1963, Monogr. Scarab. Aphod. Palaearkt. Orient. Reg., 1: 307 (Syn.) - Ochi et Kon, 1994, Elytra, 22 (2): 283 (removal from syn.). Catharsius dubius Paulian, 1936, Treubia, 15: 396 (n. nud.). Catharsius borneensis Paulian, 1936, Treubia, 15: 396 (n. nud.). Catharsius kangeanus Paulian, 1936, Treubia, 15: 395.-Balthasar, 1963, Monogr.Scarab. Aphod. Palaearkt. Orient. Reg., 1: 307 (Syn.).

Type Repository

Unknown

Diagnosis

Length: 22 mm 30 mm. The head is broad with front margin almost imperceptibly excised in the middle. The clypeus closely transversely rugulose. Pygidium finely punctured in the middle and finely granular at the sides and metasternal shield acutely angular in front and rather sparsely punctured, rather smooth and shining. The sides of metasternum very densely clothed and the front tibia bears three external teeth, the middle and hind tibiae are broad and the terminal spur of the hind tibia is truncate at the extremity.

Material Examined

4♂1♀, Swat, 29.VII.2005; 1♂, Dir, 04.VIII.2006; 3♂ 4♀, Swabi, 09.IX.2007; 4♂ 3♀, Mardan, 12.VI.2010; 2♂, Mansehra, 08.VII.2012; 4♂ 5♀, Chitral, 12.VIII.2012; 4♂ 5♀, Abbottabad, 13.VII.2013.

Worldwide Distribution

Sri Lanka, Tibet, Vietnam, Laos, Cambodia, Thailand, India, Nepal, Afghanistan, Indo- nesia, Malaysia, Timor, Hong Kong and China.

Remarks

Recorded from Pakistan by Hashmi and Tashfeen (1992). The species is abundant in cattle dung and can be collected at light. It is found in sandy to heavy loams in grasslands, shrub lands and pastures. Wide spread and versatile species. After more than Catharsius davidis Deyrolle et Fairmaire mode and Catharsius molossus (Linnaeus, 1758) naming specimens (BMNH), authors found Catharsius davidis for Catharsius molossus a new synonym.

Figure 26.1: Catharsius molossus

Figure 26.2: Distribution map of Catharsius molossus

3.9 Genus METACATHARSIUS Laporte, 1840

Clypeus rounded at the front, with emargination barely indicated. Upper side entirely smooth and shining; the elytra also more or less shining. Meso-etasternal line straight; pronotum granular at the sides only.

3.9.1 Metacatharsius inermis (Laporte, 1840) (Figs. 27.1 & 27.2) Catharsius inermis, Arrow 1931, Fauna British India Ceylon and Burnt(Co1eoptera: La- mellicomia: Coprinae),3: 101. Catharsius inermis, Balthasar1963 Monographie der Scarabaeidae und Aphodiidae der Palaearktischen und Orientalischen Region(Coleoptera: Lamellicornia),1: 316·317.

Type Repository

Unknown

Diagnosis

Length: 20 mm 28.5 mm. Head is very broad and transversely rugose with granular ocular lobes, clypeal margin rounded feebly excised in middle and frons bear a short tubercle between eyes. Prothorax punctured with a slight median longitudinal groove upon basal half and the hind angles obliterated and front angles broadly rounded. Elytra with fine punctures and the pygidium closely and strongly punctured. Metasternal shield angulate in front sparsely punctured.

Material Examined

2♂ 2♀, Mansehra, 13.VIII.2006; 4♂ 2♀, Chitral, 12.VII.2007; 3♂ 2♀, Allai, 18.VI.2008; 2♂ 3♀, Abbottabad, 23.VII.2010; 1♂ Peshawar, 12.VI.2012; 4♂, Nowshera, 14.VIII.2013.

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Worldwide Distribution

India, Africa, Ethiopia and Arabia.

Remarks

Arrow (1931) recorded previously from Pakistan. This specie was found in pastures and found abundant in cattle dung.

