JOURNAL )F THE ^lOMRAY NATURAL HISTORY SOCIETY , .^RIL 2000 Vol. 97 (1) / .... V M.R. ALMEIDA BOARD OF EDITORS Editor J.C. DANIEL AJITH KUMAR M.K. CHANDRASHEKARAN T.C. NARENDRAN B.F. CHHAPGAR A.R. RAHMANI R. GADAGKAR J.S. SINGH INDRANEIL DAS R. WHITAKER A.J.T. JOHNSINGH ^ - Assistant Editor GAYATRI WATTAL UGRA > INSTRUCTIONS TO CONTRIBUTORS 1. Papers which have been published or have been offered for publication elsewhere should not be submitted. 2. Papers should be submitted in duplicate, typed double space. Preferably an additional copy should be submitted on a floppy diskette (3.5") using Word Star. 3. Trinomials referring to subspecies should only be used where identification has been authentically established by comparison of specimens actually collected. 4. Photographs for reproduction must be clear, with good contrast. Prints should be at least 9 x 12 cm and on glossy glazed paper. Text-figures, line drawings and maps should be in Indian ink, preferably on tracing paper. Maps and figures will not be acceptable if labelled free hand. 5. References to literature should be placed at the end of the paper, alphabetically arranged under author’s name, with the abridged titles of journals or periodicals in italics and titles of books or papers in roman type, thus: Aluri, Raju J.S. & C. Subha Reddi (1995): Ecology of the pollination in two cat-mint species. J. Bombay nat. Hist. Soc. 92(1): 63-66. Prater, S.H. (1948): The Book of Indian Animals. Bombay Natural History Society, Mumbai, pp. 35-48. 6. Each paper should be accompanied by an abstract, normally not exceeding 200 words, and 6-8 keywords. Keywords should include the scientific names of important species discussed. 7. 25 reprints will be supplied free of cost to authors of main articles. In the case of new descriptions, reviews and miscellaneous notes, authors will be sent a free copy of the Journal. 8. The editors reserve the right, other things being equal, to publish a member’s contribution earlier than a non-member’s. Hornbill House, Shaheed Bhagat Singh Road, Mumbai-400 023. Editors, Journal of the Bombay Natural History Society POPULATION DENSITIES OF THE BLACKNAPED PlARE LEPUS NIGRICOLLJS NIGR1COLLIS AT ROLLAPADU WILDLIFE SANCTUARY, KURNOOL DISTRICT, ANDHRA PRADESH ( With six text-figures ) By Ranjit Manakadan and Asad Rafi Rahmani 3 BREEDING BIOLOGY OF THE MALABAR GREY HORNBILL ( OCYCEROS G RISE US ) IN SOUTHERN WESTERN GHATS, INDIA (With one text-figure) By Divya Mudappa 15 SOCIOECONOMIC TRANSITION AND WILDLIFE CONSERVATION IN THE INDIAN TRANS-HIMALAYA By Charudutt Mishra 25 AN ECOLOGICAL STUDY OF CROCODILES IN RUHUNA NATIONAL PARK, SRI LANKA ( With three text-figures) By Charles Santiapillai, Mangala de Silva, Sarath Dissanayake, B.V.R. Jayaratne and S. Wijeyamohan 33 SEXUAL HARASSMENT AMONG FEMALE LION-TAILED MACAQUES (MAC AC A SILENVS) IN THE WILD ( With three text-figures) By Ajith Kumar 42 SEASONAL CHANGES OF TROPICAL FOREST BIRDS IN THE SOUTHERN WESTERN GHATS ( With seven text-figures) By E.A. Jayson and D.N. Mathew 52 PLODIA INTERPUNCTELLA (HUBNER) (PHYCITIDAE : LEPIDOPTERA) AS A POTENTIAL PEST OF DRY FRUITS By S.P. Rad, H.R. Pajni and Neelima Talwar 62 FRESHWATER CLADOCERA (CRUSTACEA : BRANCHIOPODA) OF THE ANDAMAN AND NICOBAR ISLANDS ( With one text-figure) By K. Venkataraman 67 LONGICORN BEETLES (CERAMBYCINAE, PRIONINAE : CERAMBYCIDAE) OF BUXA TIGER RESERVE, JALPAIGURI, WEST BENGAL ( With twelve text-figures) By Dinendra Raychaudhuri and Sumana Saha 74 FISHES OF THE CYPRINID GENUS SEMIPLOTUS BLEEKER 1859, WITH DESCRIPTION OF A NEW SPECIES FROM MANIPUR, INDIA ( With one text-figure and one plate) By Waikhom Vishwanath and Laishram Kosygin 92 FOOD AND FEEDING HABITS OF INDIAN BARBETS, MEGALAIMA SPP. ( With three text-figures) By Hafiz S.A. Yahya 103 NEW DESCRIPTIONS SPINY EELS OF THE GENUS MACROGNATHUS LACEPEDE FROM MANIPUR, WITH DESCRIPTION OF A NEW SPECIES ( With four text-figures ) By L. Arunkumar and H. Tombi Singh 117 THREE NEW GENERA OF WHITEFLIES M OH A NA S UN DA RA MI ELLA , SHANTHINIAE AND V A SA NTH A RAJ I ELLA (ALEYRODIDAE : HOMOPTERA) FROM INDIA ( With three text-figures) By P. Manidurai Manoharan David 123 LYSIONOTUS PALINENSIS — A NEW SPECIES OF GESNERIACEAE FROM ARUNACHAL PRADESH, INDIA ( With one text-figure) By G.D. Pal 131 REVIEWS 1 . BIRDS OF NEPAL: FIELD ECOLOGY, NATURAL HISTORY AND CONSERVATION Reviewed by Asad R. Rahmani 133 2. BIOGEOGRAPHY OF THE REPTILES OF SOUTH ASIA Reviewed by Meghana Gavand 133 3. MOSSES OF KH AND ALA AND MAHABALESHWAR IN THE WESTERN GHATS, INDIA Reviewed by P.K.K. Nair 134 MISCELLANEOUS NOTES MAMMALS 1 . Instances of fruit bat mobbing the barn owl By Sunil Zaveri 136 2. Possible occurrence of the lesser woolly horseshoe bat ( Rhinolophus beddomei) in Chinnar Wildlife Sanctuary By Kumaran Sathasivam 136 3. Dead snow leopard Uncia uncici at Yabuk, Dongkung (5500 m) in north Sikkim By Usha Ganguli-Lachungpa 137 4. On the longevity of the tiger {Panther a tigris) in captivity By L.N. Acharjyo, B.C. Prusty and S.K. Patnaik 138 5. Sighting of barking deer ( Muntiacus muntjac) in Kalakad-Mundanthurai Tiger Reserve, Tamil Nadu By Jayanti Ray, Justus Joshua and J. Ronald 139 6. Type specimens of mammals in the collections of the Bombay Natural History Society By Meghana Gavand and Naresh Chaturvedi 1 40 AVES 7. Night herons and little cormorants in Thrissur, Kerala By Leela Madhavan 142 8. Grey heron wresting fish from herringgull By Lavkumar Khacher 142 9. Additional site records of black stork Ciconia nigra (Linn.) in Andhra Pradesh By V. Vasudeva Rao. V. Nagulu and C. Srinivasulu 143 10. Stealing of redwattled lapwing Vanellus indicus (Boddaert) and yellow-wattled lapwing Vanellus malabaricus (Boddaert) eggs by cowherds By K. V. Srini vas and S. Subramanya 143 11. A note on the feeding of lesser coucal {Centropus toulou) BySamiranJha 144 12. Occurrence of the yellowbrowed bulbul Hypsipetes indicus (Jerdon) in the Nalamalla Hills, Andhra Pradesh By Srinivasulu and V. Vasudeva Rao 144 13. Termite attack on nest material leading to desertion of eggs by birds By K. V. Srini vas and S. Subramanya 145 14. Range extension of the purplerumped sunbird Nectarinia zeylonica By Lavkumar Khacher 146 15. Water acquisition strategy adopted by goldfinch ( Carduelis carduelis) By R. Suresh Kumar 147 REPTILES 16. Occurrence of draco or flying lizard Draco dussumieri in Chittoor district, Andhra Pradesh By S. Balachandran and Aasheesh Pittie 1 47 1 7. Occurrence of yellow-bellied Pelamis platurus (Linn.) Reptilia : Hydrophidae, in coastal waters off Digha, West Bengal By S. Mitra, J. Sarkar and T.K. Chatterjee .. 148 AMPHIBIA 1 8. A record audio feat by an anuran By Sanjeev B. Nalavade 149 FISHES 19. Range extension of Pangio goaensis (Cyprini formes : Cobitidae) to the Chaliyar drainage of Kerala By K. Rema Devi, K.G. Emiliyamma and R.S. Lalmohan 150 20. Fishes of Nambiyar river, Kalakad- Mundanthurai Tiger Reserve, Tamil Nadu By M. Arunachalam, A. Sankaranarayanan, J.A. Johnson, A. Manimekalan, R. Soranam, P.N. Shanthi and C. Vijaykumar 1 53 21 . A profile of the food and feeding of hillstream teleosts ofGarhwal Himalayas By N. Singh and R. Subbaraj 155 INSECTS 22. A supplementary list of the host-plants of Indian Lepidoptera By Peter Snietacek and Rajani Smetacek .... 157 23. On the predation of the Giant Redeye Gangara thyrsis (Fabricius) (Family : Hesperiidae; Order : Lepidoptera) By S. Karthikeyan 160 24. Mating behaviour of the Common Mormon Papilio polytes (Family : Papilionidae) By Arnab Bose 160 OTHER INVERTEBRATES 25. Mycophagous arthropods from the Andaman Islands By Prashanth Mohanraj and K. Veenakumari 1 61 26. On Daphniopsis tibetana Sars, 1903, (Cladocera) collected from a high altitude Himalayan lake, India By K. Venkataraman 162 BOTANY 27. fndigofera mysorensis Rottler ex DC. (Leguminosae : Papilionoideae) — An endemic species of Peninsular India from West Bengal By S. Mitra, S. Bandyopadhyay and A. K. Sarkar 165 28 . Range extension of Nepenthes khasiana i n the Jaintia hills, Meghalaya By Anwaruddin Choudhury 166 29. Scleria laxa R. Br. (Cyperaceae) - A new record for India from Nicobar Islands By P.V. Sreekumar 167 30. Rhaphidophora calophyllum Schott (Araceae) — An addition to the flora of the Andaman & Nicobar Islands By K. Sasikala and E. Vajravelu 169 Cover photograph: Wild Tusker Editorial The problems facing the Asian elephant in India are a reflection of the state of environ- mental conservation in India. As a species able to live in a wide spectrum of vegetational types, the elephant acts as an indicator species of the condition of its biotic environment. A sub-optimal habitat is unable to meet the demands made on it by a herd of elephants, whose presence will result in further deterioration. Elephants in such habitats are compelled to seek sustenance elsewhere, and come into conflict with man. At the present rate of habitat loss, and degradation of existing habitats, it is doubtful if present populations can survive. One has to consider seriously the possibility that the Asian elephant will be known mainly as a domesticated animal in the 21st century. In India, an enormous area of prime elephant habitat has been lost since 1 860, to the plantations of coffee, tea, rubber and teak which were carved out of existing forests. After 1950, hydroelectric projects ravaged elephant habitat through the submerging of forests and unscrupulous exploitation of the remnant forests. In central India, the forests holding elephants cover the single largest deposit of iron-ore in Asia, and mining has been a continuing process since 1909. The states of northeast India, which used to be the stronghold of the elephant in India, are the areas where the main human-elephant conflict has developed. Exploding human populations have destroyed crucial elephant habitat for cultivation and plantations, extinguishing traditional migratory routes; and slash-and-burn cultivation has devastated habitats, making unlikely the survival of the elephant in some of the states. There is also the question of ivory poaching. Though not on as massive a scale as of the African species, the selective removal of tuskers has played havoc in the sex ratio of many populations. The elephant is an apex species, able by its size and its interaction with its habitat, particularly in its quest for food, to influence the direction of development of its biotic environment. It has been one of the causes for the process of change in its ecosystem. Such a function is no longer acceptable in an environment managed by man, where the process of change has been speeded up. The range of the elephant has, through the ages, shrunk considerably. This process was accelerated, however, as the industrial revolution in the latter half of the last century brought a mechanized commercial culture into the countries of its occurrence. The tools used by man in a region decide its future, and the tools of an alien culture, now in use for gathering natural resources for commerce and to meet the needs of an ever-increasing human population, have destroyed a natural slow-moving ecosystem. The elephant has become in the process too large an animal to find sustenance and living room in the shrinking world of nature. The conservation of the Asian elephant in Asia cannot be the concern of only the forest departments and environmentalists. Conserving the elephant involves the conservation of prime wildlife habitats. This needs a multidisciplinary effort, where the local people, the administrators and land-use planners have to be involved at all levels. Conserving the elephant, therefore, means conserving the human environment, and it has to be a part of the development plans of each state of Asia as a whole. The Asian elephant is a part of the culture of man in tropical Asia. It is an integral part of the religions of the region and one hopes, will not be sacrificed in the search for a better life for the people of the region. J.C. DANIEL ACKNOWLEDGEMENT We are grateful to the Ministry of Science and Technology, Govt, of India, FOR ENHANCED FINANCIAL SUPPORT FOR THE PUBLICATION OF THE JOURNAL. JOURNAL OF THE BOMBAY NATURAL HISTORY SOCIETY April 2000 Vol. 97 No. 1 POPULATION AND ECOLOGY OF THE INDIAN FOX VULPES BENGALENS1S AT ROLLAPADU WILDLIFE SANCTUARY, ANDHRA PRADESH. INDIA1 Ranjit Manakadan and Asad Rafi Rahmani2 ( With six text-figures) Key words: Indian fox, Vulpes bengcilensis , Ardeotis nigriceps , population, diet, breeding season, Rollapadu Wildlife Sanctuary, Andhra Pradesh The population of the Indian fox Vulpes bengcilensis , its spatial and temporal abundances, den distribution, characteristics and use, predation on eggs and chicks of the great Indian bustard Ardeotis nigriceps , and general ecology were studied from February 1993 to April 1995 at the Rollapadu Wildlife Sanctuary (RWS), Andhra Pradesh state, India. The population and spatial abundance of the fox was estimated by enumeration and monitoring of dens, animal sightings at den sites and from censuses. The population of the fox at RWS was estimated to be around 40-50 adult animals in 1993 and 1994, which declined to about 10 animals in 1995 due to an epidemic. Densities of the fox were significantly higher in the protected grasslands {0.65/40 ha ± 0.99 (S.D)} than unprotected grasslands (0. 1 5/40 ha ± 0.49). A total of 1 35 dens (active and non-active), comprising of 33 'den groups', were located in the study area. There was a concentration of dens in and around protected grasslands. Den use by the Indian fox at RWS was confined to the pup rearing season (February to June/July). We did not record any evidence of fox predation on bustard eggs and chicks. increase after the establishment of the Sanctuary in the early 1980s to protect the great Indian bustard and its habitat (Manakadan and Rahmani 1989, 1993, 1997). The Indian fox is known to be a predator of eggs and probably chicks of the bustard (Rahmani and Manakadan 1987). This was suspected to be one of the reasons for the decreasing numbers of the great Indian bustard at RWS over the years, in spite of good protection to the bird and its habitat. We undertook this study to estimate the population of the Indian fox at RWS; compare its abundance in protected and unprotected sites in the Sanctuary; assess reasons for the differences in abundance between sites (which could explain the increase in Introduction The Indian fox Vulpes bengcilensis is a widespread species in India, ranging from the foothills of the Himalayas to Kanyakumari (Prater 1980). In spite of its wide distribution and proximity to human habitation in many areas, it has not been studied adequately (Johnsingh 1978). The population of the Indian fox in Rollapadu Wildlife Sanctuary (RWS), Andhra Pradesh had undergone a remarkable 'Accepted April, 1999 2Bombay Natural History Society, Hombill House, Shaheed Bhagat Singh Road, Mumbai 400023, Maharashtra, India. JOURNAL . BOMBAY NATURAL HISTORY SOCIETY, 97(1). APR. 2000 3 POPULATION AND ECOLOGY OF THE INDIAN FOX 4 JOURNAL, BOMBAY NATURAL HISTORY SOCIETY. 97(1). APR. 2000 Fig. I : Location of fox dens in the study area POPULA TIONAND ECOLOGY OF THE INDIAN FOX populations over the years after protection); investigate the role of the fox as a predator of bustard eggs and chicks; and collect other ecological information on the species. Study Area Rollapadu is 18 km southeast of Nandikotkur (15°58' N lat. & 78° 18' E long.), Kurnool dist., Andhra Pradesh. It lies in the plains between the Nallamalai and Yerramalai hills, at about 200 m above msl. The terrain is gently undulating with predominantly poor red soil. The region is semi-arid with an average annual rainfall of 668 mm, received from both the southwest (June to August) and northeast (September to December) monsoon. Summer (March to May) peaks at 42°C and winter (November to February) is mild at 17° C. Rollapadu Wildlife Sanctuary (area: 6.14 sq. km) was established in 1982, after the "rediscovery’ of the great Indian bustard A rdeotis nigriceps, and was declared a sanctuary in 1988. The sanctuary proper consists primarily of three grassland plots or enclosures: Enclosure-I (320 ha), about 500 m north of Rollapadu, and Enclosure-II (40 ha) and III (120 ha), both about 1.5 km to the northeast of Rollapadu (Fig. 1). These enclosures are demarcated by trench-cum- mound (TCM) walls to exclude livestock and people. However, Enclosure-Ill was opened to grazing after protests by the locals about the lack of grazing land for their livestock. The extent of protection to Enclosure-II varied from year to year during the study. The three enclosures are separated from each other by grazing lands and crop fields. Both the grazing lands and the enclosures are predominantly grasslands, with scrub dominated areas along streams. The other major fauna of the Sanctuary include the blackbuck Antilope cervicapra , wolf Canis lupus, jackal Canis aureus, jungle cat Felis chaus, common mongoose Herpestes edwardsi, blacknaped hare Lepus nigricollis nigricollis, common Indian monitor Varanus bengalensis and lesser florican Sypheotides indica. The grassland is a major roosting ground for harriers (largely Circus pygargus and C. macrourus ) wintering in the Indian subcontinent. For more details, see Rahmani and Manakadan (1986) and Manakadan and Rahmani (1989, 1993 & 1997). Methodology Studies were conducted from February 1993 to April 1995, during daylight hours on unmarked animals. Prior to the studies, we had a fairly good idea of the population and distribution of the fox in RWS from July 1992, due to our field visits during other multi- disciplinary studies of the project. Population: A pilot survey was conducted during the breeding season in 1993 to assess den distribution in the study area. The survey was concentrated in the three enclosures and grazing lands adjoining them, to get an insight into the breeding season, den characteristics and distribution of the fox in the Sanctuary. Den searches were more intensive during the breeding season of 1994 and 1995. Searches in 1994 began in February, when the dens located in 1993 were found to have been dug up afresh, indicating the start of the breeding (pup rearing) season. The area searched (Fig. 1) was divided into smaller blocks and combed intensively for dens by two or three people. The locations of these dens were plotted on a map (Fig. 1) and details, such as active or non- active, number of holes per den, distances between dens, and site characteristics were recorded. After the survey, all the dens were visited once a week to collect data on den use. Sightings of animals (adults and young) at den sites were recorded. We also looked for indirect signs of animal presence, such as freshly unearthed soil, additional holes dug up, pugmarks, presence of scats and food remains at den sites. Visits were made till June (when the animals abandoned the dens with the onset of the monsoons) in 1994, and till May in 1995 (after the breeding season). JOURNAL . BOMBAY NATURAL HISTORY SOCIETY, 97(1), APR. 2000 5 POPULA r I ON AND ECOLOG Y OF THE INDIAN FOX Population estimate: Based on the number of dens located, den use data, and number of adult foxes seen at den sites, a rough estimate of the adult population at RWS was determined. Where animals were never seen at active dens throughout the study period, and especially if the den formed part of a complex of dens (termed den group) as in the majority of cases, we presumed that the den / den group belonged to a pair, as two animals for each den group was the norm in most of the den groups. Densities in grazing land and enclosures: Four sites of 40 ha each were selected in each of the two habitat types. Except for one site in the grazing land, which was predominantly scrub, all the other sites were grasslands. The sites were thoroughly covered on foot fortnightly — on different days — in the evenings from July 1994 to April 1995. Though the sites were searched on different days, repeated flushing of animals from the same areas suggested that the animals were territorial and that there was no significant movement between sites. Each site was searched in an hour’s time, by walking at a steady pace, in an irregular and generally zigzag manner. Some light noise (humming, dragging of feet, tapping with a stick) was made to flush the resting, sleeping or hidden foxes inside dens or among vegetation. Loud noise was avoided as it would alert the animal a good distance away, allowing it to slip away without being detected. On flushing a fox, the direction in which it ran and the place it stopped was observed to avoid duplication of counts. The fox sightings were expressed as number of foxes/40 ha. Food Availability: Data on the abundance of the known food items of the fox, such as fruits (number of fruiting trees) and grasshoppers in the two habitat types was obtained from other studies carried out during the project. Grasshoppers were sampled by the sweep net sampling method (100 sweeps per site), and was done fortnightly at all four sites in both the habitat types. The density of fruiting trees was enumerated by laying 40 quadrats (size 50 x 50 m) each in both the habitat types, and noting the species of trees or shrubs, their numbers and heights. An index of rodent abundance was obtained by enumeration of burrows along one kilometre transects (with a width of two metres), laid at random in both the habitat types. The transects were done during summer (breeding season of the fox). Fifteen transects each were laid in the enclosure and grazing land during 1994 and 1995. For more details, see Manakadan and Rahmani (1997). Diet: Scats of fox were collected whenever seen, but mostly during the breeding season, when they were available around den sites. The scats were mixed with warm water, strained and dried. After drying, the remains of animal and plant parts were recorded visually. The percentage composition was not estimated systematically, as the main purpose of the exercise was to look for remains of bustard eggs or chicks. Results Dens: The breeding (pup rearing) season of the fox in RWS was determined to be between February to May from 3 years observations. The breeding season was heralded by the re- excavation of old dens or digging of new ones in February. Scats of pups were found around den sites during April and May. Pups were seen around the den sites till the onset of the monsoon, after which the dens were abandoned. Thus, den use by the Indian fox at RWS was largely restricted to the pup rearing period. Fox dens were recorded in grassland or light scrub habitats — none in dense scrub areas. Dens were dug in the flat ground or in trench cum mound walls (TCM) of the enclosures. Two dens were recorded along the slopes of a stream. The number of holes or openings per den varied from one to as high as 43, but two to seven holes were most common (Figs. 2, 3). All the holes of a den were not used, two to seven active holes per den were most frequent. The frequency of 6 JOURNAL, BOMBAY NATURAL HISTORY SOCIETY, 97(1), APR. 2000 POPULA TION AND ECO LOG Y OF THE INDIA N FOX Frequency of dens Number of holes Fig. 2: Number of holes per den Frequency of dens Fig. 3: Number of active holes per den (1994) JOURNAL BOMBAY NATURAL HISTORY SOCIETY, 97(1), APR. 2000 7 POPULA TION AND ECOLOGY OF THE INDIAN FOX Table 1 DETAILS AND STATUS OF DEN GROUP (BASED ON 1994 DATA) Den Group No. No. of dens per group No. of foxes recorded/ estimated 1994 Status in 1993 1995 1 1 0 NA - NA 2 4 2* A A A 3 6 2** LL-A A NA 4 5 0 NA A NA 5 3 0 0 A NA 6 5 2* A A NA 7 8 9 NA A NA 8 5 2* A A NA 9 5 0 0 NA NA 10 5 2* B A NA 11 1 0 0 NA NA 12 4 2** B A NA 13 7 9 A B NA 14 2 1* A A NA 15 9 2* B B NA 16 2 0 0 - NA 17 1 2(?) LL-A - NA 18 2 1* A B NA 19 7 2* B A A 20 4 2** B - NA 21 5 2* B A NA 22 3 0 0 - NA 23 3 0 0 - NA 24 8 2**(j *\ A A A 25 3 9 LL-NA - NA 26 3 2(?) A - NA 27 3 7** B - NA 28 1 1* A A NA 29 5 2**(1 *) A - A 30 3 9 A - A 31 4 2**(1*) A - B 32 4 2(?) A A NA 33 4 9 LL-NA B NA * - from sightings ** - from signs (scats or intensive burrowing) 2**( 1 *) - 1 seen, but probably used by a pair. ? - uncertain A - Active burrows regularly used, dug or redug NA - Not active, dug early in the season, but later largely or totally unused. O - Dens of previous years: not dug at all during the year of survey . B - Breeding (pups or scats of pups seen) LL-A - Located late (afterbreeding season); -probably active LL-NA - Located late (after breeding season); - probably not active - Not located - all or some of the dens of the den group were not located. active openings in the eight breeding dens of 1994 were six for three dens, five for two dens, three for two dens and nine for one den. Many of the dens in the grazing land had rodent burrows around them, indicating that these sites had been appropriated from rodents. In some cases, the rodents continued to live in some of the burrows not enlarged by the fox. Re- use of dens by rodents after the fox had abandoned the dens during the monsoon was recorded in some cases. On two occasions, large monitor lizards Varanus bengalensis were recorded entering active fox dens. Once, a large monitor lizard, flushed by us near a den site, ran into a fox den, from which a family of gerbils rushed out and ran into their burrows a few metres from the fox den. Seven of the fox dens were appropriated by jackals or wolves (Fig. 1). During the preliminary non-intensive searches for dens in 1993, a total of 52 dens were located (33 active and 19 non-active). Breeding activity was detected in 4 dens: 1 in Enclosure-1, 2 in Enclosure-II and 1 in the grazing land. During intensive searches in 1994, a total of 135 dens were located, of which 52 were active. Of the 135 dens, 51 were in Enclosure-1, 15 in Enclosure-II, 9 in Enclosure-Ill and 60 in the* grazing land. As much as 31% of the dens in the grazing land were close to Enclosure-I and II. Breeding activity was recorded in eight dens: three dens each in Enclosure-I and Enclosure- II; one each in Enclosure-Ill and grazing land. During the breeding season in 1995, no additional dens were located. Of the dens located in 1994, only eight dens were reused (active). Breeding was confirmed at only one den in the grazing land. From the data on sightings of animals and den use, it was evident that many of the foxes used more than one den. From this data, the 135 dens located during the intensive survey in 1994 were grouped into 33 den groups, of which 22 were active (Table 1 & Fig. 1). Dens of a group generally tended to be clumped in an area, the distances between dens varying from as close as 8 JOURNAL, BOMBAY NATURAL HISTORY SOCIETY, 97(1), APR. 2000 POPULA TJONAND ECOLOGY OF THE INDIAN FOX No. of occurrences Den group size classes Fig. 4: Number of dens per den group (1994) No. of occurrences Den group size classes Fig. 5: Number of active dens per den group ( 1 994) JOURNAL BOMBAY NATURAL HISTORY SOCIETY, 97(1), APR. 2000 9 POPULA TION AND ECO LOG Y OF THE INDIA N FOX 12 m to 100 m. In some cases, the distance of a den from the main cluster was more than 200 m (e.g., den group no. 10 - Fig. 1), but these were clumped to the group, based on sightings and movements of adult and young foxes between dens. In some cases, we grouped two clusters of adjoining dens into one (e.g., den group no. 7), as one of these clusters was hardly used and was probably the denning site of the pair in the area during a previous year. The distances between den groups varied, and were less in the enclosures {Enclosure-I: 463.6 m ±283.8 (S.D.), Enclosure- II: 275.0 m ±302.9, Enclosure-Ill: (400.0 ±424.3) than the grazing land (633.3 m ±314.3)}. The number of dens per den group varied from one to nine dens, with three to five dens being most frequent (Fig. 4). However, not all dens in a den group were active during a year, one to three active dens was most common (Fig. 5). We presume that each (active) den group belonged to either a pair of foxes or rarely individuals, but cannot be certain as the animals were not collared, the nocturnal movements were not monitored, and a few dens showed all signs of regular use (especially those in the grazing lands), but no animals were sighted in them. Population: We regularly saw five pairs of foxes around Enclosure-I (den group 3, 6, 8, 10 & 13), four pairs around Enclosure-II (den group 14, 15, 19, 21), three pairs in the grazing lands east of Enclosure-Ill (den group 25,31 and 33), and a single individual at den group 28 during our field trips in 1993 and 1994 — a total of 25 foxes. Judging from the number of dens and groups, den use data, and sightings of the animal around dens during the census, it is estimated that about 40-50 foxes were present in the study area during the 1994 breeding season. About the same numbers should have been present during 1993. In 1995, the population dropped to about 10 animals due to an epidemic. The foxes were usually seen in pairs around the den-groups. Two instances of four adult animals frequenting a common area was 1.2 1 0.8 0.6 0.4 0.2 0 July-1 July-ll Aug- 1 Aug-ll Sep-I Sep-ll Mean numbers/40 ha Fortnights (1994) □ Grazing Land □Enclosure Fig. 6: Abundance (sighting / 40 ha) of the Indian fox in the two habitats 10 JOURNAL. BOMBAY NATURAL HISTORY SOCIETY. 