ECOLOGICAL ACCOUNT OF THE MAMMALS OF RAJASTHAN DESERT

TAXONOMICAL AND ECOLOGICAL ACCOUNT OF THE MAMMALS OF RAJASTHAN DESERT ISHW AR PRAKASH Gentral Arid Zone Research Institute, Jodhpur ( Continued from p. 142, Vol. I, No.2) ADJUSTMENT OF MAMMALS TO DESERT ENVIRONMENT Diurnal fluctuations of temperature and humidity, sparse vegetation, scanty rainfall, loose sand and strong winds are some of the important factors which animals 0f the desert have to contend with. Adaptations to desert conditions are anatomical as well as behavioral. The former is beyond the scope of this paper and only the latter is discussed here. TEMPERATURE-Toavoid the high day temperatures and the consequent body desiccation, the smaller mammals have taken to a nocturnal and subterranean life (Allee et al, 1950 ; Dice, 1952). Of the 38 species and subspecies of mammals like Artiodactyles and primates a few small mammals such as squirrels and desert geroils are diurnal. Nineteen (50 %) species and subspecies are fossorial, while eight types of bats live in caves, six live near water and four take shelter on trees where the temperature is considerably lower than on the ground and in the open. - The temperature inside the burrow of the gerbils was found to be lower than outside (Prakash et al, 1962). Schmidt-Nielsen and Schmidt-Nielsen (1950) also record a difference of about 10 F. inside the burrow of Dipodomys in the Arizona desert. They found in a single diurnal den, a very high relative humidity.!average 84%, range 62-91 %). The daily activity rhythm of the diurnal mammals is greatly influenced by day temperatures and they adjust their activities accordingly. In summer, the desert gerbils, Meriones hurrianae come out of their burrows just after dawn and retire after a few hours, before it gets too hot. They again come out late in the evening and retire before dusk. In winter, however, they are out of their burrows through~ut the day but not during mornings and evenings which are quite cold (Prakash, 1962). Kashkarov and Kurbatov (1930) have observed similar adjustments in the daily life of the gerbil, Rhombomys and ground squirrel, Shermophilopsis in tr.e Central Kara Kum desert. Schmidt-Nielsens (1952) also observed similar shifting of activities in the ground squirrel, Gitellus in the Arizona desert.

MAMMALS OF RAJASTHAN DESERT 151 During very hot days, the fruit bats, Pteropus 'giqanteus giganteus climb down.to the shade of leaves of Fious trees and fan themselves with their wings (Prakash, 1960). In winter, however, they are found on the tree top throughout the day, basking in the sun. Black bucks and gazelles trek long mornings in search of their food but return to their pond during the day. In the evening they again start their feeding rhythm. The burrowing animals are also sensitive t.o strong winds and try to avoid, them, in which their fossorial habit is an advantage. The burrow openings of hedgehogs and gerbils are, therefore..found in the direction opposite that of normal winds. FOOD AND WATER-Food is one of the primary. scarcity iteml! of a desert. Due to low rainfall the desert is poor in the vegetation. This has initiated the habit of storing food when it is available in mammals of other deserts. But it is interesting to note that with the sole exception of Tatera. indioa indioa no other Indian desert mammal stores food material. Mammals are found around the following six food centres:- 1. Vegetation-Nineteen species and sub-species of mammals depend on vegetation. The jackal and fox are occasional and are grouped almost in every centre. The flying fox, primates, ardiodactyles, hares and rodents are the chief dependents of vegetation. Rats and gerbils, feed sometimes on insects in summer. Most of the larger mammals restrict themselves to vegetation around lakes and ponds. 2. Inseots-They are the busiest foc>dcentre. Twenty species and subspecies of animals are grouped around this centre. Hedgeh.ogs, shrews; bats and pangolin depend mainly on insects, The sq' irrel and the desert gerbil are only occasional intruders.in this centre. The false vampire bat (Prakash; 1959 and 1959 c), foxes, mongoose are primarily grouped in other centres but their st9mach contents also show quite a large variety of Scorpion (Palemneu'9 sp.) and spider, (Galeodes agilis) not included in Insecta are worth mentioning since some of the stomachs of cats and foxes were full of these.arachnids. 3. Lizards-All lizards (except the Fat-tailed lizard, Eublepharus maoularia and Ophiomorus tridaotylus) are diurnal and some of them are fossorialliving in shallow burrows. While resting during night, they are easijy ex,c<l:vatedby hedgehogs, foxes, cats and mongoose. The false vampires also feed on them. 1;'he lizards which are, usually consumed are: Stenoda.otylus orientalis, Gymnoq,actylus scaber, Varanus griseus, Eublepharus maoularia, Mabuya aurata, Mabuya mao/daria, and Uromastix hardwickii. 4. Birds and Eggs-A large variety of birds lay eggs in shallow pits on the