Figure 27.1: Metacatharsius inermis

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Figure 27.2: Distribution map of Metacatharsius inermis

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3.10 Key to the collected genera of Tribe Scarabaeini

1. Front tarsi absent, elytra not excised behind the shoulder …………………………………………………….… Scarabaeus Linnaeus, 1758

1” Front tarsi present, elytra excised behind the shoulder …………………..…….. ………………………………………………………...Gymnopleurus Illiger 1803

3.10.1 Genus GYMNOPLEURUS Illiger 1803

The clypeus is entirely merged with the ocular lobe and gives 2 or 4 short teeth at the front margins. Short and stout forelegs. The convex and large prothorax is dorsally deco- rated with the several patches. Three external teeth are alongwith the front tibia. Visible scutellum and 6 tarsi are filiform.

3.10.2 Gymnopleurus flagellatus (Fabricius, 1787) (Figs. 28.1 & 28.2) Scarabaeus flagellatus Fabricius, 1787, Mant. Ins., 1: 17 (Type specimens transfered from ZMUK to ZMUC). Copris flagellatus: Olivier, 1789, Ent., 1(3): 162; 1790: 174. Scarabaeus coriarius Herbst, 1789, Nat. Kafer, 2: 309. Ateuchus flagellatus: Fabricius, 1801, Syst. Eleuth., 1: 59. Gymnopleurus flagellatus: Illiger, 1803, Mag. Ins., 2: 199. Gymnopleurus serratus Fischer von Waldheim, 1821, Lettre a Pander, : 11. –Olsoufieff, 1918, Mém. Mus. Cauc., 7(A): 38, 77 (Syn.). Gymnopleurus clypeolatus Mulsant, 1842, Col. France, Lamell., : 58. Gymnopleurus confusus Mulsant, 1842, Col. France, Lamell., : 57. –Olsoufieff, 1918, Mém. Mus. Cauc., 7(A): 77 (Syn.). Gymnopleurus asperatus Mulsant, 1842, Col. France, Lamell., : 58 Gymnopleurus rugulosus Mulsant, 1842, Col. France, Lamell., : 58. Gymnopleurus suturalis Mulsant, 1842, Col. France, Lamell., : 58.

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Gymnopleurus variolosus Motschulsky, 1849, Bull. Nat. Mosc., 22(3): 102 (ZMUM). Gymnopleurus conflagratus Motschulsky, 1849, Bull. Nat. Mosc., 22(3): 102 (ZMUM). Gymnopleurus hornei Waterhouse, 1890, Ann. Mag. Nat. Hist., 5 (6): 410 (BMNH). – Arrow, 1931, Fauna Brit. India, Lamell., 3: 55 (Syn.). Gymnopleurus minor Seabra, 1907, Estud. Anim. Ut. Agric., 4: 75. Gymnopleurus rufipes Seabra, 1907, Estud. Anim. Ut. Agric., 4: 75. Gymnopleurus barovskyi Kiseritzky, 1928, Izv. Kur. Prikl. Zool. Fit., 4: 45.

Type Repository

Copenhagen Museum

Diagnosis

Length: 19 mm 22.5 mm. The head is coarsely and confluently punctured with two strongly elevated oblique bisinuate carinae converging behind entirely black dull and coarsely sculptured above. The clypeus is notched in the middle and forms two rounded lobes in front and the pronotum is very coarsely and closely pitted with an incomplete median smooth longitudinal line. Sides are straight in front and strongly rounded behind with the acute front angle, very obtuse hind angle and very feebly curvy base. Slightly vaguely striate, with the interval broken into irregular shining elevations and opaque de- pressions.

Material Examined

3♀, Kurram Agency, 14.VI.2007; 1♂ 2♀, Dir, 19.VI.2008; 2♂, Kohat, 26.VIII.2010; 2♂, 1♀, Khyber Agency, 12.VII.2012; 2♂ 1♀, Bajaur Agency, 12.III.2012; 4♂ 2♀, Mohmand Agency, 23.IV.2013.

Worldwide Distribution

104

Afghanistan, Iran, Turkey, Syria, Palestine, Morocco, Turkestan, Spain and S. France.