97(1). APR. 2000 POPULA TIONAND ECOLOGY OF THE INDIAN FOX observed. In the first instance, it was during the early monsoon period, when four animals (probably pairs from nearby waterlogged dens) regularly sheltered under a fallen tree as the grass cover was burnt off in a summer fire. In the other instance four adult animals were flushed from a non-breeding den in summer. Otherwise, pairs were the norm and even in clumped den areas, the pairs kept to themselves. Solitary animals were flushed from dens on a few occasions, but the possibility of the mate sleeping elsewhere unnoticed cannot be ruled out. The only den where solitary animals were repeatedly flushed, was at den group 28, which comprised of only one den with a single enhance. Additionally, this was the only single-hole den from which we had actual sightings of the animal. Abundance of the fox was significantly higher (U=140, P<0.05) in the enclosure (mean 0.65/40 ha, S.D. ± 0.99) than in the grazing land (mean 0.15/40 ha S.D. ±0.49). The fox was recorded during all the fortnights in the enclosures from the first week of July 1994, till the first week of September 1994 (Fig. 6). In the grazing land, the fox was recorded only during two fortnights between July to August 1994. In both cases, the foxes were close to Enclosure-II, and ran into it on being approached. All the sightings in the grazing land and enclosures were m ‘grassland habitats’, none in scrubland. After the second week of September 1994, there were only rare sightings of the fox in the Sanctuary. The remains of five foxes and two wolves were found at different places between July to September. The locals too reported seeing dead foxes. After the epidemic, the only sightings were of single animals, one each in Enclosure-II and the grazing land (den group No. 31) during March and April 1995. Food Availability Fruits: Of the two species of fruits recorded to be eaten by the fox, the density of Cassia fistula was higher in the enclosure (1.8 trees/ha) than in the grazing land (0.2 trees/ha). Though the density of Zizyphus mauritiana was about the same in the enclosure and the grazing land, the trees were relatively taller in the enclosure (mean = 1.5 m) than in the grazing land (mean = 0.65 m), and yielded more fruit. Other fruits that could probably be part of the diet of the fox are Morinda tinctovia and Phoenix sylvestris. Trees of these two species were more abundant in the enclosure (0.6 and 5.2 trees/ha respectively) than in the grazing land ( 1 tree/ha for P. sylvestris ; M. tinctoria not recorded). The higher densities of fruiting trees and fruit yield, and restrictions on harvesting of fruits in the enclosures, make the availability of fruits greater in the enclosure than grazing land. Grasshoppers: Insect sampling showed that there was a slightly higher abundance of grasshoppers in the enclosure than in the grazing land. Besides numerical abundance, there was greater insect biomass availability in the enclosure due to the predominance of a larger species of grasshopper ( Acorypha ), compared to a smaller species ( Chrotogonus ) in grazing land. Studies on the great Indian bustard have shown that Acorypha is preferred to Chrotogonus , especially by adult birds (Manakadan and Rahmani 1990). The fox would also find feeding on the larger species more profitable. Rodents: Rodent burrows were recorded only in the grazing land in 1994 and 1995. Of the 1 5 transects each laid in the grazing land for both the years, a total of seventeen burrows (2 active and 15 non active) were recorded in five transects during 1994, and nine burrows (6 active and 3 non-active) were located in 5 transects during 1995. Diet: Analysis of 58 scats showed the presence of rodents, hare, monitor lizard and grasshoppers (predominantly Acorypha sp.) among the animal matter. Among vegetable matter, seeds of groundnut Arcichis hypogea , Zizyphus mauritiana and Cassia fistula were recorded. Remains of eggs or chicks of the great Indian bustard were not recorded. Scats of pups were almost solely made up of rodent fur. 1 1 JOURNAL BOMBAY NATURAL HISTORY SOCIETY, 97(1), APR. 2000 POPULA TIONAND ECOLOGY OF THE INDIAN FOX Threats: The potential predators of the fox at RWS are the wolf, jackal, jungle cat, monitor lizard (on young), and large raptors. Wolves and jackals were seen digging (to eat cubs?) or appropriating fox dens during the breeding season. Large monitor lizards were also seen entering dens on a few occasions. The remains of two foxes, with the flesh stripped off neatly from the bones, were found, indicating that the kills were made by birds of prey. A local reported seeing a dog killing a fox (during the epidemic), but we observed play between a fox and a half grown dog (at a good distance away from each other). The local people do not eat the fox, but two communities, the Pardhis based at Nandikotkur and a nomadic beggar community, hunt and eat them. We recorded three dens that were smoked and dug out in the grazing land. Poachers do not hunt in or near the enclosures, for fear of being caught by the Forest Department. Discussion Fox populations often increase steadily with the years, reach levels of overpopulation or saturation, and then decline rapidly due to epidemics (Rausch 1958, Prater 1980, Wandeler et al. 1974, Malcolm 1986, and Ginsberg and Macdonald 1990). In RWS, the population of the fox had increased from half a dozen animals during 1985-87 (Manakadan and Rahmani 1987) to about 40-50 animals during 1992-94. It then dropped down to about 1 0 animals in 1 995 due to an epidemic. Canine distemper and rabies are common among canids and could be an important factor in controlling populations, especially of the fox, due to their greater numbers and density (Mech 1970, Wandeler et al. 1974 and Malcolm 1986). The increase in population of the fox in the Sanctuary could have been a natural occurrence, or brought about by the protection of the species and its habitat after the establishment of the Sanctuary. Scrub control is suggested as a manage- ment tool to aid detection and avoidance of terrestrial predators of the San Joaquin kit fox Vulpes macrotis mutica (Warrick and Cypher 1998) and the desert kit fox Vulpes macrotis arsipus (Zoellick et al 1998). Tree and shrub growth at RWS has increased significantly, especially bordering streams (Manakadan and Rahmani 1997), and the fox or its dens were not recorded in such habitats. Scrub control appears necessary in such areas, as it gives cover to potential predators of the fox, such as wolf, jackal and jungle cat, to stalk the species. The fox was recorded in light scrub areas, which appear important for resting and shelter during the day (especially during the non-denning period), and may be vital to the species to escape aerial predators (such as eagles), especially in over- grazed or burnt areas. Digging of dens in trench cum mound (TCM) walls is easier due to the loose soil and mbble on the trenches, and this may explain the concentration of dens in the enclosures and TCM walls. Most areas of grazing land had shallow soil, exposed rock beds and a calcareous layer, which made digging of dens difficult. In the case of the Arctic fox A lope x lagopus, Eberhardt et al. (1982) mentioned that den sites were restricted to areas where the permafrost was sufficiently deep and soil characteristics allowed burrowing. It is also likely that absence of poaching results in the concentration of dens in an area. This is because the young have greater chances of survival, and on maturity, some of them dig dens in the vicinity of their parents’ dens, especially since foxes are social canids. This may explain the clumped distribution of dens and den groups in the protected enclosures, in contrast to relatively dispersed distribution in the grazing land. Although TCM walls may attract the fox for denning, it is primarily protection, habitat improvement and lack of disturbance that have attracted them to the enclosures. This explains why dens were concentrated in Enclosure-I and II (protected plots), but not in Enclosure-Ill (unprotected). Malcolm (1986). and Ginsberg 12 JOURNAL, BOMBAY NATURAL HISTORY SOCIETY. 97(1), APR. 2000 POPULA TIONAND ECOLOGY OF THE INDIAN FOX and Macdonald ( 1 990 - quoting various sources) reason that clumping of fox dens is an indication of good habitat. Trottier (1992) mentioned that the swift fox Vulpes velox prefers grass of moderate height. This may also be true for the Indian fox, for respite from the heat and protection from aerial predators, especially eagles. The protected enclosures were relatively free of human disturbance. Foxes in the grazing land were frequently disturbed by the movements of people, graziers and dogs. During our visits to the dens, sightings of animals were common around den sites in the enclosures, but were rare in the grazing land. It was not clear whether the foxes in the grazing land come out of their dens only at dusk to avoid frequent disturbance (and were hence missed during our visits), or they took refuge elsewhere during the day. Multiple den use is likely to be both, a strategy to confuse predators (jackal and wolf) and for sanitation. In the desert kit fox Vulpes macrotis cirsipus, individuals were reported to use 3-16 dens, while pahs use 9-16 dens (Zoellick et al. 1998). Canids are known to move their pups regularly to different dens (Sargeant et al. 1975), and this has also been reported in the Indian fox (Johnsingh 1978). Sargeant et al. (1975) recorded splitting of litters among two or more dens in the red fox Vulpes vulpes. In this study, it was observed that usually after breeding and occupancy of a den for about two months, die pair shifted to another den nearby and even to a third den later on. Half grown pups then frequent all such dens of the den group. Johnsmgh (1978), from his studies in Madurai dist., Tamil Nadu, recorded dens with either two holes or the more common multiple opening dens (maximum of 23 holes). In this study, except for a few single hole dens, the rest were multiple hole dens (up to 43 openings). A greater number of holes per den probably indicates the use of the dens by the same pair for many years, as stated by Johnsingh (1978). However, unlike Johnsingh’ s findings, areas around dens in RWS had less vegetation compared to the surrounding areas. This is because the soil at RWS has a calcareous layer. This layer when brought to the surface by the foxes digging, hinders plant growth. The extent of predation on bustard eggs and chicks by the fox was not established. Remains of eggs or chicks were not recorded in the scats analysed, probably because most scats were not collected in the major breeding season of the bustard. It is also unlikely for egg shell pieces to appear in the scats, as the fox might lick the egg contents and leave the shell. In some cases of nest predation recorded during this and the earlier study [predator not known] shell pieces were found strewn around the nest sites. As for chicks, not much identifiable matter could be expected in the scats, except for the bill or claws. A major drawback of our studies on the Indian fox was that we could not investigate the nocturnal activities of this largely nocturnal species. Also, it was not possible to identify individuals from body characteristics since the animals were not marked. A study of radio- collared animals with the help of night vision equipment is essential to get precise information on the species. Acknowledgements This study is a part of the Grassland Ecology Project of the Bombay Natural History Society and the Centre of Wildlife & Ornithology, Aligarh Muslim University, funded by the U.S. Fish and Wildlife Service, and sponsored by the Ministry of Environment and Forests, Govt, of India. We thank the Andhra Pradesh Forest Department for permission to work in the Sanctuary, and the cooperation and help rendered by the staff of Rollapadu Wildlife Sanctuary. JOURNAL, BOMBAY NATURAL HISTORY SOCIETY, 97(1), APR. 2000 13 POP U LA TION AND ECOLOG Y OF THE INDIA N FOX References Eberhardt, L.E., W.C. Hanson, J. Bergston, R.A. Garrot & E.R. Hanson (1982): Arctic fox home range characteristics in an oil development area. J. Wild/. Manage. 46: 183-190. Ginsberg, J.R. & D.W. Macdonald ( 1 990): Foxes, wolves, jackals and dogs. An action plan for the conservation of canids. IUCN, Gland, Switzerland. Johnsingh, A.J.T. ( 1 978): Some aspects of the ecology and behaviour of the Indian fox Vulpes bengalensis. J. Bombay nat. Hist. Soc. 75: 397-405. Malcolm, J.R. (1986): Socio-ecology of bat-eared fox ( Otocyon megalotis). J. Zool. London (A) 206: 457- 467. Manakadan, R. & A.R. Rahmani (1989): Rollapadu Wildlife Sanctuary. J. Bombay nat. Hist. Soc. 86: 368-380 Manakadan, R. & A.R. Rahmani (1990): Growth and development of a captive great Indian bustard chick. Avicidtural Magazine 96: 133-140. Manakadan, R. & A.R. Rahmani (1993): A decade of conservation of the great Indian bustard at Rollapadu Wildlife Sanctuary, Kurnool district, Andhra Pradesh. Proc. Changing Scenario of Bird Ecology and Conservation (Ed: A. Verghese, S. Sridhar & A.K. Chakravarthy), Ornithological Society of India, Bangalore. Manakadan. R. & A.R. Rahmani (1997): Rollapadu Wildlife Sanctuary (pp: 1 17-180). In: A study of the ecology of grasslands of the Indian plains with particular reference to their endangered fauna. Final Report, (Ed: A. R. Rahmani). Bombay Natural History Society. Mumbai. Pp 549. Mech. L.D. (1970): The Wolf : Ecology and Behaviour of an Endangered Species. Natural History Press, Doubleday. New York. Prater, S.H. (1980): The Book of Indian Animals. 3rd Edition. Bombay Natural History Society, Bombay. Rahmani, A.R. (1989): The Great Indian Bustard. Final Report. Bombay Natural History Society. Bombay. Rahmani, A.R. & R. Manakadan (1986): Study of the Ecology of Certain Endangered Species of Wildlife and their Habitats: The Great Indian Bustard. Rollapadu Wildlife Sanctuary. Bombay Natural History Society, Bombay. Rahmani, A.R. & R. Manakadan (1987): Interspecific behaviour of the Great Indian Bustard Ardeotis nigriceps. J. Bombay nat. Hist. Soc. 83: 17-31. Rausch, R. (1958): Some observations on rabies in Alaska, with special reference to wild canids. J. Wild!. Manage. 22: 246-260. Sargeant, A.B., W.K. Pfeifer & S.H Allen (1975): A spring aerial census of red foxes in North Dakota. J. Wildl. Manage. 39: 30-39. Trottier, G.C. (1992): Conservation of Canadian Prairie Grasslands: A Landowner’s Guide. Canadian Wildlife Service, Canada. Wandeler, A., .1. Muller, G. Wachendorfer. W. Schale, U. Forster & F. Stack (1974): Rabies in wild carnivores in Central Europe. Ill Ecology and biology of the fox in relation to control operations. Zbl. Vet. Med. B. 21: 765-773. Warrick, G.D. & B.L. Cypher (1998): Factors affecting the spatial distribution of San Joaquin Kit Foxes. J. Wildl. Manage. 62: 707-717. Zoellick, B.W., N.S. Smith & R.S. Henry ( 1 989): Habitat use and movements of Desert Kit Foxes in Western Arizona J. Wildl. Manage. 53: 955-961 . ■ ■ ■ 14 JOURNAL, BOMBAY NATURAL HISTORY SOCIETY, 97(1). APR. 2000 BREEDING BIOLOGY OF THE MALABAR GREY HORNBILL {OCYCEROS GRISEUS) IN SOUTHERN WESTERN GHATS, INDIA.' Divya Mudappa2 ( With one text-figure) Key words: hornbill, tropical rainforest, frugivory, seed dispersal, cavity- nesting, breeding biology The Malabar grey hornbill ( Ocyceros griseus) is a frugivore, endemic to the tropical rainforests and moist deciduous forests of the Western Ghats hill ranges, India. I studied its breeding biology in the Anamalai hills (Indira Gandhi Wildlife Sanctuary), Tamil Nadu state, by monitoring 10 nests and their middens, and conducting intensive observations at a focal nest. The nesting period lasted an average of 86 days (N=4), and observations at the focal nest revealed the pre- and post-hatching phases to be 40 and 46 days, respectively. At the end of the nesting period, the females and the young simultaneously broke out of the nests. A total of 2397 items of food were delivered by the male hornbill to the inmates of the focal nest. They included 6 species of lipid-rich and 8 species of sugar-rich fruits, and at least 14 kinds of animal matter. Lipid-rich fruits formed a major component (c. 37%) of the diet during nesting. Ficus fruits formed 26%, and animal matter 13.8% of the diet of the incarcerated hornbills. The frequency of sugar- and lipid-rich fruits delivered per hour of observation was significantly greater in the pre-hatching phase. While the frequency of animal food delivered was higher in the post-hatching phase. Although the Malabar grey hornbill used a wide range of food resources, it was observed that a few species of rare, tropical trees producing lipid-rich fruits during the nesting period, play an important role in the maintenance of the species. Introduction Hornbills (Aves : Bucerotidae and Bucorvidae) are a group of large, forest and savanna birds restricted to the Old World tropics. There are 54 species of hornbills in the world (Kemp 1988, 1995), nine of which occur in India (Ali and Ripley 1987). Only in the last two decades, a few studies have provided valuable insights into the ecology of these unique cavity- nesting birds (Hussain 1984, Kannan 1994, Kemp 1976, 1978, 1988, Kinnaird 1993, Leighton 1982, Poonswad 1995, Poonswad and Tsuji 1989, 1994, Reddy et al. 1990, Reddy and 'Accepted June, 1 998 :Centre for Ecological Research and Conservation 3076/5 IV Cross, Gokulam Park Mysore 570 002, Karnataka, India. Basalingappa 1995). Hornbills are secondary cavity-nesters, and the forest-dwelling species are predominantly fmgivorous. Their breeding cycles are synchronous with food productivity of the forest (i.e., fruiting phenology; Kannan 1994), but they are also dependent on keystone resources like Ficus for their survival in times of low food availability. They exhibit wide-ranging movements to meet their specialized food requirements (Poonswad 1994). Functionally, they have been described as keystone mutualists (Gilbert 1980) as they play an important role in the dispersal of many rare rainforest tree species (Kinnaird 1998, Whitney et al. 1998). The present study aimed to determine the nesting habitat requirements and breeding biology of the Malabar grey hornbill, endemic to the Western Ghats. The former aspect is dealt JOURNAL, BOMBAY NATURAL HISTORY SOCIETY, 97(1), APR. 2000 15 BREEDING BIOLOGY OF THE MALABAR GREY HORNBILL with elsewhere (Mudappa and Kannan 1997). This paper describes in detail: 1) nesting activities and behaviour of the male and female hornbill, 2) duration of nesting period and distinct phases of the nesting cycle, and 3) qualitative and quantitative data on the food delivered by the male to the incarcerated female and young, in relation to the phases of the nesting period. The results are compared with other hornbill species, their reproduction and survival strategies, and the implications for the conservation of this rainforest endemic are discussed. Study Area The study was undertaken between December 1993 and May 1994 at the Indira Gandhi Wildlife Sanctuary (10° 13' - 10°33' N and 76°49' - 77°21' E, an area of 968 km2) in the Anamalai Hills of the southern Western Ghats, in Tamil Nadu state, India. A one-month long preliminary study was carried out in the area in May- June 1993, when 15 nests were discovered and seeds from the middens were collected and identified for future reference. The nests selected for intensive observation and monitoring were in the 5.1 km2 wet evergreen' forest patch of Karian Shola National Park. This forest, classified as a Southern Tropical Wet Evergreen Forest (Champion and Seth 1968), receives an annual rainfall of about 1500 mm. The terrain is hilly, and the altitude ranges from 350 m to 2400 m above msl in the Sanctuary, which extends into Parambikulam Wildlife Sanctuary and Eravikulam National Park in the adjacent Kerala state. The forest is contiguous with moist deciduous, teak ( Tectona grandis) and bamboo forests in the surrounding areas. Study Species Of the 9 species of hornbills in India, the Malabar grey hornbill ( Ocyceros griseus), is the smallest. It is endemic to the Indian subcontinent. occuring only in the heavy rainfall tracts of the Western Ghats hill ranges. Most of the information on the Malabar grey hornbill and other Indian hornbills is anecdotal- with notes on natural history. Early papers dealing with nidification of the Malabar and the common grey hornbills ( Ocyceros birostris) are those of Bingham (1879), Hall (1918), Lowther (1942), and Abdulali (1942). More comprehensive information on their ecology and behaviour was provided by Ali and Ripley (1970, 1987) and Kemp (1978). The Malabar grey hornbill is sexually dimorphic: the male has a large, bright orange bill and golden brown iris, while the female has a relatively small and pale-coloured bill and dark brown iris. The species is monogamous, the nesting pair usually exhibiting high nest-site fidelity, occupying the same nest-cavities every year (Kemp 1978, Ali and Ripley 1987, Mudappa and Kannan 1997). The Malabar grey hornbill exhibits biparental care like most other monogamous birds with altricial young (Clutton- Brock 1991). While the incubating female is incarcerated, the male provisions her and the other inmates of the nest. Methods Active nests of the Malabar grey hornbill were located with the help of a local field assistant, by following the parent birds, and by checking for signs of previous nesting, such as seeds and faecal remains (midden) at the base of the nest trees. Fifteen nests were located during the preliminary study in May 1993. Seeds collected from the midden were catalogued and used for reference during the study. Twelve additional nests were discovered during the initial half of the study (December 1 993 to March 1994). Ten nests were chosen for monitoring during the nesting period (the period of incarceration of the female and the young) in Karian Shola National Park. Of these, one was 16 JOURNAL, BOMBAY NATURAL HISTORY SOCIETY, 97(1), APR. 2000 BREEDING BIOLOGY OF THE MALABAR GREY HORN BILL selected on the basis of logistics for intensive observation. Intensive observation of a focal nest: The ✓ focal nest was observed from the last week of January 1994 to May 1994, for approximately six-hour intervals on every alternate day (more or less uniformly) throughout the three-month nesting period. Observations were made between 0700 h and 1300 h. The forenoon was chosen for nest observation, while the rest of the day was used to visit the other nine nests. I observed the nest to gather information on the quantity and quality of food delivered by the male to the incarcerated inmates. The food was broadly classified as plant and animal food. The plant food was further categorised as: a) figs, b) sugar-rich non- fig, and c) lipid-rich fruits, based on McKey (1975) and Snow (1981). Observations were made from a ground hide about 18m from the base of the focal nest through a 7x50 binoculars or a 20x50 spotting scope. For each visit by the male hornbill to the nest, I recorded the number and type of food items delivered, the duration of the visit (to the nearest 5 seconds), and the total number of visits during each sampling/observation session. Ad libitum observations on other activities like nest-cavity sealing, cleaning, excretion, begging by the inmates, and the behaviour of the male during the time of food delivery were recorded. At the end of each session, the seeds and other faecal remains in the midden were examined, identified, classified, and counted. Nest midden monitoring: Ten nests (including the focal nest) were visited regularly to note the status of nesting, quantify the regurgitated or excreted seeds of the fruits eaten by the inmates, and to identify the other debris in the midden. Of the food items consumed by the nest-cavity inmates, only non-digestible parts such as seeds of fruits, elytra of insects, and reptile scales occur in the midden. All distinguishable midden remains were collected, identified, counted, and recorded. The midden below the nest-tree was cleared of all debris after each visit. Small seeds and animal matter in the faecal remains could not be quantified. The presence of Malabar grey hornbill feathers in the midden was taken to indicate moulting. Similarly, the presence of egg-shell in the midden, or the characteristic begging calls of the young, were evidence of hatching or the presence of chick(s) in the nest. Statistical analyses: The frequency of food items delivered during the nesting period was calculated. Differences between the food (type and quantity) consumed between the two distinct phases (pre- and post-hatching) of the nesting period were tested for statistical significance using Mann- Whitney U test (Seigel and Castellan 1988), using SPSS/PC+ computer software (Norusis 1990). The difference in the occurrence of seeds (frequency) in the midden was tested for significance, using the non- parametric Mann- Whitney U Test similar to the analysis of direct feeding observation. Results Characteristics and occupation of focal nest: The focal nest cavity was located at about 14 m on an Artocarpus lakoocha (Moraceae) tree. The diameter at breast height (1.2 m) of the focal nest tree was 56 cm, the height 25 m, and the estimated diameter at nest height was 50 cm. The cavity entrance was circular in shape, and oriented towards northwest. My field assistant observed a bird entering the nest cavity in the first week of February. This was probably an instance of nest preparation, cleaning, and widening of the nest entrance. After this, there was regular movement of the breeding pair in the vicinity of the nest- tree. On February 17, the female hornbill was seen entering the nest-cavity. The cavity entrance was then half-sealed. The male and the female visited the nest (8 times in 6 hrs). During these visits, they appeared to be enlarging the cavity entrance. JOURNAL , BOMBAY NATURAL HISTORY SOCIETY, 97(1), APR. 2000 17 BREEDING BIOLOGY OF THE MALABAR GREY HORNBILL The female was in the nest on February 18, and was seen sealing the cavity entrance, leaving only a slit, through which the male fed the inmates during the nesting period. The male was never observed to be involved in nest sealing, repair, or delivering any kind of sealing material, unlike the female which often repaired the seal with its bill. The female was seen cleaning the nest-cavity by throwing out a lot of seeds and woody debris. The female hornbill used her own excreta, rich in Ficus seeds, as material for sealing the cavity entrance. The inmates effected nest sanitation by squirting their excreta out through the slit-like opening of the cavity entrance. Nesting period: The nesting season lasted for about three months, between February and May in the study population of the Malabar grey hornbills. The nesting period could be distinguished into two main phases: the pre- hatching and the post-hatching phase. However, each phase in turn has been further divided into 3 sub-phases (fortnightly) for analysis. The nesting period in the focal nest was 86 days, commencing from February 18 (incarceration of the female) to May 15 (emergence of chick and female from the nest). The mean duration of the nesting period was 86 days (± 2.7 S.D.; N=4). In the focal nest, the young hatched 40 days after the incarceration of the female. The post- hatching phase was 46 days. Only one chick appeared to have fledged. The female and young broke out of the nest together. Details of the nesting period in the ten nests are given in Table 1 . Clutch size and moulting: The clutch size in the breeding population could not be determined. In the focal nest, only one young was seen. One nest when examined on March 1 , 1994, had only one egg. A week later, there were two eggs in this nest. The female resealed the cavity entrance and bred successfully. Flight feathers were collected from the midden occasionally, particularly in the month of April. The rectrices were never found and the Table 1 DATES OF INCARCERATION AND FLEDGING IN THE STUDY NESTS Nest number Date of incarceration Fledging date 1. 