152 ISHWAR PRAKASH ground and over bushes making them easily accessible. It is not an uncommon 5cene to witness an attacked cluster of eggs. Eggs of some, like game birds, partriges and sandgrouse are also destroyed by mammals. Middleton's (1935) analysis showed that the mammals are generally responsible for the destruction of eggs. It is worth examining such losses in the Rajasthan Desert since about 25% of the mammals destroy eggs. 5. Gerbils and hares-both are quite prolific in the desert. Mer iones is diurnal while Tatera is nocturnal. The carnivores depend on this food centre. Panther depends on artiodactyles and sometimes on hares. Artiodactyles have not been given a separat": status of a food centre since only one species; Panthera pardu8 is linked to them; moreover their number is limited. 6. Carrion-Man is responsible for keeping large number of cattle, goats and sheep. The unused parts of the dead animals are thrown away and the flesh and carrion is eaten by lhe jackal and the hyaena. Animals have to adapt themselves more to the scarcity of water than that of the food as the former is a more acute problem. Cattle have to trek long distances in search of water. Certain mammals, gerbils in particular, do not drink water or take it at very long intervals. During the intervening period, they depend on their food (vegetation or animal) for the necessary water supply. The gerbils, Tatem indica indica, Meriones hul"rianae; hare, Lepus nigr icollis dayanu8; squirrel, Funambulu8 penna1bti,and porcupine, Histrix indica in:lica, for instance. live on dry seeds, roots and stems of desert plants Similar observations have been reported for the mammals of other deserts of the world (Buxton 1923, Kashkarov & Kurbotov 1930 & Schmidt-Nielsens 19.32). Another very important source providing large amount of water consists of insects, particularly the larvae and c?terpiliars. They are full of watery juice and meet the water requirements of the animals which feed on them. Robinson ( 1928) gives the percentage of water contents in the body of insects, which varies from 41) to 92% of the entire body. This may be one of the reasons why the desert gerbils also feed on locusts during summer (Prakash, 1962). The large mammals (black buck, gazzelle, wild boar and nilgai) do not, u:mally, go very far off from the water source and hence are not found deep in the waterless tracts of the arid region; water, therefore, becomes a factor restricting their activities around the aquatic bodies. The economie use of water is as important as is the need for obtaining it. Desert animals show various adaptations for it. As most of them are nocturnal in habit when it is generally cool and the humidity is higher than what it is during the day, they are able to prevent water loss from their bodies. The fossorial habitat of the mammals is also helpful in economising the loss of body water. During the day some of them plug the openings of their burrows to avoid the heat which causes

MAMMALS OF RAJASTHAN DESERT 153 loss of water. There is considerable difference in temperature of the outer surface and that inside the burrow. When an animal respires, it gives out certain amount of moisture which is retained in burrow and helps in maintaining higher humiduty. The urine passed in the tunnel is also a helpful factor. There is less loss of water when the animal is inside the burrow, than when if stays out in the hot day. According to Schmidt-Nielsen and Schmidt-Nielsen (1952) other means of meeting the requirement of water by the desert rodents is that gained from oxidation of food stuffs ( oxidation or metabolic water ). BIOTIC ASSOCIATIONS In addition to the factors mentioncd above, lack of adequate shelter is an unfavourable desert factor. There are not many types of habitats or biotic associations in the Rajasthan desert- Blatter and Hallberg ( 1918-21) recognised five formations in this desert. They have defined a formation as "The community of plants as determined by the qualities of the soil" Krishna & Dave ( 1956) have classified Rajasthan desert into three subregions according to the climatological data and occurrence of reptiles: 1. Arid Subregion :-- RainfuU 5", N:)rth western portions of Jaisalmer district. 