Remarks

Previously recorded by Arrow (1931) in cattle dung at open pastures. The species is most active from late spring to summer. This species is the eastern-most distribution.

Figure 28.1: Gymnopleurus flagellates

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Figure 28.2: Distribution map of Gymnopleurus flagellates

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3.10.3 Scarabaeus sacer Linnaeus, 1758 (Figs. 29.1 & 29.2) Scarabaeus sacer L., Syst. Nat. ed. X, 1758, p. 347; Reitt., Verh. Ver. Brunn, xxxi, 1893, p. 41. Ateuchus sacer F., Syst. Eleuth. I, 1801, p. 54. Scarabaeus sacer Linoaeu 1758. s, Syst. Nat., 10 : 347. Scarabaeus sacer- Asrp w, Faun a of British India including Ceylon and Myanmar (Col- eoplera : Lamellicoroia : Coprioae), 3 ; 40-41, pI. n, Fig. 7. 8 Rec. zool. Surv. India. Occ. Paper No. 296 1963. Scarabaeus sacer- BaKthasar, Monographie der Scarabaeidbe und AphodUdae der Palaearktisc hen und Orientalischen Region (Coleoptera : Lanellicomia), 1 : 145-161.

Type Repository

Unknown

Diagnosis

Length: 21 mm to 26.5 mm. Head is densely and rugosely punctured in front and tubercu- late from behind behind. Frons with a pair of tubercles placed transversely between the eyes. Clypeus with four teeth separated by rounded notches. Pronotum is unevenly punctured and the lateral margins are strongly rounded and closely denticulate. There is no visible scutelluma and the elytra are lightly striate and lightly punctured. The front tibia bears four strong external teeth and two small teeth and the tarsi are inserted at a distance from the middle and hind tibiae.

Material Examined

1♀, Kohat, 12.VII.2007; 1♂ 2♀, Kurram Agency, 16.V.2010.

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Worldwide Distribution

Egypt, France, Hungary, northern India. Persia, Romania, Portugal, Russia, Spain, Tunis, Northern Sudan, Palestine, France Turkestan, Iran, Israel, Afghanistan, The whole of North Africa; and Ethiopia.

Remarks

This is the beetle generally represented in ancient Egyptian sculpture. Scarabaeus sacer and Scarabaeus typhon are confused with each other. Scarabaeus sacer pronotum has the sides basally a circular area devoid of granulating or punctuation. Scarabaeus sacer also has with two strong tubercles in frontal suture. Scarabaeus typhon pronotum is all granules or dotted, smooth with no spaces on either side at the base. It is abundant in open, non- forested areas in the foothills and valleys.

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Figure 29.1: Scarabeus sacer

109

Figure 29.2: Distribution of Scarabeus sacer

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3.11 LIST OF THE SPECIES

1. Digitonthophagus gazella (Fabricius, 1787) 2. Digitonthophagus bonasus (Fabricius, 1898) 3. Onthophagus (Palaeonthophagus) marginalis (Gebler, 1817) 4. Onthophagus (Palaeonthophagus) nuchicornis (Linnaeus, 1758) 5. Onthophagus (Phanaeomorphus) gagates Hope, 1831 6. Onthophagus (Altonthophagus) kukunorensis Kabakov, 1990 7. Onthophagus (Coloboonthophagus) tragus Fabricius 8. Onthophagus (s. str.) proletarius Harold, 1875 9. Onitis falcatus (Wulfen, 1786) 10. Onitis philemon Fabricius, 1801 11. Onitis virens Lansberge, 1875 12. Onitis lama van Lansberge, 1875 13. Onitis subopacus Arrow, 1931 14. Catharsius molossus (Linnaeus, 1758 15. Catharsius pithecius (Fabricius, 1775 16. Metacatharsius inermis (Laporte, 1840 17. Copris repertus WALKER, 1858 18. Corpis sacontala L. Redtenbacher, 1844 19. Copris punctatus Gillet 1910 20. Heliocopris bucephalus (Fabricius, 1775) 21. Heliocopris midas (Fabricius, 1775) 22. Gymnopleurus flagellatus (Fabricius, 1787) 23. Scarabaeus sacer. Linnaeus, 1758 24. Cheironitis arrowi Janssens, 1937