1 7 February 1 6 May 2. 1 5 February 3 May 3. 1 8 February 13 May* 4. 1 8 February 15 May* 5. 21 February 1 6 May 6. 1 8 February 1 8 April** 7. 4 March* 1 1 May* 8. 1 7 March* 13 May 9. 3 March* 1 6 May 10. 1 8 February 1 5 May * — The chick fledged between this day and 20 May ** — Abandoned * - Nests discovered after the nesting had commenced female of the focal nest had tail feathers throughout the nesting period. These could be seen while the bird was ejecting the faecal matter through ‘the slit. However, rectrices had been collected from the midden of six nests during the preliminary study in 1993. Thus, it is likely that the moult in this species is partial. Food delivery by the male hornbill: The focal nest was observed for a total of 161 hours and 45 minutes. All through the nesting period, the male provisioned the incarcerated female and later, the young also. A total of 2,397 food items, which included 1 1 kinds of fruit, 5 species of vertebrates, and at least 8 types of invertebrates, including 6 types of insects, were delivered by the male (Appendix). Lipid-rich fruits predominated in the diet of the incarcerated hornbills, constituting 36.9% of the food delivered. Other food categories were Ficus 26%, sugar-rich fruits 22.6%, and animal matter 13.8%. If there were several items, these were regurgitated one by one. Large fruits and vertebrate prey were usually brought as single items. The number of food items delivered peaked during the pre-hatching phase, and declined thereafter, being minimum before the fledging of the young. The frequency of lipid-rich and 18 JOURNAL, BOMBAY NATURAL HISTORY SOCIETY. 97(1). APR. 2000 BREEDING BIOLOGY OF THE MALABAR GREY HORN BILL APPENDIX PLANT AND ANIMAL FOOD DELIVERED AT THE NEST BY THE MALE MALABAR GREY HORNBILL A: Plant food (fruit) S.No. Species Habit Number Number (Family) in pre- in post- hatching hatching phase phase Sugai 1. r-rich Fruit Ficus spp. (Moraceae) Tree/ Strangler 443 123 2. Mi mu sops elengi (Sapotaceae) Tree 17 _ 3. B ridel ia sp. (Eupborbiaceae) Climber 417 13 4. Elaegnus conferta (Elaegnaceae) Climber 4 _ 5. Linocera intermedia (Sapindaceae)+ Tree 6. Syzygium spp. (Myrtaceae)+ Tree _ 7. Filicium decipiens (Oleaceae)* Tree _ 8. Zizyphus nummularia (Rhamnaceae) Shrub 61 9. Glycosmis pentaphylla (Rutaceae) Shrub 11 _ Lipid-rich Fruit 10. Uvaria sp. (Annonaceae) Climber 510 63 11. Neolitsea sp. (Lauraceae) Tree 173 52 12. Cinnamomum sp. (Lauraceae) Tree 13. Persea macarantlia (Lauraceae)* Tree _ 14. Litsea sp. (Lauraceae )+ Tree _ _ 15. Beilschmedia sp. (Lauraceae) Tree _ 19 16. Myristica dactyloides (Myristicaceae)* Tree A: Plant food (fruit) (contd.) S.No. Species (Family) Flabit Number in pre- hatching phase Number in post- hatching phase 17. Knema attenuate (Myristicaceae)* Tree 18. Polya Ithia sp. (Annonaceae)+ Tree - - 19. 20. Other Fruits Strychnos nux-vomica ( Logan iaceae)* Unidentified** Tree - + — Found in the midden of the focal nest * — Found in the middens of other (non-focal) nests ** — Ten species whose seeds were found in small numbers in the middens (three were found in the midden of the focal nest) B: Animal Food Vertebrates 1. Young bird 2. Snake 3. Lizard ( Calotes sp.) 4. Gecko 5. Frog Invertebrates 1 . Beetle 2. Cricket/Grasshopper 3. Cicada 4. Stick Insect 5. Caterpillars 6. Winged insect (wasp, termite, etc.) 7. Millipede/Centipede 8. Scorpions Total number of animal food items delivered during the nesting period = 491. non-fig sugar-rich fruits was significantly higher in the pre-hatching phases (Mann- Whitney U test, N=16, U=24, pO.OOl and U=36, p<0.001, respectively). Figs were eaten consistently throughout the nesting period. The frequency (number per hour of observation) of animal matter delivered was greater in the post-hatching phase (Mann-Whitney U test, U=41, p=0.047 for invertebrates and U=64, p=0,014 for vertebrates; Fig. 1). Within the pre-hatching phase, the frequency of lipid-rich fruits was significantly higher than the other types (Kruskal-Wallis JOURNAL BOMBAY NATURAL HISTORY SOCIETY, 97(1), APR. 2000 19 Minutes/hour Number/hour Minutes/hour BREEDING BIO LOG Y OF THE MA LA BA R GRE Y HORNBIL L Fig. 1 : a. Time spent at nest by the male, b. Visiting rate of the male, and c. Frequency of different food items delivered to the inmates by the male during the nesting period. 20 JOURNAL . BOMBAY NATURAL HISTORY SOCIETY. 97(1). APR. 2000 BREEDING BIOLOGY OF THE MALABAR GREY HORN BILL X2= 1 3 .48, df=3, p<0.001), while in the post- hatching phase, animal food was significantly greater (%2=23.26, df=3, p<0.001). Time spent at nest and visitation rate of the male: The time spent and the visitation rate of the male hornbill was influenced by the number and type of items delivered. Time spent (minutes per hour of observation) was significantly higher (Mann- Whitney U test, U=38, p<0.01) in the pre-hatching than in the post-hatching phase, as a greater number (69.5%) of small fruits (< 1.5 cm) was delivered (each had to be regurgitated individually). The visitation rate did not differ between the phases (Fig. 1). Feeding habits — evidence from middens: Supplementary data from the middens of the ten nests showed that nine additional species of fruits were consumed by the incarcerated hornbills (e.g. Strychnos nux- vomica, Litsea sp., Persea macarantha, see Appendix). A few seeds of ten unidentified plant species were collected from some middens. There was no significant difference between the pre- and post-hatching phases in the frequency of the lipid-rich fruit seeds collected in the midden. The frequency of non-fig sugar-rich fruit seeds in the midden was found to be significantly greater in the pre-hatching phase (Mann- Whitney U test, N=21, U=T 16, p=0.007). Predation on Malabar grey hornbill and nest intrusion: Two cases of mortality of Malabar grey hornbills were recorded. The first was of a young bird found towards the end of the nesting period during the preliminary study in 1993. The second was presumably an adult, whose remains were found in the middle of the nesting period in 1994, close to a regularly monitored nest which had been abandoned five days earlier. The focal nest was once visited by three hill mynas ( Gracula religiosa ) that flew away at the approach of the male hornbill. A Malabar giant squirrel ( Ratufa indica) and the dusky- striped palm squirrel ( Funambulus sublineatus ) were other inquisitive visitors to the nest, but were apparently disregarded by the incarcerated female. Discussion The 32 species of Oriental hornbills are essentially forest-dwelling, arboreal birds (Kemp 1988, 1995). These species, including the Malabar grey hornbill, are long-lived, and have a distinct and relatively long nesting period. The nesting period of the Malabar grey hornbill lasted an average of 86 (± 2.7 days) during this study. The success of this bird as a rainforest specialist can be attributed to its life-history strategies (the long and peculiar nesting behaviour), and the adaptation in food habits. Predation of adult Malabar grey hornbills by animals other than man is rare. Even during the vulnerable period of incarceration, the chances of predation are low, because the nest- cavity entrance is sealed, and the female with her large, armoured bill can protect the nest from intruders. This protection, along with the cavity nesting habit, can be the reason for the long incubation period of these birds. Overall, the nesting periocl and food delivery by the Malabar grey hornbill in the area, as in the case of great pied hornbill ( Buceros bicornis ), seems to be associated with fruiting phenology, and the onset of the southwest monsoon (Kannan 1994). Studies in Thailand (Poonswad et al. 1988) have found the nesting of hornbills to commence and terminate later than in this region, probably because of the later monsoon. Hornbills subsist on an array of diverse, locally rare, tree species (e.g. members of the Lauraceae; Kannan and James 1999). The nesting period coincides with the peak in fruit availability, as shown by the fruiting phenology study of Kannan and James (1999). Large numbers of rainforest trees of the families Lauraceae, Burseraceae, and Myristicaceae {op cit.) contribute to the abundance of fruit. JOURNAL, BOMBAY NATURAL HISTORY SOCIETY, 97(1), APR. 2000 21 BREEDING BIOLOGY OF THE MALABAR GREY HORN BILL Lipid-rich fruits formed the most abundant component of the food delivered. The coincidence of nesting with peak in lipid-rich fruit availability could be as a result of long-term co-evolutionary process (McKey 1975). The high lipid content of these fruits may be necessary to meet the requirements of the nesting, moulting, and growing birds (Snow 1981). Protein, carbohydrate, and water is obtained from sugar-rich fruits (including figs) and animal matter, which supplement the lipid- rich diet of the nesting hornbills. Notably, the Malabar grey hornbill fed less on Ficus fruits (26%) than the great pied, oriental pied (Anthracoceros coronatus ), and wreathed {Aceros undulatus) hornbills (Kannan and James 1997, Tsuji 1996). The smaller white-throated brown hornbill ( Ptilolaemus tickelli ), however, is shown to feed less on figs. Smaller-sized hornbills are able to feed on a wider range of fruit and animal food, probably due to their smaller body size which enables them to access even the understorey shrub species, thus reducing the predominance of any one type of food. The Malabar grey hornbill consumes a greater variety of sugar-rich, particularly understorey fruits, as well as fruits of small trees and climbers, unlike the larger syntopic great pied hornbill which prefers large, canopy and emergent trees (Kannan 1994). A wide range of food items are fed to the nest inmates. The kind of food delivered influenced the visitation -rate, and the time spent at nest by the male. The time spent was significantly higher in the pre-hatching phase as there was a greater number of small fruits (both lipid- and sugar-rich fruits, i.e., 61% of all small fruits) delivered at the nest. The time spent at the nest decreased towards the end of the nesting period, when large fruits and animal food were brought for the inmates and delivered as a single item per visit. The visitation rate did not differ between the phases, though the number of fruits delivered per visit decreased in the post-hatching phase. This was probably compensated by the nutritive quality (lipid-rich fruits and animal food), and larger size of the food items delivered (eg. fruits of Myristica sp., Beilschmedia spp.). There was a drastic fall in the number of visits during the last few days of the nesting period. Welty (1982) proposed that the steady decline in feeding frequency may be a naturally evolved strategy of the parent to encourage the nearly- fledged young to leave the nest. The differences in the food delivered during the nesting period can be explained by one or a combination of the following factors: (i) It could be related to the availability of fruits due to the usually high seasonal and synchronous fruiting of tree species bearing lipid-rich fruits (Snow 1981, Leighton and Leighton 1983, Kannan and James 1999), while the sugar-rich fruits are available all through the year. Community fruiting patterns in the study area were found to be largely determined by the trees producing lip id-rich fruits like Lauraceae, Annonaceae, which form a major proportion of tree species in the area (Kannan 1994). It was observed that certain fruits such as Alseodaphne semecarpifolia , Litsea sp., and Persea macaranthci , which were common and abundant in the middens during the preliminary study in 1993, were absent in 1994. So, inter-annual differences in fruiting patterns, and intra- seasonal staggering in the fruiting patterns of the Lauraceae in the rainforests is likely to play a major role in the nesting and nesting success of the hornbills (Snow 1981, Leighton 1982, Leighton and Leighton 1983, Kannan and James 1999). (ii) Another possibility is that the hornbill selects high quality nutritive food for the growing chicks in the post-hatching phase, feeding them largely lipid-rich fruits and animal matter, which may be of co-evolutionary significance. The increased delivery of animal food toward the end of the nesting season may reflect an increase in abundance of insect prey in the forest just after 22 JOURNAL BOMBAY NATURAL HISTORY SOCIETY, 97(1), APR. 2000 BREEDING BIOLOGY OF THE MALABAR GREY HORNBILL the rains. The supplementation of high quality animal matter, however, coincides with the hatching of the chick and may provide the growing chick with essential nutrients. (iii) Hombills are known to be territorial, ranging between 3 to 30 km2 (white-throated brown and great pied hornbills, respectively) depending on the size of the bird (Poonswad and Tsuji 1994). Seeds of some fruits (eg. Filicium decipiens , Polyalthia sp.) were found in the middens of only a few nests, probably because these fruiting trees were abundant in the territories of the hornbills inhabiting those nests. Conclusion The Western Ghats have been identified as one of the biodiversity hotspots in the world (Myers 1990, 1991). However, large scale deforestation for dam construction, agriculture and other developmental activities has resulted in the loss of over 40% forest cover in the last 70 years (Chattopadhyay 1985, Menon and Bawa 1997). This in turn has restricted the range of many species, including many endemics such as Refer Abdulali, H. (1942): The nesting of the Malabar Grey Hornbill. J. Bombay nat. Hist. Soc. 43:102-103. Ali, S. & S. D. Ripley. (1970): Handbook of the birds of India and Pakistan. Vol 4. Oxford University Press. Pp. 130-133. All S. & S. D. Ripley. ( 1 987): Compact handbook of the birds of India and Pakistan. Oxford University Press, Delhi. Bingham, C. T. (1879) Notes on the nidification of some hornbills. Stray Feathers 5:459-463. Chattopadhyay, S. (1985): Deforestation in parts of Western Ghats region (Kerala), India. J. Environ. Manage. 20: 219-230. Champion, H. G. & S. K. Seth (1968): A Revised Survey of the Forest types of India. Manager of Publications, Government of India, New Delhi. Clutton-Brock, T. H. (1991): The evolution of parental care. Princeton University Press, Princeton. Gilbert, L. E. (1980): Food-web organization and the the Malabar grey hornbill. Hornbills play an important role in the dynamics of their habitats because of their specialised frugivorous habits (McKey 1975, Snow 1981, Leighton 1982) and as effective dispersers of many tree species (Kinnaird 1998, Whitney et al. 1998). The Malabar grey hornbill, like other members of the family Bucerotidae, act as keystone species in the range of its distribution (Gilbert 1980). This endemic, specialist frugivore of the rainforest of the Western Ghats plays an important role in the dynamics of the moist evergreen forest it inhabits, dispersing the seeds of a few rare rainforest tree species. Conservation of their habitat is imperative as they have specialised feeding and nesting requirements (Mudappa and Kannan 1997). ACKNO W LEDG EM ENTS The research was supported by a grant from the Oriental Bird Club, U. K. I thank R. Kannan for guidance and encouragement and Ganesh, my assistant, for help in field work. I thank the Tamil Nadu Forest Department for permission to carry out the study. ENC ES conservation of neotropical diversity. Pp. 1 1-34. In Soule M. E. and Wilcox, B. A. (eds.). Conservation Biology, Sinauer, Sunderland, Massachusetts. Hall, E. F. (1 91 8): Notes on the nidification of the Common Grey Hornbill (Lophoceros birostris). J ’. Bombay nat. Hist. Soc. 25: 503-505. Hussain, S. A. (1984): Some aspects of the biology and ecology of Narcondam Hornbill ( Rhyticeros narcondami). J. Bombay nat. Hist. Soc. 81: 1-18. Kannan, R. (1994): Ecology and conservation of the Great Pied Hornbill ( Buceros bicornis) in the Western Ghats of southern India. Unpubl. Ph. D. thesis, University of Arkansas, Arkansas. Kannan, R. & D. A. James (1997): Breeding biology of the great pied hornbill ( Buceros bicornis ) in the Anamalai hills of southern India. J. Bombay nat. Hist. Soc. 94: 451-465. Kannan, R. &D.A. James (1999): Fruiting phenology and the conservation of the great pied hornbill ( Buceros JOURNAL. BOMBAY NATURAL HISTORY SOCIETY. 97(1), APR. 2000 23 BREEDING BIOLOGY OF THE MALABAR GREY HORN BILL bicornis ) in the Western Ghats of Southern India. Biotropica 31 : 167-177. Kemp, A. C. (1976): A study of the ecology, behaviour and systematics of Tockus hombills (Aves: Bucerotidae). The Transvaal Mus. Memoir 20. Kemp, A. C. ( 1 978): A review of the hombills: biology and radiation. The Living Bird 17: 105-136. Kemp, A. C. ( 1 988): The systematics and zoogeography of Oriental and Australasian hombills (Aves: Bucerotidae). Bonn. zool. Beitr. 39:315-345. Kemp, A.C. (1995): The hombills. Oxford University Press. Oxford, England Kemp, A. C. & M. I. Kemp (1975): Report on a study of hombills in Sarawak, with comments on their conservation. World Wildlife Fund Project Report 2/ 74. Kinnaird, M. F. (1993): Variation in fruit resources and the effects on vertebrate frugivores: the role of disturbance regimes in Sulawesi rainforests. Report, LIPI. Jakarta, Indonesia. Kinnaird, M. F. (1998): Evidence for effective seed dispersal by the Sulawesi red-knobbed hornbill Aceros cassidix. Biotropica 30: 50-55. Leighton, M. (1982): Fruit resource and patterns of feeding, spacing and grouping among sympatric Bornean hombills (Bucerotidae). Unpubl. Ph. D. thesis, University of California, Davis. Leighton, M. & D. R. Leighton (1983): Vertebrate responses to fruiting seasonality within a Bornean rainforest, pp 181-196. In: Sutton, S. L., Whitmore, T. C., and Chadwick, A. C. (eds.) Tropical Rain Forest: Ecology and Management. Blackwell Scientific Publications, Oxford Lowther, E. H. N. (1942): Notes on some Indian birds. J. Bombay nat. Hist. Soc. 43:386-401. McKey, D. ( 1 975): The ecology of coevolved seed-dispersal system. Pp. 159-199. In Gilbert L. E. and P. H. Raven (eds.). Coevolution of animals and plants. University of Texas Press, Austin. Menon, S. & K. S. Bawa (1997): Applications of geographical information systems, approach to bio- diversity conservation in the Western Ghats. Curr. Sci. 73: 134-145. Mudappa, D. (1994): Nesting habitat of the Malabar Grey Hornbill ( Ocyceros griseus ) in the Anaimalais, southern Western Ghats, India. M. S. dissertation, Salim Ali School of Ecology, Pondicherry University, Pondicherry. Mudappa, D. & R. Kannan ( 1 997): Nest-site selection by the Malabar Grey Hornbill ( Ocyceros griseus ) in southern Western Ghats, India. Wilson Bull. 102: \ 1 1- 119. Myers, N. ( 1 990): The biodiversity challenge: expanded hot-spots analysis. Environmentalist 10: 243-256. Myers, N. (1991): Tropical forests: present status and future outlook. Clim. Change 19: 3-12. NoruSis, M. J. (1990): SPSS/PC+: Statistics 4.0. SPSS Inc., Chicago. Poonswad, P. (1995): Nest-site characteristics of four sympatric species of hombills in Khao Yai National Park, Thailand. Ibis 137: 183-191. Poonswad, P. & A. Tsuji ( 1 989): Conservation of hombills in Thailand. Presentation at the joint meeting of ICBP Asian section and east Asian bird protection conference, 1989. Bangkok, Thailand. Poonswad, P. & A. Tsuji ( 1 994): Ranges of the males of the Great Hornbill Buceros bicornis , Brown Hornbill Ptilolaemus tickelli and Wreathed Hornbill Rhyticeros undulatus in Khao Yai National Park, Thailand. Ibis 736:79-86. Poonswad, P., A. Tsuji, R. Liewviriyakjt & N. Jirawatkavi (1988): Effects of external factors on hornbill breeding and population. World Conference on breeding endangered species in captivity, Cincinatti, Ohio. Reddy, M. S., K. S. Muralidhar, M. R. Gandhi & S. BASALiNGAPPAtl990): Distribution and variation in number of Malabar Pied Hombills Anthracoceros coronatus (Boddaert) in selected areas of north Kanara forest of Western Ghats in Karnataka (India). The Indian Zoologist 1 4:63-13. Reddy, M. S. & S. Basalingappa (1995): The food of Malabar Pied Hornbill. Jour. Ecol. Soc. 8: 23-28. Seigel, S. & N. J. Castellan, Jr., (1988): Non-parametric Statistics for the behavioural sciences. McGraw-Hill, New York. 399 pp. Snow, D. (1 98 1 ): Tropical frugivorous birds and their food plants: a world survey. Biotropica 13: 1-14. Tsuji, A. (1996): Hombills: Masters of tropical forests, Hornbill Research Foundation, Bangkok Welty, J. C. (1982): The Life of Birds. 3rd edn. Saunders College Publishing Whitney, K.D., M.K. Fogiel, A.M. Lamperti, K.M. Holbrook, D.J. Stauffer, B.D. Hardesty, V.T. Parker & T.B. Smith (1998): Seed dispersal by Ceratogymna hombills in the Dja Reserve, Cameroon. / Trop. Ecol. 74:351-371 24 JOURNAL, BOMBAY NATURAL HISTORY SOCIETY, 97(1), APR. 2000 SOCIOECONOMIC TRANSITION AND WILDLIFE CONSERVATION IN THE INDIAN TRANS -HIMALAYA1 Charudutt Mishra2,3 Key words: management, protected area, policy, livestock, Uncia uncia, Canis lupus The founding postulate of the preservationist conservation philosophy — that local human communities cause land degradation and biodiversity loss — is increasingly being questioned for its scientific validity. That this postulate may not hold in many cases is being used, inter alia , in support of calls for more inclusive conservation policies in developing countries. Such policies would allow, or even encourage, consumptive human use of natural resources within designated wildlife-protected areas. However, the latter approach again rests upon the assumption that local human communities and their impacts on natural resources are constant. The present paper questions this assumption using a case study from a hitherto isolated region of the Indian Trans-Himalaya. I describe the ongoing socio-economic flux in an agropastoral Buddhist community dependent upon the resources of a protected area, and the impacts of this transition on wildlife conservation. The analysis shows radical changes in the local economy and land use in the last decade, that ultimately proceed from extrinsic factors (market forces, changes in Government policy). Immediate conservation problems have proximately arisen from both extrinsic (uncontrolled tourism) as well as intrinsic (escalation of livestock stocking rate) changes. The analysis underscores the need for conservation policies to be sensitive to the transient nature of local human communities, even in seemingly isolated protected areas. Introduction The thrust of India’s conservation policy has been preservationist, wherein emphasis has been placed on minimising or eliminating consumptive human uses within areas designated for protection of wildlife. Despite such an exclusionary official policy, more than 80 % of Indian wildlife reserves are inhabited by local human communities that continue to use the natural resources in them, albeit within state-imposed restrictions (Kothari et al. 1989). Such restrictions on traditional resource use following the creation of protected areas are responsible for local hostility and the absence of local support for conservation efforts (Kothari et al 1995, Guha 1997, Saberwal 1997). This 'Accepted June, 1998 ^Centre for Ecological Research and Conservation, 3076/5, IV, Cross Gokulam Park, Mysore 570002, Karnataka, India. 3 Present address: Tropical Nature Conservation and Vertebrate Ecology Group, Wageningen University, 69 Bomsesteeg, 6708 PD Wageningen, The Netherlands. hostility gets further aggravated in the face of serious human-wildlife conflicts in many protected areas, and the subsequent bureaucratic apathy faced by the local people (Guha 1997, Mishra 1997a, Saberwal 1997, Saberwal et al. 1994) Not surprisingly then, as in many other developing countries (Prins 1992), the merits of the Indian preservationist approach are being increasingly questioned on social, economic, ethical, political, pragmatic and even ecological grounds. Critics have contended that the preservationist policy has been based on scientifically unsubstantiated assumptions that local human communities cause land degradation and the loss of biodiversity (Saberwal 1996, Guha 1997). There is an increasing call for 'rethinking conservation’ and embracing a more inclusive policy, which, in theory, allows for biodiversity conservation alongside local human resource use (e.g. Kothari et al. 1995, Saberwal 1996). However, the latter thesis again rests upon an important yet unsubstantiated assumption that views local human communities, their life-styles, JOURNAL , BOMBAY NATURAL HISTORY SOCIETY, 97(1), APR. 2000 25 SOCIOECONOMIC TRANSITION AND WILDLIFE CONSERVA TION and the magnitude of their impacts, as static and immune to change (Mishra and Rawat 1998). It is this assumption that is questioned here. The aim is neither to denounce nor advocate the demands for ‘democratic’ multiple use policies; under the complex sociopolitical situations in most developing countries, strict adherence to either stand will prove counter-productive for wildlife conservation. The purpose of this paper, instead, is to show that irrespective of the official conservation policy ( 1 ) local human communities even in the remotest regions of the develop- ing world are undergoing rapid social and land use transition, (2) this transition has potentially important consequences for wildlife conserva- tion, and following from this, (3) conservation policies need to be extremely sensitive to these changes. Focusing on three agropastoral Buddhist villages (80 households) dependent upon the resources of a protected area, this paper describes the ongoing socio-economic transition in the Spiti region (31° 42' to 32° 58' N lat. and 77° 21' to 78° 35' E long.) of the Indian Trans-Himalaya. Located close to the politically sensitive Sino-Indian border, in difficult mountainous terrain, Spiti remained a remote area with restricted geographical as well as administrative access until 1992. In this paper, I specifically document the socio-economic trends in the region over the last 25 years, and subsequently discuss their consequences for wildlife conservation. The urgent research and management inputs required for conservation both at the local and regional levels are also outlined. Study Area The Trans-Himalayan region includes the high altitude plateau of Tibet and the Tibetan marginal mountains, an area of over 2.6 million km2. The c. 186,000 km2 within India, despite its conservation significance, forms one of the least represented biogeographic zones in the Indian protected area network (Rodgers and Panwar 1988). The Spiti region in the Trans-Himalayan Lahaul and Spiti dist. (Himachal Pradesh) spans an area of 12,210 km2 in the catchment area of the Spiti river, with a human population of 9,59 1 (in 1991; Directorate of Economics and Statistics 1996) which is largely Buddhist (Kaushik 1993). Spiti had no wildlife reserves until the last decade. The establishment of the 675 km2 Pin Valley National Park (31° 44' to 32° IT N lat., and 77° 45' to 78° 06' E long.) in 1987, and the 1400 km2 Kibber Wildlife Sanctuary (32° 5' to 32° 30' N lat. and 78° 1' to 78° 32' E long.) in 1992, has resulted in 17% of Spiti’s land area being designated as wildlife reserve. The protected area boundaries, however, are only nominal, considering they were drawn around existing settlements and villages whose inhabitants continue using these areas for grazing, fuel and fodder collection. Kibber Wildlife Sanctuary lies in the northern catchment of Spiti and is flanked by Ladakh to the north and Tibet to the east. The Sanctuary, like the rest of the Trans-Himalaya, lies in the rain shadow of the Greater Himalaya, and ranges in altitude from c. 3,600 m to 6,700 m above msl. Temperatures range between -30°C to 3°C in the winter, and between 1°C to 28°C in summer (Rana 1994). Vegetation in the area has been broadly classified as dry alpine steppe (Champion and Seth 1968). The Sanctuary is flanked by 13 villages along its southern boundary inhabited by an agropastoral Buddhist community, whose agricultural activities are restricted to the short growing season between May and September. Barley Hordeum vulgare and green pea Pisum sativum are the main crops. Livestock includes goat, sheep, cattle, yak, dzomo (female hybrid of cattle and yak), donkey and horse. Goat, cattle and dzomo are used for both milk and meat. Sheep are used for wool and yaks for ploughing, in 26 JOURNAL. BOMBAY NATURAL HISTORY SOCIETY. 