2. Semi-arid 8ubregion :- Rainfull 5 to 10", Central Desert; 3. Sub-humid 8ubregion:- Rainfall 10- IS", South-East Barmer, North- East Jodhpur, Ganganagar. Sikar, and Jhunjhunu districts. No mammal is restricted to anyone of the subregions suggested by these authors. Most of the mammals are found in all the subregions. Another reason for doubting the validity of their classification is that in the desert the Biotic districts (Cockrum, 1953) are so inter-mixed that it is not easy to demarcate them geographically on the basis of the mammalian fauna. The following associations and sub-associations have been recognised in the Rajasthan Desert. The occurrence of these mammal associations, in most cases, coincides with the landscape, i.e with the physiographic or the edaphic conditions and with the plant associations which are edaphically controlled. I. Sandy Association: (a) Sand dunes (b) Sand hills (c) Sandy plains (d) Artificial Mines. II. Rocky Association: (a) Rocks (b) Lakes and surrounding gardens (c) Caves and Tunnels. III. Ruderal Association: These three associations are found in each of the subregions of Krishna and Dave (1956). Each association is characterised by its vegetation. landscape and soil formation.

154 ISHWAR PRAKASH I. Sandy Assoc'iation: A major part of the desert region is covered by this association. The sand consists mainly of fine grains of quartz, flakes of horneblende and felspar, as well as fragments of local locks. This association is dividetl into three sub-associations. At certain places all the three are adjacent to one another but at other places they are wide apart. I. (a) SAND DUNES General: This sub-association is usually unstable and is formed by aeolian sands. The dunes movements are more from April to July, particularly in May and June. These dunes are sometimes 20 to 30 ft. in height. Mammal Association: After the rainy season when the sand dunes,become stationary and harder, the desert gerbils, Meriones hurrianae dig their tunnels which are usually; associated with the creepers of Cdrullus colocynthis. Hedgehogs, foxes and hares visit the dunes in search of food but they do not generally make tunnels or dens in this habitat. The sand dunes are also inhabited by the Scincid lizard, Mabuya macularia and M. aurata, These form one of the food items of hedgehogs and foxes. The beetles, Bucephalus helicopris, Anthia sexguttata, Juliodes atkinsoni and the scorpion, Palaemneus eaten by these mammals. sp. which visit the dunes in search of food are also (b) SAND HILLS General: These are the large stable hills formed by aeolian sands which have become stabilized. Perennial plants are found growing over them. The composition of the sand is the same as that of the dunes. At Munabao and Gadra Roaa they attain a height of 150 ft., but are small in other places. Mammal Association: The desert gerbil is the commonest mammal to be found in this habitat. Their burrow system is extensive and causes considerable damage to the sand by loosening the soil which is easily eroded by wind. The Sind hare, Lepus n. dayanus in the Jodhpur division and the Indian hare, Lepus n. ruficaudatus in Bikaner, Jhunjhunu and Sikar districts are found in this shrubby region. The foxes Vulpes v. pusilla and V. bengalensis and cats, Felis chaus prateri and F. libyca ornata dig their tunnels through the sand hills. Large mammals also visit the sand hills but do not venture far from their water source. The lizard Mabuya sps., Varanus griseus, Stenodactylus orientalis, Gymnodactylus scaber and the nocturnal, fatty-tailed lizard, Eublepharus macularis inhabit this sub-association and form the food of the mammals. Several types of snakes have also been collected and observed from this sub association. They are Eryz johnii, Zemenis mucosus, Echis carinata, Bungarus sindanum and B. caeruleus. Snakes influence the