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4. CONCLUSION AND RECOMMENDATIONS

In the present study, 4250 specimens of the dung beetles belonging to subfamily scarabaeinae of family scarabaeidae were collected from northern parts of Pakistan and examined for morphological characters, which yielded the presence of 24 species in ten genera and twenty-four species of dung beetles in subfamily scarabaeinae. These include Digi- tonthophagus gazelle, D. bonasus, Heliocopris midas, H. bucephalus, Onthophagus (Phanaeomorphus) gagates, O. (Altonthophagus) kukunorensis, O. (Palaeonthophagus) marginalis, O. (Palaeonthophagus) nuchicornis, O. (s. str.) proletarius, O. (Colo- bonthophagus) tragus, Onitis falcatus, O. lama, O. philemon, O. subopacus, O. virens, Catharsius molossus, C. pithecius, Metacatharsius inermis, Copris repertus , C. sacontala, C. punctatus, Scarabaeus sacer, Cheironitis arrowi and Gymnopleurus flagellates. Out of these, eight species were newly recorded for Pakistan including Onthophagus gagates, Onthophagus kukunorensis, O. (s. str.) proletarius, Heliocopris bucephalus, Copris punctatus, C. repertus, C. sacontala and Onitis lama. The new distributional and temporal data of the dung beetles with notes on their descriptions was also provided. For each taxa, the synonymies were also provided. The diagnosis of each genus, identification keys to genera and species, colored photographs and geographical distribution maps for each species were also provided. The hypothesis of the present studies on the expected diversity of dung beetles in northern Pakistan stands true.

The most diverse genus of the study area was Onthophagus, represented by 7 species, followed by Onitis (5 species), Copris (3 species); Heliocopris, Digitonthophagus and Catharsius represented by 2 species each and Metacatharsius, Gymnopleurus, Scarabae- us and Cheironitis represented by single species each. Modern systematic approach was followed to recognize 3 subgenera in genus Onthophagus, comprising of subgenera Pal- aeonthophagus, Phanaeomorphus, Altonthophagus and Coloboonthophagus.

The species of genus Onitini strongly attracts towards light and are among the first to come in the light trap. The species in this tribe are tunneller. In spite of rolling away the dung balls for brood construction, they bury the dung balls in vertical chambers in prox-

112

imity to original deposition or under the dung pats. The species in the genus Onitis prefer herbivore dung (cow, buffalo, pig and horse) but O. philemon attracts towards human dung while showing less preference. O. philemon, O. virens and O. subopacus are distributed widely in northern Pakistan.

Dung beetles are prone to habitat changes such as deforestation or shifts in the mammali- an fauna and are considered to be useful indicators of ecosystem health. The study made by Mittal (2005) noticed decline in the richness and diversity of these beetles in North India because of the loss of habitat, altered food quality, pollutants and increased cattle antibiotics and other environmental changes such as the rising atmospheric temperature.

Recommendations

The northern parts of Pakistan at the amalgamation of Palearctic and Oriental realms is biologically a diverse region. The occurrence of presently recorded 24 dung beetles species with 8 new distributional records indicates very clearly that more collection efforts and intensive surveys can bring about more accomplishments in terms of new country records or species new to science.

The future studies needs to be carried out mainly focusing on reproductive biology, conservation biology and phylogenetic analysis of the group for solving more biological questions.

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5.2. Web References

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5.3 Publication (International)

 Abbas M., Ming Bai and Xingke Yang. 2015. Study on dung beetles (Coleoptera: Scarabaeidae:Scarabaeinae) of northern Pakistan with a new record from Pakistan “Entomotaxonomia” (2015) 37(4):–, DOI: 10.11680/ entomo-tax.2015044, ISSN 2095–8609 1 (Published)

 Abbas M., Ming Bai Hakeem A. and Xingke Yang. 2016. New Record of Onitis lama (Coleoptera: Scarabaeidae: Scarabaeinae) from Pakistan “En- tomo-taxonomia” ((proof reading near to submission)

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