97(1). APR. 2000 SOCIOECONOMIC TRANSITION AND WILDLIFE CONSER VA TION addition to meat. Donkeys are used as draught animals, and raised partly for trade. Horses, apart from being used for religious ceremonies, are raised mainly for trade (Mishra 1997a). The mammalian fauna of the Sanctuary includes snow leopard Uncia uncia , wolf Canis lupus , red fox Vulpes vulpes, pale weasel Mustela altaica, stone marten Mcirtes foina , Himalayan mouse hare Ochotona sp., bharal Pseudois nayaur , and ibex Capra ibex. Methods Unpublished archival records of the State Government were scrutinised (see Mishra 1997a for details of sources) for information relating to human population and past literacy rates, livestock population, and developmental changes in the region over the last 25 years. Of the 13 villages surrounding Kibber Wildlife Sanctuary, three, which together comprised 1 9% of the population living around the park, were selected as samples for the study (for details see Mishra 1997a). This included Kibber, the largest in the area (316 inhabitants), and two small villages nearby, Gete (36) and Tashigang (24). Structured interviews were conducted with at least one member from each household in the three villages, to obtain information regarding present family size and literacy, livestock and land holdings, and past and current agricultural practices. Human and livestock population growth rates (r) were calculated using the exponential growth curve equation (Nt = N0ert where Nt is the population at time t, N0 is the starting population, and e the base of natural logarithms). Crop yield per unit area was obtained for different crops by interviewing two experienced farmers, and the lower limit of the reported range used to obtain a conser- vative estimate of crop production. Casual interviews and observations during the course of field work yielded information on tourism and its impacts. Results Human population and development The human population in the thirteen villages bordering Kibber Wildlife Sanctuary increased only marginally (at an annual growth rate of 0.09%) between 1971 and 1991 (1985 people in 1991; data for 1996 not available). Likewise, between 1971 and 1996, the three study villages saw a total population increase of only 6.5%, an average annual growth rate of 0.25% (Mishra 1997a). Children <18 years comprise 49% of the present population of the study villages. Literacy rate has doubled (from 22% to 48%) in the last 25 years. Presently, 31% of the adult males (n = 91), and 26% of the adult females (n = 100) are literate. In the school-going age group (c. 5 to 1 8 years), there is 97% literacy (n = 127). Among other indicators of development, this period has seen an increase in the number of schools and the electrification of all three study villages (Table 1). Two of the three villages, Gete and Tashigang, which earlier had no roads, have been connected by motorable roads. Agriculture The number of people per unit of irrigated land has remained nearly constant over the last 25 years (Table 1), with the current average land holding per household at 1.13 ha. The cropping pattern, however, has changed in the last decade. Prior to 1986, agriculture was for subsistence. Table 1 PATTERNS IN SOME INDICATORS OF DEVELOPMENT OVER THE LAST 25 YEARS IN THREE SAMPLED VILLAGES OF KIBBER WILDLIFE SANCTUARY Indicator 1 971 1996 No. of medical care centres 1 1 No. of post offices 1 1 No. of schools 1 4 Irri gated land (ha) 83 91* People per ha irri gated land 4.2 4.1 No. of villages with electricity 0 3 No. of villages connected by motorable road 1 3 *in 1987 JOURNAL, BOMBAY NATURAL HISTORY SOCIETY, 97(1), APR. 2000 27 SOCIOECONOMIC TRANSITION AND WILDLIFE CONSERVA TION The main crops were barley and a local variety of pea (the latter largely for supplementing livestock feed), cultivated on two-thirds and one-third of the land holding respectively. Since 1986, however, one-third of the land holding is cultivated for green peas, one-third for barley, and the remaining is partly planted with local pea and partly left fallow. The entire harvest of green pea is sold as a cash crop. The estimated annual production of green pea per household is 2,587 kg, which translates into a per capita profit (corrected for transport costs) of US$ 210 per year (1994-95 conversion rate of 1 US$ = 31.4 Indian Rupees; World Bank 1996). The estimated annual production of barley per household is currently 1,294 kg. This change in cropping pattern has significantly affected an age-old barter trade between the inhabitants of the study area and a semi-nomadic pastoral community, the Changpa of Ladakh. Changpa herders have been coming into Spiti for at least a few centuries (Kapadia 1996). They come in summer with their livestock (> 1,000 goat and sheep) when the high mountain passes ( c . 5,600 m) become negotiable. The main trade involved barley, which earlier was in surplus, and was bartered with the Changpa largely in exchange for wool, salt and rugs. Owing to the replacement of barley with the commercially valuable green pea, and the resulting absence of surplus barley, the development of a market economy, and the improvement in transportation, communication, and supplies in Spiti, this trade is on the verge of breakdown. However, the trade continues for Spiti horses and donkeys, which are still in demand with the Changpa. Livestock The annual growth rate of livestock holdings in the study villages increased from 2.6% (between 1971 and 1987) to 3.5% after 1987 (up to 1996; Mishra 1997a). The growth rate of livestock throughout Spiti after 1987 was 3.2% (10,458 heads in 1988 to 1 1,881 heads in 1992). In the last 25 years, the ratio of livestock to human population in the study villages has increased from 1.85 (the year 1971, in Gete and Tashigang) to 2.80 (1996, in all three villages). In terms of herd composition, Gete and Tashigang (data for Kibber for the year 1 97 1 were not available) show an increase in all livestock species between 1971 and 1987, though the maximum increase was accounted for by goat and sheep (42%). After 1987, the number of donkeys and cow Idzomo declined, while the other species continued to increase (Table 2). In Kibber, the trend after 1987 was almost the same with all the species except cow Idzomo continuing to increase. Thus, in the last ten years, the population of cow Idzomo in all the sampled Table 2 LIVESTOCK POPULATION TRENDS OVER 25 YEARS IN THREE SAMPLED VILLAGES (DATA POOLED FOR THE VILLAGES GETE AND TASHIGANG) OF KIBBER WILDLIFE SANCTUARY Species 1971 Gete and Tashigang * 1987 1996 Kibber village* 1987 1996 Yak 9 14 29 28 110 Cattle/ dzomo 13 32 28 113 98 Horse 6 11 18 34 57 Donkey 11 17 11 93 114 Sheep/ goat 76 101 137 322 452 Total 115 175 223 590 831 *data for 1 971 were not available villages has declined marginally, while yak has increased more than threefold (Table 2). Goat and sheep again accounted for the maximum increase (57 %) during this period. Tourism Prior to 1992, foreign nationals were not allowed in Spiti, and even non-domicile Indians needed to obtain special permits from the State Government to enter the region. With the relaxation of Government policy since 1992, there has been a sudden growth in tourism. 28 JOURNAL, BOMBAY NATURAL HISTORY SOCIETY, 97(1), APR. 2000 SOCIOECONOMIC TRANSITION AND WILDLIFE CONSERVA TION Kibber, one of the study villages, had three functional hotels and one more under construction when this study was conducted, as opposed to none before 1993. These small hotels (3-4 rooms), catering to both Indian and foreign tourists, are run by local villagers. Many villages of Spiti now have makeshift hotels. The tourist inflow is restricted to between June and September. Between June and August 1996, a hotel owner reported a net profit of c. US$ 637 (Chering Dorje, Kibber, pers. comm. 1996). Demand for local guides and donkeys by trekking tourists also causes a substantial inflow of money at the local level, which could not, however, be quantified. Discussion Human population Most habitat change and biodiversity loss in developing countries has been attributed to socio-economic change in growing rural populations (Machlis 1992). The Indian population, 74% of which is rural, has indeed grown at an annual rate of 2. 1 7% in the past two decades, yielding a current density of close to 300 per sq. km (Repetto 1994). In contrast, the absolute human density of Spiti is very low (0.78 per sq. km). The unusual absence of population growth could largely be a consequence of the relatively intact system of primogenitary inheritance over most of Spiti (and polyandry in one region) where the younger siblings become celibate monks (Mamgain 1975, Punjab Government 1994). The stable population size seems to have stabilised the pressure for fuelwood on the protected area. However, it is important to keep in mind that most of the area in Spiti is uninhabited due to its inhospitable cold desert mountainous environment. Consequently, 31% of Spiti’ s present population is concentrated in and around the two protected areas, and is dependent on them for grazing and fuelwood (Pin Valley has a human population of 1500 inside and around the National Park area; Mishra 1997b). A study estimates an annual per capita extraction of 2 17 kg of shrubs and dung (for fuel), and fodder (for winter supplemental feeding) by the resident population from Pin Valley (Bhatnagar 1996). It is also prudent to note that, faced with modernization, other trans-Himalayan Buddhist communities are undergoing rapid population growth following a breakdown of social population regulation mechanisms, and this might happen in Spiti as well (Goldstein 1 98 1 , Fox et al. 1 994, Mishra and Humbert-Droz 1998). Changes in agriculture and animal husbandry The most significant socio-economic change in the region during the last decade has been the shift from a barter-based sub- sistence economy, to a market economy, resulting from, inter alia , changes in cropping pattern. The return per household from green pea harvest, the new cash crop, is almost as high as the average annual per capita income for Himachal Pradesh (US$ 248, 1994-95; World Bank 1996). Along with agriculture, there is indication of commercialisation of animal husbandry as well (livestock trade was earlier restricted to barter with the Changpa). This is evidenced in the three-fold increase of yaks in the last decade, which are now partly being raised in the villages of Kibber Wildlife Sanctuary for selling in other areas of Spiti (Chhewang D. Zangpo, Pin Valley, pers. comm. 1996). This contrasts with other yak rearing communities in the Himalaya, where the yak population is known to be declining rapidly (Negi and Gadgil 1997, J.L. Fox pers. comm. 1996.). Between 1988 and 1992, the yak population of Spiti increased from .786 to 897 heads. Livestock of the study villages graze in the Sanctuary area nearly throughout the year, though their diet is supplemented by stall feeding in winter. This supplemental forage is partly JOURNAL, BOMBAY NATURAL HISTORY SOCIETY, 97(1), APR. 2000 29 SOCIOECONOMIC TRANSITION AND WILDLIFE CONSERVA TION collected during the growing season from the Sanctuary area, and partly from the cropfields. In addition, the State Government has initiated a scheme to provide supplemental feed at subsidised rates. Given the present trend and the augmented ability to purchase supplemental feed, livestock holdings are likely to continue growing in the near future. The increasing livestock stocking rate seems to be intensifying the pressure on the protected area resources for fodder. Escalating livestock stocking rate is a countrywide phenomenon in India, the last two censuses indicating a 1.2% annual growth rate (419 million in 1982 to 445 million in 1987). With 67% wildlife sanctuaries and 83% national parks subject to livestock grazing (Kothari et al. , 1989), the urgency for evaluating the impacts of livestock on wildlife resources is obvious. In Kibber, the increase in stocking rate (together with poor anti-predatory livestock management) seems to be the main reason behind the recent escalation in instances of livestock depredation by large carnivores (the snow leopard and the wolf; Mishra 1997a). Even now, livestock outnumber bharal, the dominant wild ungulate and natural prey of the wild carnivores, by an order of magnitude. To reduce this depredation, villagers have been killing the wolf, and elsewhere, I have expressed concern that persecution of the snow leopard is likely to begin unless specific research and management measures are undertaken to understand and reduce this conflict (Mishra 1997a). At a broader level, there is a need for assessing the impact of grazing on plant communities and evaluating the forage relations between livestock and wild herbivores. The potential for regulating livestock stocking rates and range use to enhance conservation objectives has long been recognised (e.g., Anderson and Scherzinger 1975, Willms etal. 1980), and such studies are a pre-requisite to designing effective multiple-use management policies for Indian protected areas. Uncontrolled tourism Uncontrolled tourism in wildlife reserves has usually resulted in conservation problems (Budowski 1976, deGroot 1983, Kenchington 1989). Kibber presently lacks even a record of the number of tourists visiting the Sanctuary. With the sudden development of tourism, the age-old trade route between Kibber and Ladakh (used by the Changpa ; c. 125 km) has now become a popular trekking route. This route passes along wetlands in Ladakh that are important breeding sites for water birds, including rare and threatened species (Mishra and Humbert-Droz 1998). A rather conspicuous impact of this tourism has been the pollution of this route with discarded garbage (including non-degradable metal cans and polythene), especially around about 15 camping sites. In addition, Kibber Wildlife Sanctuary, like some other regions of Spiti, has deposits of nautiloid, balamnite, and ammonite fossils (Y.V. Bhatnagar, pers. comm. 1997). Locals reported that fossils were being removed from the area even before Spiti was opened to tourists. However, this was confined to geologists and amateur collectors. Tourism has now created a market for fossils, which is causing a rapid depletion of the fossil reserves of Kibber Wildlife Sanctuary and elsewhere in Spiti. Depending upon its size and quality, a fossil may fetch US$ 3 to USS 15. I could not, however, assess the magnitude of this trade. The need for a culturally and ecologically well designed tourism plan for Spiti is apparent, and has already been expressed (Kaushik 1993, 1994). Conclusions Spiti remained geographically as well as politically remote and isolated until 1992, and the so far intact social population regulation mechanisms have kept the local human population under control. However, a rapid socio-economic transition is in progress, 30 JOURNAL, BOMBAY NATURAL HISTORY SOCIETY, 97(1), APR. 2000 SOCIOECONOMIC TRANSITION AND WILDLIFE CONSERVA TION exemplified by improvements in transporta- tion, increase in literacy, changes in cropping pattern (the adoption of a cash crop), breakdown of barter trade, expansion of livestock holdings, and a sudden development of an unplanned tourism industry. This is ultimately driven by far-reaching extrinsic factors such as the influence of commercial markets and changes in Government policy. The transition from a subsistence (barter-based) economy to a market economy, and changes in land use in Kibber Wildlife Sanctuary, have resulted in conservation problems such as the escalation of human- wildlife conflict (livestock depredation by wild carnivores), increased pressure on the protected area for fodder, pollution, and the depletion of fossil reserves. These have proximately been brought about by intrinsic (escalating livestock stocking rates) as well as extrinsic (tourism) factors. This paper joins a growing body of literature documenting the significant influence of market forces even in relatively remote regions of the developing world (e.g. Goldstein 1981, Goldstein and Beall 1989, Fox et al. 1994, Negi and Gadgil 1997, Mishra and Humbert-Droz 1998). It further shows that the resultant Refer Anderson, E.W. & R.J. Scherzinger (1975): Improving quality of winter forage for elk by cattle grazing. J. Range Manage. 28\ 120-125. Bhatnagar. K. ( 1 996): A study on people’s dependence, attitudes and ecodevelopment in Pin Valley National Park, Himachal Pradesh. Unpublished report. Worldwide Fund for Nature (India), New Delhi, India. Budowski, G. (1976): Tourism and environmental conservation: conflict, coexistence, or symbiosis? Enviro. Conserv. 3: 27-31. Champion, H.G. & S.K. Seth ( 1 968): A Revised Survey of the Forest Types of India. Manager of Publications, Delhi, India, pp 404. deGroot, R.S. (1983): Tourism and conservation in the Galapagos islands. Biol. Conserv. 26: 291-300 Directorate of Economics and Statistics (1996): transition in socio-economy and landuse in local human communities can result in complex conservation problems. Conservation policies therefore, ought to bear in mind the transient nature of local human communities residing even in seemingly remote protected areas. Acknowledgements I thank Dr. K. Ullas Karanth, the Wildlife Conservation Society, Bronx, NY, and the Netherlands Foundation for the Advancement of Tropical Research (WOTRO), a body residing under the Netherlands Organization for Scientific Research (NWO) for financial support. My gratitude for technical support to the Department of Forest Farming and Conservation, Himachal Pradesh, and the Director, Wildlife Institute of India. I thank Y.V. Bhatnagar, K. Bhatnagar, T. Dorje, Dr. J.L. Fox, L. Gyalson, N. Manjrekar, and B.S. Rana, for discussions; Dr. S.P. Goyal and Dr. A.J.T. Johnsingh for encouragement, and Dr. R.S. Chundawat, Dr. S.N. Mishra, Madhusudan Katti, and T.R.S. Raman for comments. The contribution of M.D. Madhusudan in restructuring the paper is gratefully acknowledged. ENCES Economic review. Directorate of Economics and Statistics, Himachal Pradesh Government. Shimla, India. Fox, J.L., C. Nurbu, S. Bhatt, & A. C'handola (1()94): Wildlife conservation and land-use changes in the trans-Himalayan region of Ladakh, India. Mount. Res. Develop. 14: 39-60. Goldstein, M.C. (1981): High altitude Tibetan populations in the remote Himalaya: social transformation and its demographic, economic, and ecological consequences. Mount. Res. Develop. I: 5-18. Goldstein, M.C. &C.M. Beall (1989): Nomads of western Tibet: survival of a way of life. Serindia Publications. London, 191 pp. Guha, R. (1997): The authoritarian biologist and the arrogance of anti-humanism: wildlife conservation in the third world. The Ecologist 27: 14-20 JOURNAL . BOMBAY NATURAL HISTORY SOCIETY. 97(1). APR. 2000 31 SOCIOECONOMIC TRANSITION AND WILDLIFE CONSERVA TION Kapadia, H. (1996): Spiti: adventures in the trans-Himalaya. Indus Publishing Company, New Delhi, India. Kaushik, S. (1993): Towards a tourism strategy in Spiti. Equitable Tourism Options, Bangalore, India. Kaushik. S. (1994): The blossoming of an affair: a Spiti update. Equitable Tourism Options, Bangalore, India. Kenchington, R.A. (1989): Tourism in Galapagos Islands: the dilemma of conservation. Environ. Conserv. 16: 227-232. Kothari, A., P. Pande, S. Singh & D. Variava (1989): Management of National Parks and Wildlife Sanctuaries in India: a status report. Indian Institute of Public Administration, New Delhi, India, 298 pp. Kothari, A., S. Suri & N. Singh (1995): People and protected areas: rethinking conservation in India. The Ecologist 25: 188-195. Machlis, G.E. (1992): The contribution of sociology to biodiversity research and management. Biol. Conserv. 62: 161-170. Mamgain. M.D. (1975): Himachal Pradesh district gazetteers: Lahaul and Spiti. Greater Punjab Press, Chandigarh, India. Mishra, C. (1997a): Livestock depredation by large carnivores in the Indian Trans-Himalaya: conflict perceptions and conservation prospects. Environ. Conserv. 24: 338-343. Mishra, C. (1997b): Livestock grazing and wildlife conservation in the Indian Trans-Himalaya: a preliminary survey. Unpublished report for the Wildlife Conservation Society, Bronx. NY. Centre for Ecological Research and Conservation, Mysore, India, 22 pp. Mishra, C. and B. Humbert-Droz ( 1 998): Avifauna! survey of Tsomoriri Lake and adjoining Nuro Sumdo wetland in Ladakh, Indian trans-Himalaya. Forktail 14: 865-867. Mishra, C. & G.S. Rawat ( 1 998): Livestock grazing and biodiversity conservation: comments on Saberwal. Conserv. Biol. 12: 1 1 2-7 1 4. Negi, H.R. & M. Gadgil (1997): Conserving livestock genetic resources: a case study of Kinnaur in Himachal Pradesh. .1. Human Ecol. 6 (Special issue): 317-324. Prins, H.H.T. (1992): The pastoral road to extinction: competition between wildlife and traditional pastoral ism in East Africa. Environ. Conserv. 19: 117-123. Punjab Government (1994): Gazetteer of the Kangra District, parts II to IV- Kulu, Lahaul and Spiti, 1897. Indus Publishing Company, New Delhi, India. Rana, B.S. ( 1 994): Management plan of Kibber Wildlife Sanctuary. Department of Forest Farming and Conservation, Wildlife Wing, Shimla. Himachal Pradesh, 81 pp. Repetto, R. (1994): The ‘Second India’ revisited: population, poverty, and environmental stress over two decades. World Resources Institute, Washington D.C., USA. Rodgers, W.A. & H.S. Panwar (1988): Planning a protected area network in India. Vol. I - the Report. Wildlife Institute of India, Dehra Dun. India, 341 pp. Saberwal, V.K. ( 1996): Pastoral politics: gaddi grazing, degradation and biodiversity conservation in Himachal Pradesh, India. Conserv. Biol 10: 741-749. Saberwal, V.K. ( 1 997): Saving the tiger: more money or less power? Conserv. Biol. 1 1: 8 1 5-8 1 7. Saberwal, V.K., J.P. Gibbs, R. Chellam & A.J.T. Johnsingh (1994): Lion-human conflict in Gir Forest, India. Conserv. Biol. 8: 501-507. Willms, W., A.W. Bailey & A. McLean (1980): Effect of burning or clipping Agropyron spicatum in autumn on the spring foraging behavior of mule deer and cattle. J. Appl. Ecol. 17: 69-84. World Bank (1996): India: country economic memorandum. World Bank Report 1 58882 IN . World Bank. Washington, DC, USA. 32 JOURNAL, BOMBAY NATURAL HISTORY SOCIETY, 97(1). APR. 2000 AN ECOLOGICAL STUDY OF CROCODILES IN RUHUNA NATIONAL PARK, SRI LANKA1 Charles Santiapillai, Mangala de Silva2, Sarath Dissanayake3, B. V.R. Jayaratne4 and S. Wijeyamohan5 ( With three text-figures) Key words: Marsh crocodile, Crocodylus palustris, estuarine crocodile, C. porosus , Ruhuna National Park, basking, feeding, conservation A study was carried out in Block I (140 sq. km) of the Ruhuna National Park (RNP) opportunistically from October 1 99 1' to October 1 994, in order to study the two species of crocodiles occurring in Sri Lanka, viz. Crocodylus palustris and C. porosus. A total of 341 sightings of the two species were made on 77 occasions, 307 sightings on C. palustris and 34 sightings on C. porosus. Among C. palustris , solitary animals made up most of the observations (55.8%) while pairs accounted for 13.0%. Of the 22 water-holes that were surveyed, 13 (59%) had only one crocodile. Although both species could be seen at any time of the day, the number basking increased with the increase in the ambient temperature, and peaked around noon. C. porosus basked alone, and C. palustris communally. The population structure consisted of 44% hatchlings, 6% juveniles, 24% subadults and 26% adults. Only adults of C. porosus were observed. Hatchling losses can be very high through predation by birds and mammals. Both species feed on a variety of food, ranging in size from aquatic insects and Crustacea (in hatchlings) to fish, frogs, birds and large mammals (in adults). The minimum crude density values for C. palustris and C. porosus are estimated to be 0.72 and 0.07 animals per sq. km respectively. The populations of both species in Block I appear to be secure and viable. Introduction Of the 13 species of ‘true’ crocodiles (Subfamily: Crocodylinae) that are extant in the world, 8 species occur in Asia, of which 2 are found in Sri Lanka, namely the freshwater, or marsh crocodile, or mugger ( Crocodylus palustris) and the saltwater or estuarine crocodile (C. porosus). While C. palustris is listed as ‘vulnerable’ by IUCN (Groombridge, 1993), C. porosus has been transferred to the Tow risk’ category, given the tens of thousands known to ‘Accepted April, 1999 department ofZoology, University of Peradeniya, Sri Lanka ?National Wildlife Training Centre, Giritale, Sri Lanka JWasgomuwa National Park, Hasalaka, Sri Lanka 'Faculty of Applied Sciences, Vavuniya Campus of the Jaffna University, Vavuniya, Sri Lanka be present in numerous localities across its geographical range. However, in Sri Lanka, given its low number and restricted distribution, C. porosus is more threatened than C. palustris. According to Whitaker and Whitaker (1989), “Sri Lanka has more mugger crocodiles than the rest of the subcontinent put together, mostly concentrated in the two national parks, Yala (=RNP) and Wilpattu.” Even though this may not be strictly true now, it indicates the high number of mugger crocodiles still occurring in Sri Lanka. Both species found in Sri Lanka are listed in Appendix I of the Convention on International Trade in Endangered Species of Wild Fauna and Flora (CITES). Crocodiles were once plentiful in Sri Lanka. The man-made reservoirs or tanks in the Dry Zone were teeming with crocodiles (Baker JOURNAL . BOMBAY NATURAL HISTORY SOCIETY. 