MAMMALS OF RAJASTHAN DESERT 155 lives of the gerbils by feeding on them. The snakes, in turn, are eaten by the mongoose, foxes and hedgehogs. Common insects of the sandy hills which form part of mammal food are Batocera ruba, Blaps orientalis, AntMa sexguttata, Juliodes atkinsoni etc. Some birds, {>articularly sand grouse and partridges lay their eggs on the ground. Their eggs as well as nestlings form a great deal of mammalian food. All these animals, though unrelated taxonomically form a close ecological association. General: sand particles. (c) SAND PLAINS The sand plains may be composed of either fine or coarse ground Mammal Association: Oapparis and other Jow; prostrate bushes and herbs form a sort of shelter and a variety of animals are found living in them, forming an ecological community. The burrows of hedgehogs are situated under these bushes, and those of hedgehogs, cats, foxes, and mongoose are found in the open plains. The snakes, Ooluber diadema, Oligodon taeniolatus, Lycodon striatus, PsammopMs schokeri, P.lethi, Bungarus caeruleus. Echis carinata and the lizards Mabuya sps. Stenodactylus orientalis Gymnodactylus scaber, Varanus griseus, Eublepharus macularis. Uromastix hardwickii influence the life of mammals in the sandy plains of the Rajasthan Desert. Insects are also quite common during and after the rainy season. The ants, Dorilus labiatus. Monmorium indicum; beetles, AntMa sexguttata, Endema angulatum, Juliodes atkinsoni, Batosera ruba, Helicopris buciphalus, Onthophag1J,8 longicornis Oniticellus cinctus; the hoppers, Bchistocerca gregaria, Hieroglayhus negeroriphletus, Poecilcercus pictus, Acridida exatata; mantis, Hirodula condurctota; cricket Grylus; the termite, Anacanthotermes macrocephalus; all these form a rich source of food for the mammals. The Gazelle and the Black Buck are also frequent visitors. The Pangolin which is not so common lives in burrows near termite mounds. (d) ARTIFICIAL MINES In the Bikaner region there are several deep mines excavated for "FuUer's earth". In these artificial mines are found Rhinolophus lepidus and RMnopoma hardwickei. II. ROCKY ASSOCIATION II (a) General: In the Indian desert, there are rocky regions composed of (1) Volcanic-Malani Rhyolites and (2) Sedimentary-Vindhyan sandstone. They are found near about Jodhpur, Barmer, Jaisalmerand Phalodi.

156 ISHWA~ PRAKASH Mammal Msociatiofl,: Hare, Jackal and Wolf are mainl found in this habitat. Near about Jodhpur, panther and hyaena have also beea obseeved. Mongl!>oseswere cwlleced from rocky localities at Barmer and Jodhpur. Porcupine als0 djgs its long burrow in the sand which accumulate on rocks. The trees are inhabited by' langu.r and squirrel. (b) I;akes and surrounding gardens: There are several parennial lares in the desert encompassed by large gardens. Most of the tree~ are cultivated species. At Balsamand near Jodhpur, Pteropu.~ giganteus giganteus was found on Ficus trees. At Sardarsamand, Jodhpur, wild boars were observed and also nilgai. Adams (1899) repor-tffd otters from the la~es of Jodhpur and Jaisalmer but the author did not observe any. (c1 Cal1p,sand Tunnels: At several places, the rocks have deep and dark caves :which are inhabited by bats. The Bhimbharak.Jodhpur and Barmer caves hav~rat tailed and sheath-tailed bats. False Vampire bats were collected in the tunnel at Mandore,Jodhpur. III. RUDERAL ASSOCIATION The term has been used by Blatter and Hallberg (1918-21) fnr associations of plants near human habitations. In this text, this term is applied to animal associations near human habitations, such as rat, mice, squirrel, Indian gerbil. shrew, bat, ;l3.:hesusmonkey and langur. The black buckis particularly associated with certain areas of Rajasthan where they are protected by a tribe, "Vishnoi". The wild boar, Nilgai, Sambhar, Gazelle are also associated with similar protected areas. Ruins-Near many towns in the desert, huge unattended ruins of buildings are met with. These ruins form ideal bat localities. Rhinop'Jma kinneari ~s found in large numbers in the dark portions of the ruins. Associated with them are R. hardwickei, T. p. perforatus and P. m. glaucillus. Mongooses have also been observed in the ruins. ZOOGE0GRAPHY Blanford (1901) assigned the Punjab tract, including the deserts of Baluchistan, Sind, the Punjab tract, and western Rajputana to the Palaearctic region. The rest of India was placed in the Oriental region. This classification was based on distribution of the animals. Pocock (1939) also mentions that certain mammals ofwester-u!india are very nearly allied to similar types occurring in sou!ta-w6stern Asia and Af~i0a (lion, hunting leopard, caracal, desert cat, hyaena, rate!, chinkara, Indian wolf and desert f >x).