97(1). APR. 2000 33 ECOLOGICAL STUDY OF CROCODILES IN RUHUNA NATIONAL PARK 1853; Tennent 1859; Hennessey 1949). But today, both species have declined in range and number as a result of poaching and loss of habitat. Crocodiles are almost confined to the first peneplain in Sri Lanka. They represent an excellent renewable natural resource and, therefore, their conservation can be made much easier if such a resource is used for the benefit of the people who share the land with them (Child, 1987). Unfortunately, crocodiles have a poor image in Sri Lanka. They are considered dangerous, and few would really regret their disappearance. The general public is unaware of the beneficial role played by crocodiles in wetlands. Legislation alone cannot save a species if the public is against its conservation. As Sale (1985) points out, a sound scientific understanding of a natural resource is fundamental to the management of that resource. Nowhere is this more true than in Sri Lanka, where the aims of crocodile management are straightforward preservation of the species within protected areas, with no interest in utilization despite the high economic value of the skin. So far, there has been no ecological study of crocodiles in Sri Lanka. Deraniyagala (1953) provides detailed information on the taxonomy, range and ecology of the two species of crocodile in Sri Lanka, while Whitaker and Whitaker (1979) carried out the first comprehensive survey of crocodiles in Sri Lanka. More recently, Porej (1997) studied the distribution of the two species along the south-western coast of Sri Lanka. An island-wide reassessment of their status was carried out by Santiapillai & de Silva (1998, under review). Study Area The study was carried out in Block I of the Ruhuna National Park, in southeast Sri Lanka in the low country Dry Zone (Fig.l). Block I is about 140 sq. km in extent, and is separated from the rest of the park by the Menik Ganga (= river) in the northeast. The vegetation of the park has been classified by Mueller-Dombois (1972) into three physiognomic categories: (a) forest (with at least 20% of crown biomass above 5m in height), (b) scrub (less than 20% of crown biomass above 5m), and (c) grassland or plains. The dominant forest trees are Manilkara hexandra (palu), Drypetes sepiaria (weera) in well drained soil, and Limonia acidissima (divul) and Salvadora persica (malithan) in poorly drained areas (Balasubramaniam et «/., 1980). The coastal region in Block I has numerous water-holes of varying size and salinity, surrounded by grasslands where the main species are Eragrostis viscosa, Dactyl otaenium aegyptium , Sporobolus diandrus , Echinochloa colonum, Setaria pallidifusca and Alloteropsis cimicina (Balasubramaniam et al. , 1980). The fauna includes threatened species such as the Asian elephant Elephas maximus (E), leopard Panthera pardus (T), sloth bear Ursus ursinus (I), and water buffalo Bubalus buba'lis (V). In addition, there are several herbivores: wild pig Sus scrofa , sambar Cervus unicolor , spotted deer Axis axis and mouse deer Tragulus meminna , which are potential prey species of the crocodiles. Other reptiles include the common monitor lizard Varanus bengalensis, cobra Naja naja, Russell’s viper Daboia russelli. At least three species of sea turtles, the green Chelonia mydas (E), olive Ridley Lepidochelys olivacea (E) and leatherback Dermochelys coriacea (E), nest along the beach (Hewavisenthi, 1990). The most numerous crocodile in Ruhuna National Park is the marsh crocodile or mugger (C. palustris). Methods The study on crocodiles was incidental to a much larger study on the mammals of the Ruhuna National Park and was carried out in Block I opportunistically from October 1991 to October 1994. All observations were made from a vehicle, using a pair of 7 x 52 binoculars, from 34 JOURNAL, BOMBAY NATURAL HISTORY SOCIETY, 97(1), APR. 2000 ECOLOGICAL STUDY OF CROCODILES IN RUHUNA NATIONAL PARK Menik Pannagamuwak~-o Fig. 1 : Map of Block I of Ruhuna National Park showing the waterholes 0600 to 1 900 hr, during which time most of the water-holes in the park were visited. At every sighting of crocodiles, their number, location, habitat and behaviour were noted. Whenever possible, the species was identified based on field criteria such as the shape of the dorsal osteoderms — subquadrangular plates transversely sutured to one another in C. palustris , and ovoid and separated by skin in C. porosus (Deraniyagala, 1953). But this was not easy, for as Daniel (1983) points out, the two species are difficult to distinguish in the field. When the two species JOURNAL BOMBAY NATURAL HISTORY SOCIETY, 97(1), APR. 2000 35 ECOLOGICAL STUDY OF CROCODILES IN RUHUNA NA TIONAL PARK are in water, they are almost impossible to tell apart. Besides, smaller individuals are difficult to distinguish in the field. Wherever possible, the length of the animals was estimated visually. Four categories were recognized: hatchlings (<0.5 m), juveniles (0. 5-1.0 m), subadults (1. 1-2.0 m), and adults (>2 m). The crocodiles were also monitored from 0600 to 1900 hrs at Buttuwa Wewa during the peak of the dry season in early October 1991, just prior to the northeast monsoon rains, to study their basking behaviour. An attempt was made to estimate the minimum number and density of crocodiles by taking into account the maximum number recorded from each waterhole within a sampling session (7-10 days). Results A total of 341 crocodiles (of both species) were recorded in 77 observations, of which 307 sightings were on C. pcilustris and 34 on C. porosus. Among C. porosus, solitary animals made up 55.8%, while pairs accounted for 13.0% (Fig. 2). The largest group seen during the survey consisted of 44 animals (39, C. pcilustris and 5, C. porosus ), in the Buttuwa reservoir. It is likely that many of the pairs observed in Buttuwa reservoir are adult male and female marsh crocodiles. Of the 22 water-holes that were surveyed, 13 (59%) had only one crocodile (C. pcilustris) each. Crocodiles were observed to move from one waterhole to another during the dry season. As the dry season progresses from May to September, many of the smaller water- holes become bone dry, and the crocodiles (C. pcilustris ), move either to large water-holes such as the Buttuwa Wewa, Wilapala Wewa, Keen Wewa and Katagamuwa tank, or concentrate along the Menik Ganga. In the dry season, one crocodile (C. pcilustris) was observed more than a kilometre from the nearest water- hole in the neighbouring Block II. At the peak of the drought, marsh crocodile numbers along % of observations 36 JOURNAL BOMBAY NATURAL HISTORY SOCIETY, 97(1), APR. 2000 ECOLOGICAL STUDY OF CROCODILES IN RUHUNA NATIONAL PARK the banks of Menik ganga can be as high as 35 animals per km. Furthermore, if the drought is prolonged, the Menik ganga mostly dries up, leaving scattered pools of water along the banks. These pools, which are no more than 0.5 m in depth, and a few sq. m in area, may be inhabited by up to 4 marsh crocodiles. The largest estuarine crocodile seen measured about 3.0 m at Diganwala, while the largest marsh crocodile was about 2.5 m at Gonalabba lagoon. Table 1 SIZE AND COMPOSITION OF MARSH CROCODILES (C. PALUSTRIS) IN RNF (N = 50) size class (m) number percentage category <0.5 22 44 hatchling 0.5-1 .0 3 6 juvenile 1 .0-2.0 12 24 subadult >2.0 13 26 adult When the maximum number observed in each waterhole within a sampling session (7-10 days) was taken into account, there were 101 marsh crocodiles and 10 estuarine crocodiles in Block I. This amounts to a minimum crude density of 0.72 per sq. km of C. pdlustris , and 0.07 per sq. km of C. porosus in Block I. Among C. palustris, 44% were hatchlings, 6% were juveniles, 24% subadults, while sexually mature animals made up 26% (Table 1). The observed C. porosus were all adults. However, the hatchlings and juveniles taken as C. palustris may have included some C. porosus as well, since these two species are difficult to distinguish in the field from a distance, especially when they are small. Crocodiles could not be sexed in the field. Crocodiles were seen throughout much of the day, either in water, or basking on land. In Block I, both species were observed basking on the embankment of the reservoirs or on the banks of rivers and streams. The pattern of basking observed at Buttuwa Wewa was generally the same in both species (Fig. 3). The ambient temperature increased as the day progressed, and there was a substantial increase in the number number of crocodiles 600 700 800 900 1000 1100 1200 1300 1400 1500 1600 1700 1800 time C. palustris C. porosus Fig. 3: Pattern of basking activity shown by both species of crocodile JOURNAL, BOMBAY NATURAL HISTORY SOCIETY, 97(1), APR. 2000 37 ECOLOGICAL STUDY OF CROCODILES INRUHUNA NA TIONAL PARK of C. palustris observed basking, with the maximum number recorded from 1 1 00 to 1 200 hrs. A much smaller number of C. porosus, while showing a similar trend, were observed basking from 0800 hrs, reaching a peak from 1100 to 1500 hrs, and subsequently declining until 1700 hrs. Another behavioural difference that may help in the identification of species in the field concerns basking. Marsh crocodiles were seen basking communally, while estuarine crocodiles were never observed basking together. However, the estuarine crocodile was also seen basking in the company of marsh crocodiles. While basking, one C. palustris was observed defaecating, after which it moved its hind leg over the pile of faeces and shifted its hind parts a little away, then continued basking. Basking crocodiles varied in the length of time they kept their mouths open, the maximum period being 2 hrs. Both species of crocodile were observed feeding on frogs, which are abundant in almost all the water-holes in Block I. In the dry season, frogs may form a substantial part of the crocodiles’ diet at the smaller water-holes where there are no fish or crustaceans such as crabs or prawns, since the water-holes dry up. However, in the lagoons such as Gonalabba, Uraniya and larger water-holes at Heenwewa, Wilapala Wewa or Palatupana, into which Tilapia were introduced, crocodiles fed largely on such fish. Two marsh crocodiles were seen at night attacking a dead buffalo, in Uraniya plains. Marsh crocodiles were also observed feeding on the carcass of spotted deer, and sambar. In the present study, estuarine crocodiles were not observed feeding on carrion, although it is quite likely that they do. They were not observed doing so, though they were seen at night away from the water-holes. Marsh crocodiles were seen pulling the carcasses either from land or near the water’s edge into water and eating them. Once the carcass is under water, it is out of reach of other scavengers such as jackal ( Canis aureus) and wild pig ( Sus scrofa). In Ruhuna National Park, crocodiles of both species catch most of their terrestrial prey near the edge of the water. Much of the feeding appears to take place at night. Hatchling losses can be very high due to predation. In Block I, hatchlings were seen among the roots of Rhizophora trees in the mangroves at Buttuwa, where the prop-roots form a three dimensional mesh, which even some large wading birds find difficult to penetrate. The only birds large enough to attack hatchlings are the black-necked stork ( Ephippiorhynchus asiaticus), lesser adjutant stork ( Leptoptilos javanicus ), spot-billed or grey pelican ( Pelecanus roseus ), and raptors such as crested hawk eagle (Spizaetus cirrhatus), crested serpent eagle ( Spilornis cheela), brahminy kite (Haliastur indus) and white -bellied sea eagle ( Haliaeetus leucogaster). According to Park officials, egg predation by jackal ( Canis aureus ), monitor lizard (Varanus bengalensis) and wild pig can be substantial. Discussion In addition to the crocodiles that were observed in Block I of RNP, another 150-200 marsh crocodiles were recorded from the Katagamuwa Wewa (Fauna International Trust, 1993; de Silva, pers. obs.), which lies just outside the northwest comer of Block I (Fig. 1). As these marsh crocodiles regularly move in and out of Block I, they could be considered a part of the crocodile population of Block I. If these crocodiles are also taken into account, then the minimum cmde density of the marsh crocodile in Block I could be as high as 1.99-2.16 animals per sq. km. Marsh crocodiles live in groups, but male estuarine crocodiles, being aggressive and highly territorial, tend to live alone. Furthermore, in estuarine crocodiles, the large territorial males may service a number of females, and thus keep potential competitors at bay (Webb and Manolis, 1989). This may explain the movement of some 38 JOURNAL BOMBAY NATURAL HISTORY SOCIETY, 97(1), APR. 2000 ECOLOGICAL STUDY OF CROCODILES IN RU HUN A NA TIONAL PARK males far into the interior, away from the estuaries. The number of crocodiles inhabiting a particular waterhole depends not only on the productivity of the waterhole, but also on its size. Usually, large waterholes such as Wilapala Wewa and Buttuwa Wewa support relatively large numbers of crocodiles, in particular C. palustris, all year round. In general, female crocodiles grow more slowly and reach maturity at a smaller size than males, which continue growing and usually exceed females in maximum size (Ross, 1998). According to Webb and Manolis (1989), in saltwater crocodiles, the females reach sexual maturity at the age of 12 years (2.3 m total length), while the males become sexually mature at the age of 16 years (3.4 m total length). But female marsh crocodiles of 6 years and 8 months of age (2.2 m) have also been known to reach sexual maturity in India (Whitaker and Whitaker, 1989). As crocodiles cannot maintain a constant body temperature by physiological means, heating and cooling are of particulai importance to them (Webb and Manolis, 1989). Crocodilians have a preferred body temperature of about 30-33°C, and to achieve this temperature range, they move to and fro between water and land. Basking crocodiles usually orient themselves in such a way as to get the maximum exposure to the sun. But as their body gets heated, they reduce the heat uptake by turning and facing the sun, and opening their mouth to cool the brain through evaporative cooling (Webb and Manolis, 1989). Crocodiles in general are very sluggish, and their short periods of activity are usually followed by long periods of inactivity. Wading birds were seen feeding quite close to the basking crocodiles. Crocodiles are very effective aquatic predators. They are also opportunistic feeders, and catholic in their diet. Most wild crocodiles are known to be attracted to carrion (Webb and Manolis, 1989). In Katagamuwa tank, marsh crocodiles are known to feed communally on fish, when water is low (Fauna International Trust, 1993). Although game animals fall prey to crocodiles, such predation is unlikely to have a significant effect on their numbers. It is likely that the bulk of the crocodiles’ food in the park consists of fish, frogs and water birds, which are most abundant the year round. As the dry season progresses, many of the water-holes dry up. Fish become concentrated in a few water-holes, which attract crocodiles from other areas. Crocodiles can go for months without feeding (Whitaker and Whitaker, 1989). They are known to feed on a variety of food items that range in size from freshwater mussels to water buffalo (Webb and Manolis, 1989). Their food changes with their size: beginning with aquatic insects, Crustacea, small fish, and as they grow larger, vertebrates such as fish, turtles, birds and mammals (Ross, 1998). Much of the feeding appears nocturnal, for which they are well equipped with good eye sight. The retinal tapetum situated at the back of the eyeball is an image intensifier, allowing crocodiles to see better even in low light intensities (Webb and Manolis, 1989). The predators on crocodile hatchlings, apart from those observed in Block I, include larger crocodiles, freshwater turtles, large predatory fish and python (Webb & Manolis, 1989). Although crocodiles lay many eggs, only 1 % of the hatchlings may survive to maturity, largely due to predation. The estuarine crocodile also suffers heavy losses when flash floods inundate estuaries where its mound nests are found. Given the high number of crocodiles, especially marsh crocodiles, present in Block I of RNP, and the fact that these animals maintain genetic exchange with crocodiles from the rest of the Park, it is clear that both species of crocodile present in Block I constitute secure and viable populations. Factors such as desiccation of eggs during severe drought and avian predation on hatchlings appear to help regulate crocodile numbers in the Park. JOURNAL, BOMBAY NATURAL HISTORY SOCIETY, 97(1), APR. 2000 39 ECOLOGICAL STUDY OF CROCODILES IN RUHUNA NA TIONAL PARK Conservation and Management There has never been any conservation programme designed specifically for crocodiles in Sri Lanka. While, they are being killed as vermin or poached for meat and skin outside the protected areas, their prospects for long-term survival appear good in a few protected areas such as the Ruhuna National Park in the southeast and the Wilpattu National Park in the northwest. The policy of allowing nature to follow its own course appears to have benefited crocodiles within these protected areas. Crocodiles being large predators, require very large areas of undisturbed wetlands to survive (Ross, 1998). Such areas are becoming increasingly difficult to find in Sri Lanka, as a result of the increase of its human population, currently estimated to be over 18 million. Therefore, protected areas appear to be the last refuge for wildlife. There have been no recent reports of crocodiles being poached within the Park, although several were killed outside. The approach to management of crocodiles in the park is therefore a conservative one, in that the crocodile habitats are secure and remote from centres of high human population. So far, management measures have boosted the numbers of the crocodiles inside RNP. The crocodile, being an exceptionally adaptable predator, is able to survive on a broad spectrum of prey species. So the emphasis in crocodile conservation policy Refer Baker, S. W.H. ( 1 853): The Rifle and the Hound in Ceylon. Repr. 1970. Tisara Prakasakyo, Dehiwela. Balasubramaniam, S., C. Santiapillai & M.R. Chambers (1980): Seasonal shifts in the pattern of habitat utilization by the spotted deer ( Axis Erxleben, 1 777) in the Ruhuna National Park, Sri Lanka. Spixiana. 3: 157-166. Bellairs, A. d’A. (1987): The Crocodilia. 5-7. In: (eds.) G.J.W. Webb, S.C. Manolis, & PJ. Whitehead. Wildlife Management: Crocodiles and Alligators. Surrey Beatty & Sons, Chipping Norton, Australia. Child, G. F. T. ( 1 987): The management of crocodiles in must be on maintaining a variety of prey, and preventing the pollution and destruction of the Park’s wetlands. The national parks, however, remote from human population centres, are still prone to environmental disturbances outside their boundaries. The crocodile is well adapted to respond to a “sanctuary strategy”. There are good grounds to believe that it will increase in number under protection, which is by far easier, cheaper and more likely to be successful, than re-introduction. Local people strongly object to the translocation of a potentially dangerous predator such as the crocodile, to their neighbourhood. Law enforcement will become ineffective in the face of public hostility to crocodiles. The dissemination of factual information on crocodiles and their role in the ecosystem may help change the people’s attitude. In the final analysis, the survival of crocodiles is intimately linked with their acceptance by local people and the attitude of their politicians. What is needed is the widest possible acceptance of crocodiles as a renew able natural resource. Their conservation can be made easier, if this resource is used for the benefit of the people who share the land with them (Child, 1987). If crocodiles are properly managed, either in farms or as wild populations, they can become a considerable economic asset to the countries that contain them (Bellairs, 1987). E N C E S Zimbabwe. 49-62. In: (eds.) G.J.W. Webb, S.C. Manolis & P.J. Whitehead. Wildlife Management: Crocodiles and Alligators. Surrey Beatty & Sons, Chipping Norton, Australia. Daniel, J.C. (1983): The Book of Indian Reptiles.Bombay Natural History Society, Bombay. Deraniyagala, P.E.P. ( 1 953): A Coloured Atlas of some Vertebrates from Ceylon. The Ceylon Government Press, Colombo. Fauna International Trust. (1993): Yala National Park. Aitken Spence Printing (Pvt.) Ltd. Colombo. Groombridge, B.(ed.). (1993): 1994 IUCN Red List of 40 JOURNAL . BOMBAY NATURAL HISTORY SOCIETY. 97(1). APR. 2000 ECOLOGICAL STUDY OF CROCODILES IN RUHUNA NA TIONAL PARK Threatened Animals. IUCN Gland, Switzerland. Hennessey. D.J.G. (1949): Green Aisles: A story of the jungles of Ceylon. Colombo Book Centre, Colombo. Hewavisenthi, S. ( 1 990): Exploitation of marine turtles in Sri Lanka: historic background and the present status. Marine Turtle Newsletter 48: 14-19. Mueller-Dombois, D. (1972): Crown distortion and elephant distribution in the woody vegetation of Ruhuna National Park, Ceylon. Ecology. 53: 208-226. Porej, D. (1997): Crocodile Survey and Public Relations Program. Crocodile Specialist Group Newsletter. 16(3): 910. Ross, J.P. (ed.) (1998): Crocodiles. Status Survey and Conservation Action Plan. 2nd edition. 1UCN/SSC Crocodile Specialist Group, IUCN, Gland, Switzerland. Santiapillai, C. & M. de Silva ( 1 998): Status of Crocodiles in Sri Lanka, (under review). Sale, J. B. H (1985): Wildlife Research in the Indomalayan Realm. 1 37-149. In: (ed.) J.W. Thorsell. Conser\>ing Asia’s Natural Heritage: The planning and Management of Protected Areas in the Indomalayan Realm. IUCN Gland, Switzerland. Tennent, Sir. .1 .E. ( 1 859): Ceylon: an account of the island physical, historical and topographical, with notes on its natural history, antiquities and productions. 6th edn. Tisara Prakasakyo, Dehiwela, Sri Lanka. Webb, G. & C. Manolis ( 1 989): Crocodiles of Australia. Reed Books, Australia. Whitaker, R. &Z. Whitaker (1979): Preliminary crocodile survey — Sri Lanka. J. Bombay nat. Hist. Soc. 76: 66-85. Whitaker, R. (1987): The Management ofCrocodihans in India. 63-72. In: (eds.) G.J.W. Webb. S.C. Manolis & P.J. Whitehead. Wildlife Management Crocodiles and Alligators. Surrey Beatty & Sons. Chipping Norton, Australia. Whitaker, R. & Z. Whitaker (1989): Ecology of the Mugger Crocodile. 276-296. In: Crocodiles: their ecology, management, and conservation. IUCN Gland, Switzerland. JOURNAL, BOMBAY NATURAL HISTORY SOCIETY, 97(1), APR. 2000 41 SEXUAL HARASSMENT AMONG FEMALE LION-TAILED MACAQUES {MAC AC A SILENUS) IN THE WILD1 Ajith Kumar2 (With three text-figures) Key words: reproductive suppression, sexual swelling, Maccica silenus , mounting frequency Adult female lion-tailed macaques often harass sexually interacting adult male and female members of the group. The extent of harassment and its implication for reproduction by females was studied in a group in the Anaimalai (presently Indira Gandhi) Wildlife Sanctuary, Tamil Nadu, India. Nearly 1 560 hours of observation were made on the same group during nine months in 1979-80 and 15 months in 1982-84. A total of 577 sexual interactions between single adult male and females were recorded. Most of the sexual mountings occurred when the females had sexual swelling with a peak 2-4 days prior to deflation of the swelling. Most of the harassment was by females with sexual swelling. Harassment decreased the probability of mating taking place once a sexual interaction had been initiated (from 0.582 to 0.07). Aggressive harassment significantly reduced the duration of mating (from 9.12 secs to 6.16 secs), and thus probably prevented ejaculation. The percentage of sexual interactions that were harassed increased with the number of females with sexual swelling. Postponement of conception due to harassment might be a major reason for the absence of a synchrony in conceptions and births similar to that seen in sexual swelling soon after the summer amenorrhea. Sexual harassment is unlikely to serve as a behavioural means of population regulation. This is because fewer females show sexual swelling as the group becomes larger, probably due to increasing competition for food resources. The major reason for the occurrence of sexual harassment in the lion-tailed macaque might be competition among females for mating. This competition results from a high synchrony in sexual swelling among the females, the tendency for groups to have only one adult male, a high female to male (5:1) ratio, and multiple mount pattern in the male. Introduction Reproductive suppression of ovulating females occurs in some primates. In Theropithecus gelada, females actively disrupt each other’s copulation (Mori, 1979). In the same species anovulatory cycles and premature termination of menstrual cycles and implantation occur in low ranking females from social stress due to harassment by high ranking females (Dunbar, 1980). Reproductive suppression from social stress also occurs in Papio cynocephalus (Wasser, 1983). In captivity, female rhesus monkeys could be prevented from mating by 'Accepted June, 1998 :Salim Ali Centre for Ornithology and Natural History Ana ikatti, Coimbatore 641 108, Tamil Nadu, India. aggression from high ranking females (Keveme, 1983). Reproductive suppression of ovulating females has also been demonstrated in captive Miopithecus talapoin (Abbot et al. , 1986). In marmoset monkeys ( Callithrix jacchus) ovulation by subordinate females is physio- logically suppressed by the mere presence of the dominant females (Abbot, 1988). Lion-tailed macaque, Confined to the rain forests of the Western Ghats of South India, mostly live in one male units with a mean group size of 18-20 animals (Kumar, 1995a). The reproductive biology is characterized by a high sex ratio in favour of females ( 1 :5), a conspicuous sexual swelling phase to which compulatory mountings are mostly confined, and a low birth rate (0.30/female year) compared to other macaques (Kumar 1987, 1995a). There is also a 42 JOURNAL, BOMBAY NATURAL HISTORY SOCIETY, 97(1). APR. 2000 SEXUAL HARASSMENT AMONG FEMALE LION- TAILED MA CA QUES high degree of synchrony in the incidence of sexual swelling among the females. Harassment of the mating pair by others, especially by adult females, is frequent. This study examines the extent of harassment of matings pairs and its implication for reproduction by the females. Whether such harassment could play a population regulatory role is also discussed, since birth rate has been found to decrease with increasing group size (Kumar, 1995b). Methods The analysis is based primarily on data collected during an ecological study on one group in the Anaimalai (presently Indira Gandhi) Wildlife Sanctuary, Tamil Nadu State, from March 1979 to March 1980, and from December 1982 to March 1984. The group was located in Varagaliyar shola, about 25 km south of Top Slip, the Sanctuary headquarters. Varagaliyar shola is about 20 sq. km in area and is the largest of the rain forest fragments in the Sanctuary. This shola had five or six groups of lion-tailed macaque. The main study group had only one adult male during both the study periods. There was no Table 1 COMPOSITION OF THE MAIN STUDY GROUP IN THE INDIRA GANDHI WILDLIFE SANCTUARY IN 1979-80 AND 1982-84 Year Adult males Subadult males Adult females Immatures Total Jan 1979 1 0 5 6 12 Mar 1980 1 0 5 9 15 Dec 1982 1 1 6 9 17 Mar 1984 1 1 9 12 23 subadult male in 1979-80, and one in 1982-84. The number of adult females varied from 5 in 1979-80 to 9 in 1982-84 (Table 1). Data on the incidence and duration of sexual cycles come from records on the sexual status (presence or absence of swelling) of females in the study group. These records were made during five to eight days of dawn to dusk observation of the group every month, and at least once in a week during the remaining part of the month. All sexual interactions between the adult male and females were recorded ad libitum during dawn to dusk observation, along with the sexual status of the female. The copulatory calls of the females (see below), given during more than 80% of the sexual mounting and audible up to 75 m, was used as an indicator of mounting. Mounting frequency/hour was estimated for each day by dividing the number of mountings (seen and heard) by the number of hours of observation. Only days with dawn to dusk observation were selected for analyses, since mounting showed a strong diurnal variation. Five to eight days of such observations were earned out each month between March 1979 and January 1980 (except for July and August when no data was collected) and again between December 1982 and February 1984 (except for January and February 1984 when only two days of observations were done each month). A total of 631 hours of ad libitum records were made in nine months in 1979-80 and 937 hours in 15 months in 1982-84. Besides the study group, six other groups were monitored at intervals of 30-40 days in 1979-80 and 1982- 84. Data on seasonality of births were taken from these groups (see Kumar, 1987). Results Female Sexual Cycle: The female sexual cycle in the lion- tailed macaque is characterized by the cyclical appearance of sexual swelling in the perineal region and at the base of the tail which is conspicuous (Fooden, 1975). The swelling phase had a mean length of 14.1 days (range 8-19 days, n=7) and the non-swelling phase had a mean length of 16.4 days (range 6- 25, n=7). The combined duration of these phases gave a mean cycle length of 30.5 days. More than 80% of the mountings by the adult male occurred JOURNAL, BOMBAY NATURAL HISTORY SOCIETY, 97(1), APR. 2000 43 SEXUAL HARASSMENT AMONG FEMALE LION-TAILED MA CA QUES Fig. 1 : Mounting frequency (per hour) by the adult male on successive days of a sexual swelling cycle of a female: mean for six sexual cycles. The sexual cycles were aligned by the day on which the swellings disappeared (day 0). when the female had sexual swelling. Nearly 84% of these mountings were accompanied by copulatory calls of the females, compared to only 9.1% in the case of females without swelling (x2 = 24.9, df=l,/?<0.001 ). The mounting frequency started to increase 3 to 4 days before the appearance of the swelling and reached a peak (of about 3/hour) four days prior to its disappearance. It then dropped abruptly almost to zero on the last day of swelling (Fig. 1). The interval between the appearance of the swelling and peak sexual activity varied from 10 to 15 days, with a mean of 12.2 days (n=6). When data from 1979-80 and 1982-84 were combined, swellings were seen in the study group m all months of the year except March and April. In May, swelling was seen only in the last week in 1979 and none in 1983 (Fig. 2). Although there are no systematic data from the other groups, no swellings were seen in them during March-May of 1979 and 1983. It appears. therefore, that there is a summer amenorrhea in the lion-tailed macaque in the months of March and April, probably extending to May. There was a synchrony of sexual cycles in the study group soon after the first cycle following the summer amenorrhea (Fig. 2). In 1979, the sexual cycle of two females started in the last week of May, and in June all the five females of the group had sexual cycles. The sexual cycle of two subadult females started only in September-October. All the four adult females which showed swelling in 1982-83 did so in synchrony in October 1983, one sexual cycle after the first cycle of the season. (Four of the remaining five females were in post- partum amenorrhea. The fifth, the oldest female of the group, did not show swelling in 1982-84). The cycle of the subadult female started only one month later. Sexual Harassment: Sexual harassment consisted of activities by members of the group that apparently interfered with sexual 44 JOURNAL, BOMBAY NATURAL HISTORY SOCIETY. 97(1), APR. 2000 SEXUAL HARASSMENT AMONG FEMALE LION- TAILED MA CA QUES Months (1979-80) Months (1983-84) | | One sexual swelling cycle | Sexual swellings leading to conception Fig. 2: The distribution of sexual cycles and conceptions in the adult females of the study group in 1970-80 and 1982-84. interactions between adult male and female. Such interference occurred in 12.8% of the 577 sexual interactions observed. Interference occurred at the premounting stage (i.e. after the initiation of sexual interaction but before mounting) or at the mounting stage. Most of the interference were at the latter stage (70.3%). Out of 74 harassments recorded, 23.0% were by infants and juveniles. These occurred mostly at the mounting stage, and consisted of rushing to the mating pair, and then moving about rapidly in short arcs about 2-3 m away (with tail-wagging and uttering ‘ uh uh ’ sounds) until the mounting was over. Mountings involving females with and without sexual swellings were equally harassed by the immatures (Fisher exact test p= 0.33). Moreover, mounting did not appear to discontinue as a result of such harassment. Harassment by the subadult and adult females was related to the sexual status of the female interacting with the male. In 1982-84. 11.9% of the 270 sexual interactions involving females with swelling were harassed by other adult females, while none of the 69 mountings involving females without swelling were JOURNAL BOMBAY NATURAL HISTORY SOCIETY, 97(1), APR. 2000 45 SEXUAL HARASSMENT AMONG FEMALE LION-TAILED MACAQUES harassed (%2=7.7, df=l, /?<0.01). In 1979-80, 13.1% of the 145 sexual interactions involving females with swelling were harassed by other adult females as opposed to only 2.2% of 90 sexual interactions involving female without sexual swelling (%2=6.8, df=l, /?<0.01). About 5.2% of the sexual interactions were harassed by the subadult and adult females at the pre- mounting stage and a further 12.1% at the mounting stage. Harassment at the pre-mounting stage consisted of a female presenting to the adult male while another female was presenting, often between the male and the first female. Sometimes a female rushed to a presenting female with aggressive calls and chased it away from the male or physically prevented the male from mounting by pulling it by the tail or by standing in the way. Harassment at the premounting stage occasionally resulted in the redirection of mounting to the harasser (21.4%). More often it prevented mounting from taking place. The percentage of sexual initiations which ended in mounting when harassed by adult females (7.0%) was significantly lower than those which were not harassed (58.2%, %2=12.0, df=l, /?<0.001). Harassment at the mounting stage consisted of rushing to the pair with growls, and chasing and often physically attacking the female. Presenting in front of the mounted pair was also seen. Mounting of the harasser soon after mounting the harassed female occurred in 1 1 . 1 % of the cases. When harassment was overtly aggressive the harassed female often ran or jumped away before the male had dismounted. Significantly fewer of the harassed mountings were accompanied by copulatory calls (63.3%) than those which were not harassed (83.6%, X2=4.6, df=l, p<0.05). Harassed mountings had a shorter duration (mean=7.75 secs, s.e=0.89, n=12), than normal mountings (mean=9. 12 secs, s.e=0.35, n=95). However, duration of only those which were aggressively harassed (mean=6.16 secs. s.e=0.72, n=9) was significantly shorter (/-test, t= 2.6, p< 0.05). In short, harassment (a) was mostly by adult females with sexual swelling; (b) was targeted at females with sexual swelling (c) drastically decreased the probability of mounting taking place after the initiations of a sexual interaction, from 0.582 to 0.07; (d) caused a premature termination of mounting and thus probably prevented ejaculation; and (e) redirected mounting from the harassed to the harasser. Harassment and Synchrony in Sexual Swelling: The frequency of harassment varied with the number of females with swelling. At the pre-mounting stage, 1.3% of the sexual interactions were harassed with two females with swelling and 13.7% with four such females (X2=14.5, df=3, /?<0.001, Table 2). Harassment at the mounting stage also increased with the number of females with swelling in the group, although the difference was not significant (X2=5.09, df=3, /?>0.10). Harassment at the mounting stage was significantly more frequent when there were three females with swelling (33.3%) compared to when there was only one (7.3%, Fisher exact test, p=0.04). Table 2 PERCENTAGE OF SEXUAL INTERACTIONS, HARASSED AT THE PREMOUNTING AND MOUNTING STAGES BY ADULT FEMALES, AND ESTIMATED PERCENTAGE OF MATING CURTAILED Number of females with swelling Sexual interactions seen % harassed premount mount stage % harassed mounting stage 0 69 0 0 1 108 1.9 7.3 2 75 1.3 14.3 3 14 7.1 33.3 4 73 13.7 1 1.6 The frequency of mounting by the male showed significant differences between days, depending on the number of females with swelling. (Kruskal-Wallis one-way analysis of variance (K-W test), x2=13.4,p<0.005. Table 3). However, it did not increase in proportion to the 46 JOURNAL. BOMBAY NATURAL HISTORY SOCIETY, 97(1), APR. 2000 SEXUAL HARASSMENT AMONG FEMALE LION-TAILED MACAQUES Table 3 MOUNTING FREQUENCY (PER HOUR) BY THE ADULT MALE AND SUBADULT MALE WHEN THERE WERE 0 TO 4 FEMALES WITHSEXUAL SWELLING IN THE GROUP Number of females with swelling 0 1 2 3 4 Adult male Mean 0.09 0.42 1.66 1.37 1.53 Min. 0.00 0.00 1.14 0.27 1.24 Max. 2.50 1.24 2.53 2.45 1.90 Subadult Mean 0.04 0.04 0.23 0.30 0.22 Male Min. 0.00 0.00 0.00 0.00 0.00 Max. 1.50 0.10 0.38 0.82 0.36 number of females with swelling, but appeared to reach a plateau when there were two females with swelling. The single subadult male in the group in 1982-84 had a mating frequency that was considerably lower than that of the adult male, but seemed to increase as the number of females with swelling increased (Table 3). However, the duration of mounting was considerably shorter for the subadult male (often less than 5 secs), and also did not show the characteristic multiple mount pattern of the adult male. Consequences of Harassment: If harass- ment significantly reduces the frequency of ejaculatory mating, this could result in a reduction in the chances of conception by female. This is particularly so if harassment is asymmetrically distributed among the females, for example due to social dominance. Dominance interactions were relatively few and occurred mainly on major feeding trees when visibility was poor. As a result, the dominance hierarchy of females in the main study group was not precisely known. Moreover, it was often impossible to identify the females because of the speed with which harassments occurred and poor visibility. Therefore, the reproductive consequences of harassment was examined indirectly. The distribution of conceptions and births in the study group was used to test whether females were less likely conceive when there were more than one female with swelling. If this is so, then conceptions and births would not show a synchrony similar to that shown by sexual swelling, but would be more evenly spread out across the months. The date of births in the group during the study period were known. For these, the months of conception were estimated using a gestation period of 172 days (Lindburg and Lasley. 1985). Conceptions did not have a peak corresponding to that of sexual swelling at the beginning of the season (Fig. 1). Of the five females which had swellings in June 1979, only one conceived during that month. There were no data on sexual cycles in July and August, but only one each of four remaining females conceived in July and August. The cycles of the remaining two females continued in synchrony until one conceived in December. Since the second study ended before the births from the 1983-84 mating season (September 1983 to February 1984), stoppage of cycling by females was taken as indicating conception. Two females which showed swelling in September 1 983 did so again in October, when the four females which showed sexual swelling during that mating season, did so in synchrony. The cycle of only one stopped after that month. The remaining three females showed swelling in November (along with a subadult female), but only two conceptions occurred. The cycle of the remaining adult female continued until December 1983. The subadult female’s cycle continued until the end of the field study in February 1984. Population regulation: Sexual harassment could potentially play a population regulatory role since the number of females that postpone conception, especially to the next reproductive year, could increase with group size. If this is the case, then the births in the larger groups should be more dispersed among the months. This was tested with data on births from the mam study group and six other groups that were periodically monitored. The seven groups were divided into two group size classes (12-18 and JOURNAL, BOMBAY NATURAL HISTORY SOCIETY, 97(1), APR. 2000 47 SEXUAL HARASSMENT AMONG FEMALE LION - TAILED MA CA QUES Months Months Fig. 3: The distribution of births in two group-size classes, 12-18 (above) and 19-28 (below). Each square represents one birth. 19-28) based on the mean group size during the study period (Fig. 3). Both the classes had the same mean birth date (Caughley 1977), June 15, but the coefficient of variation for the smaller class (205.0%) was nearly twice that of the larger class (112.2%). Thus, contrary to what was expected, births in the smaller groups were more dispersed through the year than the larger groups. It is also noteworthy that the main study group had a shorter mating season in 1983-84 when the group size was 17, compared to that in 1979- 80 when the group size was 12 (Fig. 2). Discussion Sexual harassment by adult females probably occurs as a consequence of the high synchrony of sexual swelling among the females of a group, a high female/male ratio (5:1), and the tendency for the groups to be one-male units. These could lead to considerable sexual competition among the females. The multiple- mounting pattern of the male (Fooden, 1975; Kumar and Kurup, 1985) might also impose constraints on the mating potential of the male. 48 JOURNAL, BOMBAY NATURAL HISTORY SOCIETY, 97(1). APR. 2000 SEXUAL HARASSMENT AMONG FEMALE LION-TAILED MACAQUES This competition could increase with the number of females in sexual synchrony. The extent to which harassment could affect the probability of conception would depend on the stage of the sexual cycle in relation to ovulation and the degree of asymmetry in the direction of harassment. Even though the frequency of mounting in the first week of swelling was highly variable even when there was only one sexually active female (Kumar, 1987), the peak between 2-5 days prior to deflation indicates that mountings at this stage of the cycle might be critical to conception. Thus, harassment in the last week of swelling could severely affect the probability of conception. At extreme asymmetry, in the direction of harassment, all the curtailed mountings could be of the low-ranking females. In addition, if harassments between females of different ranks differed in aggressiveness (for example, those by dominant females being more aggressive) mounting by the low-ranking females could be curtailed more than those of dominant females since aggressive harassments were more effective in curtailing mounting. Birth rate in the lion-tailed macaques is a decreasing function of group size and the number of adult females in the group (Kumar, 1995b). Sexual harassment could lead to such an effect and thus serve as a population regulatory factor, if two conditions are met: i) the proportion of females coming into sexual synchrony during the mating season should be constant with group size, so that their absolute number would increase with group size; and ii) groups should be either one male units irrespective of group size, or when there is more than one male, only one of them is reproductively active during all the phases of the sexual cycle of the females. If these conditions are met, then the mating season should be more prolonged with increasing group size, as more females postpone conception. Therefore, births should be more dispersed in the larger groups and have a higher coefficient of variation. The limited data on the main study group shows that the mating season gets shorter, and not longer as predicted, as the group becomes larger. Also, contrary to the second prediction, births were relatively less dispersed in the larger groups than in the smaller groups. This was probably because of the violation of the above two conditions. It is known that females do not ovulate until they reach a particular nutritional level (Frisch and McArthur 1974). Since resource competition increases with group size, it could be expected that the number of females able to build up sufficient nutritional reserves, so as to start ovulation, would decrease with increasing group size. There is no systematic data on the number of females coming into sexual cycle as a function of group size. In one large group with more than 25 members, which was regularly censused, not more than 4 of the 1 2 females were ever seen with sexual swelling on the same day. Since births in the larger groups were few, it was unlikely that other females were in post-paitum amenorrhea. Moreover, although the group was seen almost every month in 1979, swellings were seen only in June and November-December (with 2 and 3-4 females respectively). In addition, the number of adult and subadult males increase with group size (Kumar, 1987). No data was collected on the sexual behaviour of males in multi-male groups. The limited data on sexual behaviour of the subadult male of the study group indicate that mounting frequency of subadult males increased with the number of sexually active females in the group (Table 3). Even if mountings by the subadult male (and probably low ranking adult males of multi- male groups) are confined to the early follicular and luteal phases of the cycle, such mountings could significantly reduce the sexual competition between the females with overlapping sexual cycles. As a result, mountings by the adult male (or dominant male in multi-male groups), even if only confined to the late follicular phase, could be less harassed by other females which are in other phases of the cycle. JOURNAL, BOMBAY NATURAL HISTORY SOCIETY. 97(1), APR. 2000 49 SEXUAL HARASSMENT AMONG FEMALE LION-TAILED MACAQUES The short birth season in the larger groups might be, therefore, a cumulative function of (a) fewer females coming into sexual cycles in each season which in itself would significantly reduce female sexual competition and (b) more adult males in the larger groups which would further reduce female sexual competition. Thus, it appears unlikely that sexual harassment could be a population regulatory factor, in the small and large groups. In the former, in spite of female sexual competition (resulting from one male and several sexually active females), postponement of conception is expected to be only within the mating season. In the larger groups, on the other hand, fewer females ovulate in the mating season. It is possible that ovulating females are still sufficiently numerous in the medium sized one male groups, so that sexual competition could be high. A few females would be forced to postpone conception to the next mating season thus leading to/reproductive suppression. Postponement of conception within the season could serve indirectly as a population R E FE Abbott, D.H. (1988): Social suppression of reproduction in primates. In: Comparative Socioecology: The Behavioural Ecology and Humans and Other Animals (Eds. V. Standen and R.A. Foely), pp. 285-304. Abbot. D.H., E.B. Keverne, G.F. Moore & U. YoDYiNGYARd (1986): Social suppression of reproduction in subordinate talapoin monkeys, Miopithecus talapoin. In: Primate Ontogeny, Cognition and Reproductive Behaviour (Eds. J.G. Else and P.C. Lee), Cambridge University Press, Cambridge. Pp 329-34 1 . Caughley, G. ( 1 975): Analysis of Vertebrate Populations. Wiley, Chichester. Drickamer. L.C. (1974): A ten-year summary of reproductive data for free-ranging Macaca mulatto. Folia Primatologica 21: 61-80. Dunbar, R.I.M. (1980): Determinants and evolutionary consequences of dominance among female gelada baboons. Behavioural Ecology and Sociobiology 7: 253-265. Fooden, .1. (1975): Taxonomy and Evolution of Liontai 1 and Pigtail Macaques (Primates : Cercopithecidae). regulatory factor. Increased mortality of infants born in late season has been reported; for example in M. mulatto (Drickammer, 1974) and in A. palliatta (Froelich et al., 1981). Since postponement of conception is expected to increase with group size within the small to medium-size range, late season births and infant mortality could be expected to increase with group size within that range. Acknowledgements I am grateful to Tamil Nadu Forest Department for facilities provided in the field; to Zoological Survey of India for funding in 1 977- 80, to Wenner-Gren Foundation, L.S.B. Leaky Foundations, WWF-US, WWF-India, and Cambridge Commonwealth Trust for grants in 1981-87; and to Wildlife Conservation Society, New York, for grants in 1987-89. Earlier drafts of this paper greatly benefited from comments by Drs. D.J. Olivers, E.L. Bennet, J.M.Y. Robertson, E. Barret, L. Fuller, E.B. Keverne, and G.W. Norton. e n c e s Bibliotheca Primatologica 10. Basel, Karger. Frisch, R.E. & E. McArthur (1974): Menstrual cycles: fatness as a determinant of minimum weight for the maintenance or onset. Science 185: 949-95 1 Froelich, J.W., Thorington, Jr., & J.S. Otis ( 1 98 1 ): The demography of howler monkeys ( Alouatta palliata) on Barro Colorado Island, Panama. International Journal of Primatology 2: 207-236. Keverne. E.B. (1983): Endocrine determinants and constraints on sexual behaviour in monkeys. In: Mate Choice, (Ed. P. Bateson). Cambridge University Press, Cambridge, pp. 407-420. Kumar, A. (1987): The Ecology and Population Dynamics of the Lion-tailed macaque (Macaca silenus) in South India. Ph.D. Dissertation submitted to the University of Cambridge, U.K. Kumar, A. (1995a): The life history, ecology, distribution and conservation problems in the wild. In: The Lion- taled Macaque: Population and Habitat Viability Assessment Workshop. Zoo Outreach. Coimbatore, India. Kumar, A., S. Molurand S. Walker (Eds.). Kumar. A. ( 1995b): Birth rate and sun ival in relation to 50 JOURNAL. BOMBAY NATURAL HISTORY SOCIETY. 97(1). APR. 2000 SEXUAL HARASSMENT AMONG FEMALE LION-TAILED MACAQUES group size in the lion-tailed macaque, Macaco silenus. Primates. 36: 1-9. Kumar, A. & G.U. Kurup (1 985): Sexual behaviour of the Lion-tailed macaque, Macaca silenus. In: The Lion- tailed Macaque: Status and Conservation (Ed. P.G. Heltne), pp, 1 09- 1 30, Alan R. Liss, New York. Lindburg, D.G. & B.L. Lasely (1985): Strategies of optimising the reproductive potential of lion-tailed macaque colonies in captivity, lit: The Lion-tailed Macaque: Status and Conservation (Ed. P.G. Heltne). pp. 34-56. Alan R. Liss. New York. Mori. A. (1979): Analysis of population changes by body weight in the Koshima troop of Japanese monkeys. Primates 20: 371-397. Wasser, S.K. (1983): Reproductive competition and cooperation among female yellow baboons. In: Social Behaviour of Female Vertebrates. (Ed. S.K. Wasser), Academic Press, New York. pp. 349-390. JOURNAL BOMBAY NATURAL HISTORY SOCIETY. 97(1). APR. 2000 51 SEASONAL CHANGES OF TROPICAL FOREST BIRDS IN THE SOUTHERN WESTERN GHATS1 E.A. Jayson2 and D.N. Mathew3 ( With seven text-figures) Key words: Seasonal changes, forest birds, Western Ghats, Kerala, India A study was carried out in the tropical forests of Silent Valley and Mukkali in the Western Ghats, Kerala from May 1988 to April 1993, to elucidate the seasonal changes of bird communities in the two vegetation types. Abundance and density of birds were assessed, using variable width line transects each month. The highest populations, 609-1 ,892 /km2 were found from December- April. Total number, monthly density and species richness of birds declined during monsoon. When compared, abundance and density of birds, observed in the evergreen forests was more (929 /km2) than in moist deciduous forests (747 /km2). However, bird population showed more stability in the moist deciduous forests. Except for two summers, significantly higher bird density was obtained in the evergreen forests during summer (1,074 /km2). Bird species diversity was high during summer and low in monsoon in both the vegetation types. A direct negative relationship was also obtained between the rainfall, total number of birds, bird density and total number of bird species in the evergreen forests. Significant positive correlation was obtained between the temperature and bird community parameters in the evergreen forests, whereas rainfall and temperature showed no significant effect on the bird community in the tropical moist deciduous forests. Introduction Tropical forests support a stable population of birds m all seasons, whereas marked variations have been noted in temperate forests (Wright, 1970; Kricher, 1975). Seasonal variation of forest birds has been reported from several other countries (Anderson, 1972, Morrison etal. 1980, Pyke, 1984). No information, however, is available on the seasonal trends of tropical forest birds of the Western Ghats of South India. An attempt has been made to monitor the seasonal changes of bird communities in the tropical evergreen forests and the southern secondary moist mixed deciduous forest of Kerala. Birds of Kerala have been studied by Ali (1969), Ali and 'Accepted April, 1999 ;Division of Wildlife Biology Kerala Forest Research Institute Peechi 680 653, Kerala, India. ■ Department of Zoology University of Calicut, Calicut University P.O, Kerala, India. Ripley (1983a) and All and Ripley (1983b) earlier. Ecological studies were carried out at Silent Valley by Balagopalan (1990) and Balasubramanian (1990). Ramakrishnan (1983) studied the ecology of birds in the Malabar forests. Daniels (1989) and Daniels et al. (1990) reported many aspects of birds of the northern Western Ghats. StudyArea Location and topography: The study areas, Silent Valley and Mukkali are located in Palakkad dist., Kerala State, between 1 1° 3' and 11° 13' N lat., and between 76° 25' and 76° 35' E long. They lie in the Western Ghats of south India and form part of the Nilgiri Biosphere Reserve (Fig. 1). After evaluating the entire area, two intensive study sites were selected: a tropical evergreen forest, Silent Valley, and a moist deciduous forest at Mukkali. The elevation of the study sites varied from 500 m to 1500 m above msl. The topography is undulating. According 52 JOURNAL, BOMBAY NATURAL HISTORY SOCIETY, 97(1), APR. 2000 SEASONAL CHANGES OF TROPICAL FOREST BIRDS JOURNAL, BOMBAY NATURAL HISTORY SOCIETY, 97(1), APR. 2000 53 Fig. 1 : Location of the study area SEASONAL CHANGES OF TROPICAL FOREST BIRDS to Udvardy (1975), Silent Valley and Mukkali fall under the Malabar Rain Forest Realm. These two study sites are at a distance of about 20 km with a difference in elevation of 400 m between them. Climate: There are two distinct seasons in the study area, monsoon starting from the end of May, up to mid-November, and the dry summer from December to April. Mukkali (4,227 mm/year) receives less rainfall compared to Silent Valley (5,096 mm/year). Heavy rainfall, 803 mm to 2,043 mm/month, was recorded at Silent Valley. From December to March, there is practically no rain. Temperature ranged froml9°C to 22°C at Silent Valley and 21°C to 27°C at Mukkali. Vegetation: A total of 966 species of angiosperms belonging to 559 genera and 134 families were recorded from Silent Valley and adjacent areas (Manilal, 1988). Pascal (1988) described the vegetation of the area as Cullenia exarillata-Mesua ferrea-Palaquium ellipticum type. It is characterised by the abundance of these three species, which may constitute about 80% of the large trees. Degraded areas and other vegetation types like grasslands are also common here. Vegetation of Mukkali is southern secondary moist mixed deciduous forest (Champion and Seth, 1968), degraded to some extent. Methods After considering all the available techniques, variable width line transect method described by Burnham et al. (1981) was adopted. Whenever a bird was spotted, it was identified up to the species level and details like the number of birds, perpendicular distance from the transect, height at which it is located in the canopy and habitat features were noted. Two line transects were selected, one at Silent Valley and the other at Mukkali; each transect was 4 km in length. The first transect covered evergreen forests and the second habitats like moist deciduous forests, rocky patches and fire burned moist deciduous forests. Census was started 30 minutes after sunrise in all the months. Transects were covered at a uniform speed. No census was done on days with very heavy rain and fog. Two samples were collected from each area in a month. The second sample was started from the end of the first sample. A total of 1 50 samples were collected between May 1988 and 1993. No systematic data was collected on nocturnal birds. All calls were considered as single individuals. Perpendicular distances were measured approximately up to metres. To help distance assessment, known distances were measured and marked on trees using a Range Finder before the census. Abundance of birds in each month obtained from the census was used for analysis. Seasonal index of birds for each month was calculated using Time Series Analysis by the method of Simple Averages (Rao, 1983). The formula used is given below: Monthly average Seasonal Index = x 100 Sum of monthly averages Analysis of variance was employed to find any significant difference existing in the total number of birds among the months. The Fourier Series Method was employed for calculating density from the ungrouped perpendicular distances from the transect. All the assumptions described by Burnham et al. ( 1 98 1 ) were followed during the census. Students ‘t’ test was applied to find out the significant difference in the number of birds between summer and monsoon. Diversity was calculated using Shannon- Wener Index (H - -X (pi In pi) with the program specdivers.bas developed by Ludwig and Reynolds (1988). Spearman Rank Correlation was used to find out the correlation between climatic parameters and bird community parameters. 