MAMMALS OF RAJASTHAN DESERT 157 (,).. Based-on the 'geographica:1 distribution the/mammals of this region are. classified into the following categories: I. Mamma~ of the Rajasthan desert iu.ittibuteclin the Palaearctic Region. Hemiechinus auritus, Rhinopoma hardwir.kei,taphozous perforatus,ganislupus, - '- Ganis aureus, Vulpes, Herpestes auropunctatus, Herpestes edwardsi, Hyaena hyaena, Felis 'Ubyca, Panthera pardus, Gazella gazella, Hystrix indica, Rattus rattus, Mus musculul/, "Tate'raindica, Meriones kurrianae. II. Mammals of the Rajasthan desert distributed East of Aravalis. Paraechinus micropus, 8uncus murinus; Pteropus giganteus, Rhinopoma kinneari, Rhinopoma hardwirkei, Taphozouz kachhensis, M egaderma lyra, Rhinolophus. lepidus, Pipistrellus mimus, Scotophilus heathi, Macaca mulatta, Presbytis entellus, Manis crassicaudata,. Oanislupus" Vulpes bengalensis, Hyaena hyaena, Felis chaus, Panthera pardus,. 8us scrofa, Oervus unico1or, Boselaphus tragocamelus, Antelope cervicapra, Gazella gazella, Lepus nigricollis, Funambulus pennanti, Hystrig; indioa; Mu.s mu.9cul1t.<j, Rattus rattu.9, Tateta indica,'meriones hurrianae (in arid tracts of Delhi only). III. Mammals 'restricted to Sindo-Raj18thani Desert Hemiechinus auritus collaris,'paraechinus' micropus.micropu8,tbuncus~11iurinus sinden,sis, Pipistrellu3' mimu"i glaucillus,feu.s libyca otn-zta, Felis chaus prateri, Antelope cervicapra rajputp,nae, Lepus nigricollill dayanu/8. IV. 'Mammals found in nearby Arid l Regionstindnot collecteilifrom the ' Rajastlian des'ert. Paraechinus hypomela8 (Sind) Suncus stolizkanub (Sino, Kathiaw.it,"punJab), Viverricula malaccensis (Cutch), Golunda ellioti, (Cutch), Gerbillus nanu's' (Cutch; Sind), Cremmomys cutchicus (Cutch), Oapra1akoneri (Sind). It has been further observed ( Prakash, 1963) that 564 percent mammal species of this desert are Palaearctic, 4r per cent Oriental and Indo-Malayan' an'd'. only 2.6 per cent species are endemic. SUMMARY The Rajasthan Desert was surveyed au the year round during 1953to 1956 for couection of mammals which, comprised of 38 species and subspecies (Ins(:;ctivora 3,Chiropteva 9, Primates 2, Pholidota 1, Carnivora 10, Artiodactyla 5, Lagomorpha 2, and Rodentia 6). Under, each species or sub-species. the author's' reference, notes on the biology, including habits and preferred habitat, stomach contents, breeding

158 ISHW AR PRAKAsH habits and the parasites found on the body have been given. Body and cranial measurements are appended. Three main mammalian associations have been recoignized in the desert region: Sandy, Rocky and Ruderal. The first comprises of 4 sub-associationssand dunes, sand hills, sand plains and artificial mines. The second consists of lakes, gardens,.caves and tunnels. The ruderal is the association which is influenced by man. Under each sub-association, mammals and other animals, particularly snakes, lizards and insects are listed and their relations are discussed. The breeding season of the mammalian listed. species is also discussed. Six food centres have been established and mammals by each centre are Simplified food chains are also given. The zoogeographical study shows that the mammalian fauna of the Indian degert is comparatively of recent origin and it is found that 56.4 per cent mammals of the Rajasthan desel'tare Palacearctic and the rest are Indo-Malayan. ACKNOWLEDGMENTS The project on the study of the