54 JOURNAL, BOMBAY NATURAL HISTORY SOCIETY. 97(1), APR. 2000 SEASONAL CHANGES OF TROPICAL FOREST BIRDS Table 1 SEASONAL INDEX OF BIRDS PRESENT IN EACH MONTH AT SILENT VALLEY AND MUKKALI Area Months .1 F M A M J .1 A S O N D Silent 114 109 88 81 95 54 58 59 119 101 136 153 Valley Mukkali 113 92 131 89 84 113 73 87 116 99 133 70 Results Patterns of change Monthly variation: During September to February, more birds were present at the Silent Valley compared to the annual average of 100 (Table 1). In Mukkali, higher number than the annual average were observed during the months of January, March, September and November. Highest Seasonal Index (133) was obtained in November. Analysis of variance showed a significant difference in the total number of birds among the months at Silent Valley (F= 6.18; P= 0.01), whereas no significant difference was obtained at Mukkali (F= 1.95; P= 0.08). Seasonal variation in a year: The total number, monthly density and species richness of birds at Silent Valley and Mukkali declined during monsoon and increased in the dry months (Table 2). No significant difference in total number was obtained between monsoon and summer at Silent Valley and Mukkali (Silent Valley ‘t’=1.63, P=0.14; Mukkali ‘f=0.28, P=0.79). Species like the black bulbul ( Hypsipetes madagciscariensis ), emerald dove(Chalcophaps indica) and the imperial pigeon ( Ducula badia) were practically absent during monsoon at Silent Valley. Seasonal change over the years: Total number of birds: Data were pooled into two seasons, monsoon and summer, to find out the seasonal differences in the total number of birds over the years. Chi-square test revealed a significant difference in the number of birds between the seasons at Silent Valley (Table 3). The highest number of birds per month (91) was observed in the 1991 summer and the lowest (53) in the monsoon of 1992. At Mukkali, there was no significant difference among seasons in the total number of birds. Significant difference in the number of birds per month between Silent Valley and Mukkali was observed during three summers. During these seasons, there were more birds at Silent Valley. But during the 1992 summer and monsoon, no significant difference in the number of birds was observed, both at Silent Valley and Mukkali. Table 2 COMM UNITY PARAMETERS OF BIRDS RECORDED DURING TWO SEASONS (1 988-1993) Area Monsoon season Summer season No. of birds (mean) 70.00 (±28.63) 90.33 (±32.25) Silent Valley Density (birds/knr ) 958. 16 (±478.58) 1286.17 (±781 .1 8) Species richness 28.33 (±6.87) 43. 16 (±7.00) No. of birds (mean) 60.67 (±12.61) 56.5 (±12.91) Mukkali Density (birds/knr) 854.33 (±400.43) 707.00 (±285.36) Species richness 30.67 (±9.35) 39.17 (±10.23) Standard Deviation is in parenthesis JOURNAL, BOMBAY NATURAL HISTORY SOCIETY, 97(1), APR. 2000 55 SEASONAL CHANGES OF TROPICAL FOREST BIRDS Table 3 MEAN NUMBER OF BIRDS RECORDED PER MONTH IN DIFFERENT SEASONS AT SILENT VALLEY AND MUKKALI Seasons Silent Valley Mukkali Total X2 P= Monsoon 1988 70 76 146 0.25 NS Summer 1989 95 52 147 12.58 0.02 Monsoon 1990 74 48 122 5.50 0.02 Summer 1991 91 50 141 11.92 0.001 Monsoon 1992 53 67 120 1.63 NS Summer 1992 83 70 153 1.10 NS Summer 1993 Total X2 P = 89 555 16.36 0.02 59 422 1 1.83 NS 148 6.08 0.02 NS= Not Significant Species Richness: There is no significant difference in bird species richness between years in monsoon (x2=4.26; P=0.05) and summer (X2=8.92; P=0.05) at Silent Valley. But a significant difference was obtained between years in both seasons at Mukkali (Monsoon x 2 =38.97; P=0.001, Summer %2= 14.64; P=0.001). Density: Significant difference in density was obtained between seasons in different years at Silent Valley and Mukkali. The values for summer and monsoon showed a significant difference (Silent Valley: %2=62.25, P=0.05, df=l; Mukkali: %2=39.33, P=0.05, df=l). Bird density was high during summer, both at Silent Valley and Mukkali. Except for two summers, significantly higher bird density was observed at Silent Valley in summer (Table 4). Diversity: Variations in the diversity of birds, based on Shannon- Wener diversity index, in different seasons at Silent Valley and Mukkali are given in Table 5. Diversity index showed high values in summer (X=3.12, 11=5) and lower during monsoon (X=2.65, n=4), at Silent Valley and Mukkali (monsoon: X=2.78. n=4 and summer: X=3.14, n=5). Table 4 SEASONAL VARIATION IN BIRD DENSITY AT SILENT VALLEY AND MUKKALI Seasons Density/sq. km Mean density Silent Valley Mukkali Total Mean X2 P = Monsoon 1036 638 1674 837 94.63 0.00 1 1988 (3.23) (5.01) Summer 2123 1662 3785 1892.5 56.15 0.001 1989 (2.21) (7.72) Monsoon 685 401 1086 543 74.27 0.001 1990 (3.03) (7.86) Summer 741.4 370 1 1 1 1 .4 555.7 124.11 0.001 1991 (3.93) (14.99) Monsoon 493 792 1285 642.5 69.57 0.001 1992 (9.11) (3.06) Summer 823 757 1580 790 2.76 NS 1992 (6.03) (4.34) Summer 608 688 1296 648.0 4.94 NS 1993 (10.91) (5.61) Total 6509.40 5308 X2 1976.52 1471.29 P = 0.001 0.001 NS= Not Significant; The values in the brackets denote coefficient of variation of the estimates. 56 JOURNAL, BOMBAY NATURAL HISTORY SOCIETY, 97(1), APR. 2000 SEASONAL CHANGES OF TROPICAL FOREST BIRDS Factors affecting the seasonal variation Rainfall: A direct relationship was obtained between rainfall and number of birds, density and total number of bird species at Silent Valley. When rainfall increased, all of these three community parameters decreased, and vice versa (Figs. 2, 3 & 4). At Mukkali also, rainfall had its influence on bird community, but not in the same magnitude as that of Silent Valley (Figs. 5, 6 & 7). At Silent Valley, significant negative correlation was obtained between the mean of monthly total rainfall (1988-1993) and number Fig J FMAMJ JASOND Months 2: Relation between rainfall and number of species at Silent Valley 1400 J FMAMJ JASOND Months Fig. 3: Relation between rainfall and number of species at Mukkali Species 19 Rainfall Species Rainfall JOURNAL, BOMBAY NATURAL HISTORY SOCIETY, 97(1), APR. 2000 57 SEASONAL CHANGES OF TROPICAL FOREST BIRDS 1600 1400 1200 1000 | 800 75 c 600 c 400 200 0 J FMAMJ JASOND Months Fig. 4: Relation between rainfall and abundance of birds at Silent Valley i ou 1400 -1200 -1000 E 800 £ 75 600 1 DC 400 200 0 J FMAMJ JASOND Months Fig. 5: Relation between rainfall and number of birds at Mukkali Birds 91 Rainfall Birds Rainfall of species in each month (r= -0.731, P= 0.01, n= 12). Significant correlation was also obtained between mean monthly rainfall and total number of birds in each month (r= -0.66, P= 0.05, n= 12). But there was no significant correlation between the density of birds in each month and rainfall (r= -0.45, P= 0.05, n= 12). At Mukkali, no significant correlation was obtained between monthly rainfall and bird community parameters. Here, monthly rainfall showed negative correlation with the number of bird species (r= -0.41, P= 0.05, n= 12) and there was no significant correlation between monthly rainfall and the total number of birds (r= -0.21, 58 JOURNAL, BOMBAY NATURAL HISTORY SOCIETY. 97(1), APR. 2000 SEASONAL CHANGES OF TROPICAL FOREST BIRDS 3000 1600 J FMAMJ JASOND Months Fig. 6: Relation between rainfall and density of birds at Silent Valley 1400 1200 1000 £ 800 » & 600 J FMAMJ JASOND Months Fig. 7: Relation between rainfall and density of birds at Mukkali Density Rainfall Density Rainfall P= 0.05, n=12) and their density (r= -0.06, P= 0.05, n= 12). This suggests that rainfall does not have any significant effect on the bird community at Mukkali. Temperature: There was significant positive correlation between temperature and bird community parameters at Silent Valley. Number of species increased with increase in temperature (Coefficient of correlation r= 0.57, P= 0.05, n= 12). Similarly, total number of birds (r= 0.83, P= 0.001, n= 12) and their density (r= 0.62. P= 0.05, n= 12) showed an upward trend as the JOURNAL, BOMBAY NATURAL HISTORY SOCIETY, 97(1), APR. 2000 59 SEASONAL CHANGES OF TROPICAL FOREST BIRDS Table 5 SEASONAL VARIATION IN DIVERSITY (H’) AT SILENT VALLEY AND MUKKALI Seasons Silent Valley Mukkali Monsoon 1988 2.77 2.50 Monsoon 1989 2.38 2.63 Monsoon 1990 2.70 2.85 Monsoon 1992 2.74 3.13 Mean 2.65 2.78 Summer 1 989 3.20 2.96 Summer 1 990 3.01 2.95 Summer 1991 3.23 3.08 Summer 1 992 3.29 3.46 Summer 1993 2.88 3.25 Mean 3.12 3.14 temperature increased during summer. At Mukkali, no such significant correlation was found (temperature and number of species r= 0.21, P= 0.05, n= 12; temperature and total number of birds r= -0.08, P= 0.05, n= 12). Discussion Patterns of change: During monsoon, there was reduction in the number of birds both at Silent Valley and Mukkali. Birds appeared to move locally to avoid the unfavourable climate. Local movements in search of optimum habitats are possible because of the availability of other habitats in the vicinity as the tracts where the study was conducted were fragmented forest patches. Similar trends were reported from the tropical forests of other countries also. Variation in rainfall and soil moisture makes tropical bird fauna seasonal (Greenberg and Gradwohl, 1986). According to them, this is due to the influence of rainfall on phenological patterns of trees, which in turn affect the population trends of arthropods. Karr (1976) also showed the effect of high rainfall on the seasonal patterns of birds. Higher numbers of birds were recorded during summer in two vegetation types. A greater abundance of birds was found at Silent Valley during summer than at Mukkali. Density of birds and their diversity indices were also higher for Silent Valley during summer, which can be attributed to the availability of more fruits at Silent Valley during summer. However, at Mukkali, the bird population showed much more stability. Factors influencing the seasonal variations: Rainfall and temperature were the major factors influencing the abundance of birds at Silent Valley and Mukkali. Price (1979) who worked on the birds of Eastern Ghats also found a similar trend in annual cycles of bird fauna due to changes in rainfall. As mentioned earlier, a few species of birds like the yellowbrowed bulbul ( Hypsipetes indicus ) showed stability in population even in the fluctuating environment. This can be attributed to the resident nature of the species, coupled with its ability to feed on various food types like berries, drupes, nectar, spiders and insects. Stiles (1978) had also shown that in tropical forests bird communities fluctuated in number as a response to the availability of food and climate changes. The relationship between food resources and bird diversity was also reported by Terborgh (1985). Even though tropical forest birds are considered sedentary, MacArthur (1972) has shown that seasonal movements are fundamental in many species as an adaptive strategy in varied forest habitats. This study also showed that rainfall and temperature influence the tropical evergreen forest bird community, whereas such climatic factors have little effect on birds of moist deciduous forests. Acknowledgements Statistical analyses were done with the help of Ms. K.A. Mercey, Asst. Prof., College of Veterinary and Animal Sciences, Mannuthy, Trichur. We thank the field staff of Silent Valley National Park for their help and the Dept, of Environment, Govt, of India for support. 60 JOURNAL . BOMBAY NATURAL HISTORY SOCIETY, 97(1), APR. 2000 SEASONAL CHANGES OF TROPICAL FOREST BIRDS References Ali, S. (1969): The Birds of Kerala. Oxford University Press, Bombay, pp. 444. Ali, S. & S.D. Rjpley ( 1 983a): Handbook of the Birds of India and Pakistan. Oxford University Press, New Delhi, pp. 737. Ali, S. & S.D. Rjpley (1983b): A Pictorial Guide to the Birds of the Indian Subcontinent, Bombay Natural History Society. Bombay, pp. 1 77. Anderson, S.H. (1972): Seasonal variations in forest birds of Western Oregon, Northwest Science 46(3) : 1 94- 206. Balagopalan, M. (1990): Soil and plant community relationships in wet evergreen forests of Silent Valley. In: Ecological Studies and Long-term Monitoring of Biological Process in Silent Valley National Park. KFRI Research Report, pp. 1 35-206. Balasubramanian, K. (1990): Establishment of permanent sample plots for long-term monitoring of ecological process. In : Ecological Studies and Long-term Monitoring of Biological Processes in Silent Valley National Park. KFRI Research Report, pp. 201-231. Burnham, K.P., D.R. Anderson & J.L. Laake (1981): Line transect estimation of bird population density using a Fourier series. In: Estimating the number of terrestrial birds. Eds. C.J. Ralph and M.J. Scott. Studies in Avian Biology No. 6. Cooper Ornithological Society. Champion, H.G. & S.K. Seth ( 1 968): A Revised Survey of the Forest Types of India. Govt, of India. 404 p. Daniels, R.J.R. (1989): A conservation strategy for the birds of the Uttara Kannada District. Ph.D. Thesis. Indian Institute of Science, Bangalore. Daniels R.J.R, M. Hegde & M. Gadgil (1990): Birds of the man-made ecosystems: the plantations. Proc. Indian Acad. Sci (Anim. Sci.) 99(1): 79-89. Greenberg, R. & J. Gradwohl (1986): Constant density and stable territoriality in some tropical insectivorous birds. Oecologia (59:61 8-625. Karr, J.R. ( 1 976): Seasonality, resource availability and community diversity of tropical bird communities. Amer. Nat. // 0:973-974. Kricher, J.C. (1975): Diversity in two wintering bird communities: Possible weather effects. Auk 92 (4):! 66-111. Ludwig, J. A. and J.F. Reynolds (1988): Statistical Ecology, John Wiley and Sons, New York. MacArthur, R.A. (1972): Geographical Ecology. Harper and Row, New York. Manilal, K.S. (1988): Flora of Silent Valley Tropical Rainforests of India. The Mathrubhmni (MM) Press, Calicut, pp. 398. Morrison, M. L.. A. Kimberly & 1. C. Timossi (1980): The structure of a forest bird community during winter and summer Wilson Bull., 98(2): 214-230. Pascal, J.P. (1 988): Wet Evergreen Forests ofthe Western Ghats of India, Ecology, Structure, Floristic Composition and Succession. Institute Francais de Pondicherry. Pondicherry. 239 p. Price, T.D. ( 1 979): The seasonality and occurrence of birds in the Eastern Ghats of Andhra Pradesh. J. Bombay nat. Hist. Soc. 76(3):319-422. Pyke, G.H. (1984): Seasonal patterns of abundance of insectivorous birds and flying insects. Emu 85(1): 34-39. Ramakrishnan, P. (1983): Environmental Studies on the Birds of Malabar Forest. Ph. D. Dissertation. University of Calicut. Rao, G.N. (1983): Statistics for Agricultural Sciences. Oxford and IBH Publ. Co. pp 280. Stiles, F.G. (1978): Temporal organization of flowering among the Humming bird, food plants of a tropical forest. Biotropica 10: 1 94-2 10. Terborgh, J. (1985): Habitat selection in Amazonian birds In: Habitat selection in birds. Ed. M.L. Cody, Academic Press, New York. 3 1 1 -338. Udvardy, M.D.R. (1975): A classification of the biogeographical provinces of the world. IUCN Occasional Paper. 18 IUCN. Gland, Morges. Wright, J.S. (1970): Competition between insectivorous lizards and birds in Central Panama. Amer. ZooL, 19:1 145-1156. JOURNAL, BOMBAY NATURAL HISTORY SOCIETY, 97(1), APR. 2000 61 PLODIA INTERPUNCTELLA (HUBNER) (PHYCITIDAE : LEPIDOPTERA) AS A POTENTIAL PEST OF DRY FRUITS1 S.P. Rad3, H.R. Pajni andNeelima Talwar2 Key words: Plodia interpunctella , dry fruits, susceptibility, weight loss, development period, moisture content Relative susceptibility of 1 2 types of dry fruits viz., almond, apricot, cashewnut, chilgoza, coconut, date, fig, hazelnut, mulberry, pista, raisin and walnut and 1 0 varieties of pista procured from Iran i.e. Ebrahimi, Fandoghi, Gholam Rezaia, Jabbary, Kallenghoochi, Momtaz, O’hadi, Rezaia, Shasti and Wahedi to the attack of Plodia interpunctella (H.) has been studied for the first time. The results showed that cashewnut and pista were the most susceptible and date the least. Out of 10 pista varieties, the varieties Rezaia and Wahedi were the most resistant while the cultivars Fandoghi and Momtaz were the most susceptible. The index of susceptibility has been calculated on the basis of weight loss of fruits and development period and progeny of the pest. Introduction The Indian meal moth Plodia inteipunctella (Hubner) (Phycitidae : Lepidoptera) is an important pest of stored cereals, legumes and dry fruits. The damage is caused by the larvae: besides consuming the product they also spoil it with their webbings and faecal matter, making it unfit for human consumption. A large number of studies have been made on its general biology. Hoppe (1981), Mbata (1987, 1990) and Stein (1990) studied the development pattern while food preference was studied by Lecato (1976). Observations on oviposition behaviour have been made by Mullen and Arbogast (1977), Mbata (1985, 1990) and Almasi et ah, (1987). Grant (1974), Grant and Brady (1975) and Grant (1976) studied the copulation while Grant (1974), Grant and Brady (1975), Ono (1981) and Rangaswamy ( 1 985) made observations on the role of pheromones. The diapause behaviour has been studied by Bell and Walker (1973) and Bell ( 1 976a, 1 976b). However, only a few dry fruits have been tested as hosts of this pest. Myers (1928) studied the relative preference of the pest for a few dry fruits. Hamlin et al.,{ 1931 ), Simmons ( 1 93 1 ) and William ( 1 964) observed development in some dry fruits and 'Accepted February, 1998 : Department of Zoology, Panjab University, Chandigarh 160014, India. 'Present Address: 28, Matyer Feeroze Lane, Caroon Street, Azarbyjan Street, Tehran - Iran 1 3448. cereals. Mullen and Arbogast (1977) studied oviposition on peanuts and dates while Mbata and Osuji ( 1 983) studied the development in whole and cracked groundnuts. The present communication deals with the relative susceptibility and extent of damage to 12 dry fruits and 10 varieties of pista, to assess the potential of P. interpunctella (Hubner) as a pest of stored dry fruit. Material and Methods Adults of Plodia interpunctella (H.) used in the present study were taken from stock cultures raised in the laboratory from small samples collected from Delhi and Chandigarh. The cultures were maintained on different foods stored in an electric incubator fixed at 30 ± 1° C and 75-85% R.H. The foods used for stock cultures as well as those selected for different experiments were sterilized at 50° C for two hours in order to eliminate any parasites or other microorganisms. The twelve selected dry fruits were Prunus amygdalus Batsch almond, Primus armeniaca L. apricot, Anacardium occidental L. cashewnut, Pinus gerarcliana chilgoza, Cocos nucifera L. coconut, Phoenix dactylifera L. date, Ficus glomerata fig, Corylus spp. hazelnut, Morus nigra L. mulberry, Pistacia vera L. pista, Vitis vinifera L. raisin and Juglans regia L. walnut. The susceptibility index of different dry 62 JOURNAL . BOMBAY NATURAL HISTORY SOCIETY. 97(1) APR. 2000 PLODIA INTERPUNCTELLA ASA POTENTIAL PEST OF DRY FRUITS fruits was studied out by keeping ten three-day old eggs mixed with 2 gm of nuts. Three replications were kept in each case. The samples were reweighed after emergence to determine the loss of weight due to consumption by the larvae. The moisture content of the samples was also calculated at the beginning and the end of the experiment and loss/ increase in weight due to moisture variation was considered while calculating actual weight loss. The percentage weight loss due to moisture content variations has been calculated by using the following relationship given by Jamieson (1970). 100 (M,-M.) G = — - 100 -M, Where M, = Initial moisture content percentage wet basis. M, = Final moisture content percentage wet basis. Knowing the value of G, the loss Or gain in weight due to variation in moisture content (d) can be calculated as under, and necessary correction in weight loss of the food made. GxW, d - — 100 Where W = Observed weight loss of the food. G =* Loss or gain percentage in weight due to moisture content variation. The data obtained were subjected to statistical analysis. Results and Discussion The relative susceptibility of twelve types of dry fruits was calculated on the basis of food consumed, the number of adults emerged, duration of developmental period and weight loss of the fruits. The results given in Table 1 showed that amount of different foods consumed by the larvae varied greatly, the largest amount being consumed in mulberry (1.816 gm) and the least in the case of coconut (0.004 gm). Appreciable differences have also been noted in the average development period. Pista registered the shortest development period of 31.71 days, whereas, date showed the longest development period of 104.25 days. However, Hamlin et al. (1931) observed more rapid development of larvae on figs among three fruits namely raisins, prunes and figs tested by them. The progeny produced was maximum in pista, walnut, cashewnut and almond, while other fruits produced comparatively much less progeny. The Table 1 WEIGHT LOSS OF 1 2 DRY FRUITS DUE TO THE ATTACK OF PLODIA INTERPUNCTELLA (H.) (based on three observations) Food Initial Weight of food mean (gm) Final Weight of food mean (gm) Moisture Content M, M, Weight loss Mean % age weight loss Corrected mean % age weight loss Mulberry 2 0.184 8.96 7.326 1.816 90.80 90.768 Fig 2 0.593 10.32 9.949 1.407 70.35 70.345 Cashewnut 2 1.149 4.38 3.307 0.851 42.55 42.541 Almond 2 1.248 3.82 3.410 0.752 37.60 37.597 Walnut 2 1.449 3.40 2.208 0.551 27.55 27.544 Pista 2 1.485 3.34 2.828 0.515 25.75 25.748 Raisin 2 1.497 12.12 6.479 0.503 25.15 25.120 Hazelnut 2 1.550 3.46 2.387 0.450 22.50 22.496 Date 2 1.713 9.26 7.764 0.287 14.35 14.346 Apricot 2 1.862 17.54 14.607 0.138 6.90 6.896 Coconut 2 1.996 2.98 2.550 0.004 0.20 0.200 Chilgoza 2 Nil Nil Nil Nil Nil Nil JOURNAL, BOMBAY NATURAL HISTORY SOCIETY, 97(1), APR. 2000 63 PLODIA INTERPUNCTELLA ASA POTENTIAL PEST OF DRY FRUITS larvae failed to survive on apricot, coconut and chilgoza as they do not get sufficient nutrition to reach maturity. In fact, the larvae did consume some food in the case of apricot and coconut but died before reaching the pupal stage. In the case of chilgoza, on the contrary, the larvae did not consume any food. The relative suitability of different foods was also determined with the help of the formula Log eY/T, given by Osuji (1976), where Y is the number of progeny, T is the time taken by 50% of the adults to emerge and e is a constant with a value of 2.303 (Table 2). Pista, walnut and hazelnut, with a suitability index value of 1.743, 1.590 and 1.393, were the most suitable food while date with an index value of 0.085 was the least suitable food. The relative susceptibility of various foods can be judged by combining the amount of food consumed with the index of suitability (Table 3). Cashewnut and pista with susceptibility index values of 49.773 and 44.878 respectively, were the most susceptible foods whereas date with the index value of 1.219 was the least susceptible food. Table 2 RELATIVE SUITABILITY OF 12 DRY FRUITS TO THE ATTACK BY PLODIA INTERPUNCTELLA (H.) (based on three replications of 1 0 eggs each) Food Progeny Y (Average) Development period (Todays) Index of suitability L°SeY/T50 Pista 24 31.71 1.743 Walnut 25 36.20 1.590 Hazelnut 21 34.71 1.393 Cashewnut 25 49.20 1.170 Almond 24 49.04 1.127 Mulberry 14 81.21 0.397 Fig 8 101.38 0.181 Raisin 5 94.60 0.121 Date 4 104.25 0.085 Apricot Nil Nil Nil Coconut Nil Nil Nil Chilgoza Nil Nil Nil Loge = 2.303 (constant) T-0 = Time taken by 50% of the adults to emerge. It is clear from the data in Tables 2 and 3 that the order of relative suitability and relative susceptibility of the foods was different. This is so because cashewnut undergoes maximum weight loss though the development period on this food is long. It is the duration of the Table 3 RELATIVE SUSCEPTIBILITY OF 12 DRY FRUITS TO THE ATTACK BY PLODIA INTERPUNCTELLA (H.) Food Suitability index value (a) ‘ Corrected mean % age of weight loss (b) Susceptibility index value (axb> Cashewnut 1.170 42.541 49.773 Pista 1.743 25.748 44.879 Walnut 1 .590 27.544 43.795 Almond 1.127 37.597 42.372 Mulbeiry 0.397 90.768 36.035 Hazelnut 1.393 22.496 31.337 Fig 0.181 70.345 12.732 Raisin 0.121 25.120 3.039 Date 0.085 14.346 1.219 Apricot Nil 6.896 Nil Coconut Nil 0.200 Nil Chilgoza Nil Nil Nil development period that pushes the cashewnut at number 4 in term of suitability index. However, maximum weight loss by cashewnut pushes its susceptibility index to number 1. There is little difference in the two indices of pista and date which occupy the same order in the lists of both indices. The resistance among ten cultivars of pista procured from Tehran Agriculture University was tested on the same pattern as followed for different fruits and susceptibility index was calculated in the same manner The obtained results are give in Tables 4, 5 and 6. Table 4 reveals that varieties kGH’ and ‘WA’ underwent a minimum weight loss of 0.439 gm and 0.465 gm respectively whereas, ‘MO’ underwent a maximum weight loss of 0.660 gm. The progeny from 30 eggs on each food varied from 20 to 28 and the developmental period differed from 22.5 to 26.83 days (Table 5). 