vertebrates of the Rajasthan desert was financed by UNESCO and was carried out under the guidance of Dr. Daya Krishna, Professor of Zoology, Jaswant College, Jodhpur. I am extremely grateful to UNESCO 'authorities and to Dr. Krishna who helped me painstakingly during the entire course of research by his mature advice and guidance. I am also indebte~ to Dr. M. L. Roonwal, Director, Zoological Survey of India and Dr. P.C. Raheja, Director, Central Arid Zone Research Institute, Jodhpur for unstinting help. The late Dr. F. S. Bodenheimer, Jerusalem was kind enough to go through the manuscript. REFERENCES ADMAS,A. 1899, The Western Rajputana States, Taylor & Francis, London. ALLEE, W. C., EMERSEN,A. E., PARK. 0., PARK, T., & SCHMIDT,K. P. 1949 Prinoiples of animal ecology, Saunders & Co. Philad. & London. ATKINSON, J. C. 1844, Notes on a hedgehog. Zoologist, 2, 791-792 (quoted by Cott, 1951). BANERJI,ARUNA1955, The family life of a five-striped squirrel Funambulus pennanti Wr.), J. Bombay nat. Hist. Soc., 53, 261-264. BANERJI,ARUNA1957, Further observations on the family life of the five striped squirrel, Funambulus pennanti Wr., Ibid, 54, 336-343. BLANFORDW. 'T. 1888-91, The fauna of British India, Mammalia., Taylor & Francis, London.

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MAMMALSOF RAJASTHANDESERT 161 PRAKASHISHWAR. 1960, Breeding of mammals in Rajasthan desert, India. J. Mamm., 41, 386-389. ---~ 1960 a, Taxonomic and Biological observations on the bats of the Rajasthan desert. Rec. lnd.mu,s., 59,,149-170. 1961, Die s:tugetiere der Rajasthan wiiste in India. Natur und Volle. 9],443-448. 1962,., Ecology of the gerbils of the Rajasthan desert, India, Mammalia, 26, 31I.:131. 1962 a, Rolling of the desert hedgehog, Hemiechinas a1~rituscollaris Gray, J. Anat Soc. India, 9, ]9-22. ---- 1962 b, Group organisation, sexual behaviour and breeding season of certian Indian monkeys, Jap. J. Ecol" 12,80-86. 1962 c, Times of emergence of the pipistrelle, Mammalia, 26, 133-135. ]962 d, Eco-toxicology and Control of desert gerbils. Pt. 1. Feeding behaviour energy requirement and selection or bait, J. Bombay nat. Rist. Soc., 59, 800-806. 1962 e, Eco-toxicology and control of desert gerbils. Pt. 2 Breeding season, litter size and post natal development. Ibid, (In Presl1). 1962 f" Eco-toxicology and Control of desert gerbils Pt. 3, Toxicity, relative acceptability and lethal. dose of certain poisons, Indian Fares er (In Press). 1962 g, Eco-toxicology and control of desert gerbils Pt. 4, Burrow temperatures. Ibid, (In Prcss). 1963, 'Zoo-geography and evolution of the mammalian fauna of Rajasthan desert, India, Mammalia 27, 342-351. & SHARMAS. C. 1955, Nematode and hedgehog mortality, J. Bombay nat. Hist. Sac, 5:J, 586. ROBINSON,W. ]928, Water conservation in Insects, J. Econ. Ent. 21, 897-902. (not seen in original). ROONWAL, M.L. 1950, Contributions to 'he fauna pi Manipur State, Assam. Pt. III Mammals, with special reference to the family muridae (order Rodentia). Rec. Ind. Mus. 47, 1-64. SCUUITDT-NIELSEN, B. & SCHMIDT-NIELSEN,K. 1950, Evaporative water loss in desert rodents in their natural habitat, Ecolo;7Y, 31, 75-85. & ---- 1952, Water metabolism in desert mammals. Physiol. Rev., 32~ 135-166. STERNDALE, R. A. 1884, Natural History of Mammals of India and Ceylon. Thacker Spink & Co., Calcutta. WALL F. 1906, The snake and its natural foes, J. Bombaynat. Hi,9t. Sac, 17,386.