64 JOURNAL, BOMBAY NATURAL HISTORY SOCIETY. 97(1 ) APR. 2000 PLODIA INTERPUNCTELLA ASA POTENTIAL PEST OF DRY FRUITS Table 4 WEIGHT LOSS OF TEN VARIETIES OF PISTA DUE TO THE ATTACK OF PLODIA INTERPUNCTELLA (H. ) (based on three observations) Food Initial Weight Final Weight Moisture Content Weight Mean Corrected of food of food M, m2 loss %age mean % age mean (gm) mean (gm) weight loss weight loss ‘MO’ 2 1.340 4.380 4.029 0.660 33.000 32.998 ‘FA’ 2 1.363 4.600 4.400 0.637 31.850 31.849 ‘KA’ 2 1.406 4.420 4.196 0.594 29.700 29.699 ‘JA’ 2 1.431 4.160 3.913 0.569 28.450 28.449 ‘EB’ 2 1.433 4.100 3.698 0.567 28.350 28.348 ‘SH' 2 1.471 4.620 4.554 0.529 26.450 26.449 ‘OH’ 2 1.498 4.680 4.270 0.502 25.100 25.098 ‘RE’ 2 1.524 4.480 3.608 0.476 23.800 23.796 ‘WA’ 2 1.535 4.360 3.452 0.465 23.250 23.246 ‘GH’ 2 1.561 4.520 4.290 0.439 21.950 21.949 MO = Momtaz FA = Fandoghi KA = Kallehghoochi JA = Jabbary EB = Ebrahimi SH = Shasti OH = O’hadi RE = Rezaia WA = Wahedi GH = Gholam Rezaia It is clear from the susceptibility index results (Table 6), that the varieties ‘FA’ and ‘MO’ with the susceptibility index values of 84.750 and 77.578 were the most susceptible foods while the varieties ‘RE’ and ‘WA’ with the index value of 41.1 19 and 45.887, were the least susceptible foods. The result of this study reveal that dry fruits like cashewnut, pista, walnut, almond, hazelnut Table 5 RELATIVE SUITABILITY OF TEN VARIETIES OF PISTA TO THE ATTACK BY PLODIA INTERPUNCTELLA (H.) (based on three replications of 1 0 eggs each) Food Progeny Y (Average) Development period (Todays) Index of suitability L°ScY/T50 ‘FA’ 26 22.50 2.661 ‘O’H’ 28 25.64 2.514 ‘GH’ 26 24.23 2.471 ‘KA’ 27 25.48 2.440 ‘MO’ 25 24.48 2.351 ‘JA’ 26 25.65 2.334 ‘SH’ 25 25.12 2.291 ‘EB’ 22 23.77 2.131 ‘WA’ 23 26.83 1 .974 ‘RE’ 20 26.65 1.728 Loge = 2.303 (constant) T = Time taken by 50% of the adults to emerge. and mulberry are preferred foods of Plodici interpunctella (Hubner) and therefore special care should be taken to save these commodities from the attack of this pest. The damage to fig and raisin is not much and therefore, no special care is required for protection of these two fruits. The remaining three fruits namely apricot, coconut and chilgoza are not attacked by the pest in nature. Some larval feeding is witnessed on apricot and Table 6 RELATIVE SUSCEPTIBILITY OF TEN VARIETIES OF PISTA TO THE ATTACK B.Y PLODIA INTERPUNCTELLA (H.) Food Suitability index value (a) Corrected mean % age of weight loss (b) Susceptibility index value (axb) ‘FA’ 2.-661 31.849 • 84. 750 ‘MO’ 2.351 32.998 77. 578 ‘KA’ 2.440 29.699 72.465 ‘JA’ 2.334 28.449 66. 400 ‘O’H’ 2.514 25.098 63. 096 ‘SH’ 2.291 26.449 60.594 ‘EB’ 2.131 28.348 60. 409 ‘GH’ 2.471 2 1 .949 54. 235 ‘WA’ 1.974 23.246 45. 887 ‘RE’ 1.728 23.796 41.1 19 JOURNAL, BOMBAY NATURAL HISTORY SOCIETY, 97(1), APR. 2000 65 PLOD1A INTERPUNCTELLA ASA POTENTIAL PEST OF DRY FRUITS coconut under experimental conditions, whereas no feeding takes place in case of chilgoza. Acknowledgements The authors are thankful to the Chairman, Refer Almasi, Radmila, Z. Srdic & T. Stojanovic (1987): Influence of food on the fecundity and fertility of Indian meal moth {Plodia interpunctella Hbn.) (Lepidoptera: Phycitidae, ZastBilja 38(4): 309-316. Bell, C.H. (1976a): Effect of cultural factors on the development of four stored product moths. J. Stored Prod. Res. 12: 185-193. Bell, C.H. (1976b): Factors governing the induction of diapause in Ephestia elutella and Plodia interpunctella (H.) (Lepidoptera, Pyralidae). Physiol. Ent. 7:83-91. Bell, C.H. & D.J. Walker ( 1 973): Diapause induction in Ephestia elutella (Hubner) and Plodia interpunctella (Hubner) (Lepidoptera, Pyralidae) with a dawn dusk lighting system. J. Stored Prod. Res. 9: 149-158. Grant, G.G. ( 1 974): Male sex pheromone from the wing glands of the Indian meal moth, Plodia interpunctella (Hbn.) (Lepidoptera : Phycitidae). Experientia. 30: 917-918. Grant, G.G. ( 1 976): Female coyness and receptivity during courtship in Plodia interpunctella (Lepidoptera : Pyralidae). Can. Ent. 108: 975-979. Grant, G.G. &U.E. Brady (1975): Courtship behaviour of phycitid moths (1) comparison of Plodia interpunctella and Cadra cautella and role of male scent glands. Can. J. Zool. S3: 813-826. Hamlin, J.C., W.D. Reed& M.E. Philips (1931): Biology of the Indian meal moth on dried fruits in California. U.S.D.A. Technical Bull No. 242. Hoppe, T, (1981): Food preference - oviposition and development of the Indian meal moth, Plodia interpunctella (H.) on different products and chocolate industry. Z. Angew Entomol. 91(2): 170-179. Jamieson, M.F.S. (1970): A simple tool for calculating loss or gain in weight resulting from a change in the moisture content of produce. Trop. Stored Prod. Int. 19-20. Lecato, G.L. (1976): Yield, development and weight of Cadra cautella (Walk) and Plodia interpunctella (H.) on twenty-one diets derived from natural products. J. Stored Prod. Res. 12:43-41. Department of Zoology, Panjab University, Chandigarh for research facilities. The first author is also thankful to the Government of Iran for providing necessary funds for studying in the Panjab University, Chandigarh. N C E S Mbata, G.N. ( 1 985): Some physical and biological factors affecting oviposition by Plodia interpunctella (Lepidoptera : Phycitidae). Insect. Sci. Appl. 6(5): 597-604. Mbata, G.N. (1987): Studies on the susceptibility of groundnut varieties to infestation by Plodia interpunctella (H.) (Lepidoptera : Pyralidae). J. Stored Prod. Res. 23(1): 57-63. Mbata, G.N. ( 1 990): Suitability of mai/e varieties for the oviposition and development of Plodia interpunctella (H.) (Lepidoptera : Pyralidae). Trop. Pest Manage. 36(2): 122-127. Mbata, G.N. & F.N.C. Osuji ( 1 983): Some aspects of the biology of Plodia interpunctella ( H.) (Lepidoptera : Pyralidae), a pest of stored products in Nigeria. J. Stored Prod. Res. 19(3): 141-151. Mullen, M.A. & R.T. Arbogast (1977): Influence of substrate on oviposition by two species of stored product moths. Environ. Ent. 6(5): 641-642. Myers, J.G. (1928): Report on insect infestation of dried fruits. Empire Marketing Board No. 12. Ono, T. (1981 ): Factors releasing the copulation attempt in three species of Phycitidae. Appl. Entomol. Zool. 16(1): 24-28. Osuji, F.N. (1976): A comparison of the susceptibility of cowpea varieties to infestation by Callosobruchus maculatus (Coleoptera : Bruchidae). Ent. Exp. and Appl. 20: 209-217. Rangaswamy, J.R. (1985): Sex pheromones of stored product insect pests. J. Sci. Jnd. Res. (India). 44(9): 491-500. Simmons, P. (1931): Fig insects in California. U.S.D.A. Circular No. 157. Stein, W. (1990): Investigations about the development of stored product insects at fruits of indigenous trees and shrubs. Anz. Schaedlingskd Pflanzenschutz Umweltschutz. 63(3): 41-46. Williams, G.C. ( 1 964): The life history of the Indian meal moth, Plodia interpunctella (H.) in a warehouse in Britain and on different foods. Ann. Appl. Biol. 53: 459-475. 66 JOURNAL BOMBAY NATURAL HISTORY SOCIETY. 97(1) APR. 2000 FRESHWATER CLADOCERA (CRUSTACEA : BRANCHIOPODA) OF THE ANDAMAN AND NICOBAR ISLANDS1 K. Venkataraman2 ( With one text-figure) Key words: Cladocera, 38 species, Andaman and Nicobar Islands During 1990-92, 221 Cladocera samples were collected from 106 freshwater habitats throughout the Andaman and Nicobar Islands. A total of 38 species belonging to 21 genera of five families were identified, of which 24 were chydorids and 14 nonchydorids. The seven regions, viz. north, middle, south and little Andaman, Carnicobar, Nancowry group and Great Nicobar, were divided into two groups for the present study. All the seven stations were compared by the Sorensen index of similarity, and Koch index of biotal dispersity. The indices are generally high, reflecting the small number of species involved and their wide distribution, but two groups of stations are easily discernible. The indices for the Nicobar group of islands are somewhat lower, being influenced by the erratic occurrence of eurytopic species such as Moina micrura , Ceriodaphnia cornuta and Macrothrix spinosa , but they are clearly interrelated. Introduction The Andaman and Nicobars consist of over 550 islands, including several archipelagoes with a land area of 8,293 sq km. Being oceanic islands, they have hilly terrain and virgin forests, free flowing streams and cavities where water logging takes place throughout the monsoon. Irrigation reservoirs or lakes are absent, except for a few very small dams used mainly for drinking water (Danikari dam and Dilthaman tank, Port Blair). A few perennial water bodies, and many temporary cavities and rice fields are the main wetlands, where this study was conducted. The Cladocera are dominant micro- crustaceans in the freshwater habitats of the Andaman and Nicobar islands, but they are not known taxonomically and ecologically as compared to those in the surrounding regions, such as the Indian mainland (Venkataraman, 1983; 1992a; Michael and Sharma, 1988; Venkataraman and Das, 1993), Sri Lanka 'Accepted November, 1 997 Zoological Survey of India 100, Santhome High Road, Chennai 600 028, Tamil Nadu, India. (Rajapaksha and Fernando, 1987), Malaysia (Idris, 1983) and the Philippines (Marnaril, 1977). Except for Venkataraman ( 1 99 1 , 1 992b, c), no worker has studied the freshwater bodies of Andaman and Nicobar Islands. Hence, this study on the occurrence of Cladocera was undertaken in the freshwater habitats of Andaman and Nicobar Islands. Material and Methods During 1990-92, 221 samples of Cladocera were collected from 106 freshwater habitats throughout the North (NA), Middle (MA) and South Andaman (SA), (Diglipur, Mayabunder, Rangat, Kadamthala, Port Blair, Havelock, Ross Islands and Little Andaman), Carnicobar (CN), Nancowry Group (NG) and Great Nicobar (GN) of Nicobar Islands (Fig. 1). Samples were collected from ponds, marshes, reservoirs, rice fields, dams, streams and rainwater pools, using a plankton net of 45 cm diameter, with circular mouth. The samples were usually collected in shallow water, among vegetation and in clear water. The net was dragged close to the bottom; excessive stirring of the mud was avoided. This technique gave a qualitative sample of shallow JOURNAL, BOMBAY NATURAL HISTORY SOCIETY. 97(1), APR. 2000 67 FRESH W A TER CLADOCERA OF THE ANDAMAN AND NICOBAR ISLANDS water invertebrates living on the substratum, among vegetation and in the water column as in rice fields or marshes. The normal annual rainfall was 3,180 mm, mean max. temp. 29.98 °C and the mean min. temp. 23. 1 3 °C. The mean relative humidity was 79%. Descriptions of Some Rare Species Brief descriptions of five rare species reported in the present study are given below. All the other 33 species recorded in this study have been described by Michael and Sharma (1988) from the Indian mainland, Malaysia (Idris, 1983) and Sri Lanka (Rajapaksha and Fernando, 1987). Family Sididae Diaphanosoma volzi Stingelin 1905 Material examined: 5 females from Bomila creek marsh, Little Andaman. Female: Body size 0.73 mm. Head rounded and small, eye relatively large. Valves straight on ventral margin, duplicature forming a wide angle, posteroventral comer rounded, without denticles except for a long spine on the posterior margin. Postabdomen with three long and sharply pointed basal spines. Remarks: Very rare. The material agrees with the description often mentioned under the name D. aspinoswn by Chiang (1956) from China, and by Idris (1983) from Malaysia. Family Macrothricidae Guernella raphalis Richard 1892 Material examined: 3 females from Hut Bay nalla, Little Andaman, several females from roadside ponds in Wandoor, Port Blair. Female: Body size 0.41 mm. Carapace 68 JOURNAL BOMBAY NATURAL HISTORY SOCIETY, 97(1). APR. 2000 FRESHWA TER CLADOCERA OF THE ANDAMAN AND NICOBAR ISLANDS slightly oval. Head rounded anteriorly and concave ventrally; eye large, ocellus at the apex of rostrum. Antennule short and broad with lateral setae. Valves with polygonal reticulations, broadly rounded distally, ventral margin rounded and serrated. Claw short without basal spine. Macrothrix laticornis (Jurine 1820) Material examined: 6 females from Kadamthala fish pond (NA). Female: Body size 0.48 mm. Head rounded; rostrum small with two antennules implanted with a bunch of subapical long spines and four rows of dorsal spines. Ventral margin with movable spines. Postabdomen thick and swollen, with numerous fine spines. Family Chydoridae Pleuroxus denticulatus Birge 1879 Material examined: 5 females from Yatrik pond, 7 females from Schoolline pond, 22 females from Coast Guard pond. Female: Body size 0.42 mm. Shape broadly oval, with striated carapace. Posteroventral comer with 2-4 denticles. Rostrum long and pointed. Ocellus situated closer to the eye than to apex of rostrum. Postabdomen with 14-16 denticles with two basal spines on the claw. Alona cf. dentifera (Sars 1901) Material examined: 9 females from Mumgan temple pond, 6 females from Schoolline pond, Port Blair (SA). Female: Body size 0.45 mm. Valves with longitudinal lines. Posteroventral corner rounded, with three denticles. Ocellus slightly smaller than eye, situated half-way between eye and tip of rostrum. Labrum rounded anteriorly, slightly pointed ventrally. Postabdomen with prominent preanal and postanal corner, with 1 1 groups of denticles. Claw long, with a long basal spine. Remarks: Rare. New record to India. Idris (1983) shifted this species from the genus Alonella to Alona. More studies are required to confirm the identity of this species. Results and Discussion A total of 38 species belonging to 2 1 genera of 5 families were identified in the 22 1 samples collected from different habitats during 1989-1991, of which 24 were chydorids and 14 nonchydorids. Of all the samples, only 10 contained no cladocerans. There were great differences between the islands in the number of cladocerans collected (Table 1). As in the Northeast (Venkataraman, 1994, 1995), as well as Tamil Nadu and Rajasthan (Venkataraman, 1983, 1992a), cladocerans of the Andaman and Nicobar Islands are a mixture of tropical and temperate species (Table 2). Ceriodaphnia cornuta , Moina micrura and Diaphanosoma excisum are considered to be typically tropical species widely distributed from the northernmost tip Diglipur, to the other end, Great Nicobar. Diaphanosoma volzi , Macrothrix laticornis , Pleuroxus denticulatus , Chydorus pubescens , Alona cf. dentifera and Leydigia acanthocercoides, which are considered to be temperate in origin, occur in Andaman and Nicobar Is. (Table 1). Cladoceran hatching and growth rate is controlled by temperature, which ranged from 29-32 °C (Table 3) in the study. The pH range of these wetlands was narrow, 7.25 to 8.90. Previous workers Bayly (1963), Moitra and Bhattacharya (1965) and Chengalath (1982) showed that Cladocera and other freshwater, zooplankton populations vary inversely with pH. However, the present study does not show any such significant variation. The study areas receive monsoon rain from March through October. This continuous rainfall dilutes the ionic strength and nutrient levels of the water, which in turn may affect the proliferation of cladoceran population. It also increases the oxygen content of the water along JOURNAL . BOMBAY NATURAL HISTORY SOCIETY. 97(1). APR. 2000 69 FRESH W A TER CL A DOCERA OF THE A NDA MAN AND N I CO BA R IS LA NDS Table 1 OCCURRENCE OF CLADOCERA (IN NUMBER OF SAMPLES) IN DIFFERENT REGIONS OF ANDAMAN AND NICOBAR ISLANDS (TOTAL NUMBER OF SAMPLES COLLECTED IS GIVEN IN PARENTHESIS) Andaman Nicobar SI. No. Cladocera species North Andaman (20) Middle Andaman (30) South Andaman (96) Little Andaman (44) Carnicobar (10) Nancowry Group (6) Great Nicobar (15) Family Sididae 1 . Pseudosida bidentata 2 - - - - - - 2. Latonopsis australis - - 3 1 - - 1 3. Diaphanosoma sarsi 3 4 10 3 - - 2 4. Diaphanosoma excisum 1 3 28 - - - - 5. Diaphanosoma volzi - - - 1 - - - Family Daphniidae 6. Ceriodaphnia cornuta 5 7 18 6 3 1 3 7. Scapholeberis kingi - 1 7 - - - - Family Moinidae 8. Moina micrura 6 13 19 5 - - 1 9. Moinodaphnia macleayi - - 5 - - - - Family Macrothricidae 1 0 . Macrothrix spinosa 1 1 9 3 - 1 3 1 1 . Macrothrix laticornis - 1 - - - - 1 2. Echinisca triserialis - - 18 2 - 1 - 1 3 . Ilyocryptus spinifer - 1 7 3 - - - 14. Guernella raphalis - - 2 1 - ■ _ Family Chydoridae Subfamily Chydorinae 15. Pleuroxus similis 4 1 6. Pleuroxus denticulatus - - 6 - - - - - 17. Chydorus ventricosus - 1 19 3 - - - 18. Chydorus reticulatus 7 13 22 5 - - - 19. Chydorus eurynotus 5 6 9 2 - 1 1 20. Chydorus parvus - 1 6 - - - ■ 2 1 . Chydorus barroisi - - 17 - - - - 2 2 . Chydorus pubes cens - 1 - - - - - 23. Dadaya macrops 1 2 8 2 - - 1 24. Dunhevedia crassa - 2 12 3 - - 1 25. Dunhevedia serrata - 1 7 - - - * Subfamily Aloninae 26. A Iona monacantha 1 - 8 - - - ■ 27 . A Iona cf. dentifera - - 3 - - - - 28. Alona pulchella - 6 12 4 - - - 29. Alona guttata - 2 - - - ■ ■ 30. Alona davidi - - 18 5 - - • 31. Alona karua 2 5 12 3 - - - 32. Alona verrucosa - - 6 - - - - 3 3 . Oxyurella sinhalensis ' - - 10 - - - - 34. Kurzia longirostris 1 - 7 2 - - 1 35. Euryalona orientalis - - 4 - - - ■ 36. Notalona globulosa 2 - 7 - - " 3 7 . Leydigia acanthocercoides 1 1 3 - - ■ 1 38. Leydigia australis - - 4 - ■ ■ ' Total number of species 14 20 32 19 1 4 10 70 JOURNAL, BOMBAY NATURAL HISTORY SOCIETY, 97(1). APR. 2000 FRESHWATER CLADOCERA OF THE ANDAMAN AND NICOBAR ISLANDS Table 2 OCCURRENCE OF SPECIES OF CLADOCERA IN DIFFERENT STATES OF INDIA SI. No. Name of the family Number of species of Cladocera Tamil Nadu Rajasthan West Bengal Tripura Andaman and Nicobar India (total) 1. Sididae 5 5 5 5 5 6 2. Daphniidae 12 12 9 7 2 17 3. Moinidae 2 4 3 2 2 5 4. Bosminidae - 1 2 2 - 2 5. Macrothricidae 4 8 6 4 5 8 6. Chydoridae 23 24 32 29 24 47 Total 46 54 57 49 38 85 with the nutrient level in the wetlands of Andaman (Table 4). The seven regions viz. north, middle, south and little Andaman, Carnicobar, Nancowry group and Great Nicobar have been divided into two groups for the puipose of the present study. The four northern regions known as the Andaman group have fourteen or more species Table 3 PHYSICOCHEMICAL PARAMETERS OF THE FRESHWATER PONDS AND LAKES STUDIED IN ANDAMAN AND NICOBAR ISLANDS. SI. No. Name of the pond Date pH Surface water Temp. °C Conductivity mmhos O, mg/I Transparency cm 1. Mayabundar (NA) 23.01.91 5.5 - 8.70 '* - 2. Schoolline pond (SA) 28.04.90 - - 3.80 - - 3. Dhobi pond (SA) 21.05.90 8.53 32.3 1.30 4.10 30 4. Yatrikpond I (SA) 21.05.90 7.95 33.0 2.00 8.10 75 5. Bay Island Hotel Pond (SA) 25.05.90 8.70 31.2 4.00 10.42 50 6. Murugankoil pond (SA) 02.06.90 7.09 29.9 2.40 6.05 7. Murugankoil pond (SA) 02.07.90 7.17 30.8 2.00 - - 8. Dhobi pond (SA) 02.07.90 7.59 30.2 7.30 - - 9. Murugankoil pond (SA) 09.07.90 7.78 31.2 6.90 - 10. Murugankoil pond (SA) 17.7.90 7.29 29.5 7.50 5.70 - 11. Murugankoil pond (SA) 18.07.90 7.04 30.4 8.00 5.90 ■ - 12. Murugankoil pond (SA) 19.07.90 7.26 30.2 3.00 5.20 - 13. Yatrikpond(SA) 27.7.90 8.94 31.5 1.90 6/00 - 14. Dhobi pond (SA) 18.07.90 8.50 31.5 6.60 10.30 - 15. Dhobi pond (SA) 19.07.90 7.34 29.7 5.70 5.30 - 16. Coastguard Pond (SA) 19.07.90 8.36 30.1 2.00 6.20 - 17. Schoolline pond (SA) 08.08.90 7.35 31.4 0.11 7.78 - 18. Yatrik pond (SA) 08.10.90 7.26 30.8 2.00 8.00 - 19. Havelock pond I (SA) 23.01.91 5.50 - 8.70 - - JOURNAL. BOMBAY NATURAL HISTORY SOCIETY. 97(1), APR. 2000 71 FRESHWA TER CLADOCERA OF THE ANDAMAN AND NICOBAR ISLANDS Table 4 PHYSICOCHEMICAL PARAMETERS OF THE SURFACE RUN OFF WATER BEFORE AND AFTER RAIN IN THE CANALS OF PORT BLAIR DURING MAY 1 990 SI. Canals of Port Blair Conductivity in pH 02mg/l Total solid Dissolved solids No. mmhos mg/1 mg/1 Before Rain After Rain Before Rain After Rain Before Rain After . Rain Before Rain After Rain Before Rain After Rain 1. Murugan temple 4.6 6.9 7.40 7.02 6.54 6.43 120 3800 500 1200 2. Shadipur canal 2.1 4.1 7.61 7.48 6.34 6.63 340 2000 500 1400 3. Phoenix Bay canal 8.0 11.3 7.51 7.49 4.27 4.73 3300 5700 1000 2000 4. Anarkali canal 9.3 8.4 7.88 7.64 6.19 6.53 770 21800 700 2200 5. Megapod Nest canal 3.3 3.4 7.68 7.70 5.88 6.36 1250 1300 200 900 Mean 6.0 8.37 7.64 7.49 5.97 6.25 683 6017 533 3000 Deviation 2.57 4.76 0.17 0.25 0.86 0,76 577 7910 287 3609 of Cladocera each, while the southern three regions known as the Nicobar group have only one to ten species. The seven stations have been compared by the Sorensen index of similarity. This was calculated for each combination of stations according to the following equation (Sorensen, 1948): S=2c/a+b x 100, where ‘c’ is the number of species common to both associations, ‘a’ the number of species in one association and ‘b’ the number of species in the other association. The results for 21 pairs of stations are given in Table 5. The indices are generally high, reflecting the small number of species involved and their wide distribution, but the two groups of stations are easily discernible. Andaman group (4 stations) have highly interrelated indices. Those for the Nicobar group are lower, being influenced by the erratic occurrence of eurytopic species such as Moina micrura , Ceriodaphnia cornuta and Macrothrix spinosa, but they are clearly interrelated. Andaman Islands closely resemble each other, the Great Nicobar closely resembles Nancowry group, whereas Camicobar is unique (Table 5). Koch (1957) has devised an index ofbiotal dispersity (IBD) which can be used to assess the wide dispersity of species between islands. IBD =T-S/S(n-1 ) x 100, where ‘T’ is the arithmetical sum of species living in each ‘iT compared associations and ‘S’ is the total list of species in ‘iT compared associations. If each station had a completely different set of species, ‘S’ should equal ‘T’ and the IBD would be 0%. If each station had an identical set of species, ‘T’ would equal n x S and the IBD would be 100%. When the Koch index for all seven stations was calculated, the resulting IBD was 27, but when separate indices were calculated for the Andaman and Nicobar groups, there was an increase in the IBD for the former (40) and a decrease for the latter (18). The large increase in IBD when the Andaman group were considered separately indicates that these Table 5 SORENSEN INDICES FOR CLADOCERA FROM SEVEN DIFFERENT ISLAND GROUPS OF ANDAMAN AND NICOBAR. 1 2 3 4 5 6 7 1 _ 53 59 53 13 33 67 2 53 - 64 60 10 25 33 3 59 64 - 68 7 24 45 4 53 60 68 - 10 58 60 5 13 10 7 10 - 40 18 6 33 25 24 58 40 - 43 7 67 33 45 60 18 43 - 1 - North Andaman; 2- Middle Andaman; 3 - South Andaman; 4 - Little Andaman; 5 - Camicobar; 6 - Nancowry Group; 7 - Great Nicobar. 72 JOURNAL, BOMBAY NATURAL HISTORY SOCIETY, 97(1), APR. 2000 FRESHWA TER CLADOCERA OF THE ANDAMAN AND NICOBAR ISLANDS regions resemble each other in Cladocera fauna much more than they resemble the Nicobar group. This agrees well with the Sorensen indices. Acknowledgements I thank the Director, ZSI, Calcutta, Officer- in-charge Andaman and Nicobar Regional Refer Bayly, I.A.E. (1963): Reversed diurnal vertical migration of planktonic Crustacea in inland waters of low hydrogen ion concentration, Nature , 200 : 704-705. Chengalath. R. (1982): A faunistic and ecological survey of the littoral Cladocera of Canada, Can. J. Zool. 60: 2668-2682. Chiang, S.C. (1956): Some species of Diaphanosoma (Cladocera) from Wuchang, China (in Chinese with English Summary). Acta Hydrobiologia Sinica, 2: 31-2-341. - Idris, B.A.G. (1983): Freshwater zooplankton of Malaysia (Crustacea : Cladocera), Penerbit Universiti Pertanian Malaysia, 153 pp. Koch, L.P. (1957): Index ofbiotal dispersity. Ecology, 38: 145-148. Mamaril, A.C. (1977): Freshwater zooplankton of Philippines (Rotifera, Cladocera and Copepoda) M.Sc. thesis, University of Waterloo, Canada, 151 pp. Michael R.G. & B.K. Sharma (1988): Fauna of India, Indian Cladocera (Crustacea : Branchiopoda : Cladocera) Ed. Director, Zoological Survey of India, 262 pp. Moitra, J.K. & B.K. Bhattacharya (1965): Some hydrological factors affecting plankton production in a fish-pond in Kalyani, West Bengal, India, Ichthyologia, 4: 8-12. Rajapaksa, R. & C.H. Fernando (1987): Redescription and assignment of Alona globulosa Dady 1 898 to Station and Marine Biological Station for facilities for preparing this paper. I also thank Dr. H.S. Mehta, Shri Bulganin Mitra, Dr. Sanjeev Kumar, Shri P.T. Rajan, Shri Sukla, Shri Ponnusamy, Shri Deivaprakasam and Shri Selvaraj of Andaman and Nicobar Regional Station for their valuable help in collecting the specimens. ENC ES new genus Notoalona and a description of Notoalona freyi sp. nov., Hycirobiologia. 144: 131-153. Sorensen, T. (1948): A method of establishing group of equal amplitude in plant sociology based on similarity of species content and its application to analysis of the vegetation on Danish commons, Biol. Skr. 5(4): 1-34. Venkataraman, K. (1983): Taxonomy and Ecology of Cladocera of southern Tamil Nadu. Ph.D. thesis, Madurai Kamaraj University, Madurai, 190 pp. Venkataraman, K. (1991): Freshwater Cladocera of Little Andaman, J. Andaman Sci. Assoc. 6: 60-62. Venkataraman, K. (1992a): I. Cladocera of Keoladeo National Park, Bharatpur and its environs, J. Bombay nat. Hist. Soc.., 89: 1 7-26. Venkataraman, K. (1992b): Freshwater Cladocera of Andaman, J. Andaman Sci. Assoc. 8: 133-137. Venkataraman, K. (1992c): Occurrence of male Cladoceran of Moinodaphnia macleayi (King) in oriental region, / Andaman. Sci. Assoc. 8: 179-180. Venkataraman, K. (1994): Cladocera In: State Fauna Series 3: Fauna of West Bengal, Part 10: 1-36. Venkataraman, K. (1995): Freshwater Cladocera of Tripura State, North Eastern India,/. Andaman Sci. Assoc. II: 15-20. Venkataraman, K. & S.R. Das (1993): Freshwater Cladocera (Crustacea : Branchiopoda) of southern West Bengal, /. Andaman Sci. Assoc.. 9: 1 9-24. JOURNAL. BOMBAY NATURAL HISTORY SOCIETY. 97(1), APR. 2000 73 LONGICORN BEETLES (CERAMBYCINAE, PRIONINAE : CERAMBYCIDAE) OF BUXA TIGER RESERVE, JALPAIGURI, WEST BENGAL1 Dinendra Raychaudhuri2 and Sumana Saha C With twelve text-figures ) Key words: Taxonomy, Cerambycidae, Cerambycinae, Prioninae, Buxa Tiger Reserve, West Bengal The paper deals with the taxonomy of 1 2 species of Cerambycidae (Cerambycinae and Prioninae) of Buxa Tiger Reserve. Of these, 7 species Ceresium leucosticticum White, C. rufum Lameere, Hoplocerambyx spinicornis Newman, Macrotoma (Zooblax) spinosa (Fabricius), Tetraommatus filiformis Perroud, Thranius simplex Gahan and Xoanodera regular is Gahan, are new records from the state of West Bengal. The species have been described and suitably illustrated. Identification keys are provided wherever necessary. Introduction Family Cerambycidae refers to the longhorn beetles. These coleopterans are wood borers in their larval stages, for which they are extremely important in any forest ecosystem. Because of their great economic importance, these beetles have received serious attention. Up to 1200 species of cerambycids are reported from the Indian region, largely dominated by the Lamiinae (Beeson, 1961). The systematics, biology and ecology of these beetles have been worked out. Khan and Maiti ( 1 983) while dealing with the biotaxonomy, biology and ecology of some of these borers have reviewed the works of others, e.g. Stebbing, Beeson, Beeson and Bhatia, Husain and Khan, Bhasin and Roonwal, Bhasin et al., and Dutt. Basak and Biswas (1993) have remarked “our present state of knowledge of longicom beetles of the state of Orissa is very incomplete and fragmentary”, and “no comprehensive work on the longicom beetles from Orissa is available”. They, however, listed 32 species belonging to 27 genera under 3 subfamilies as the cerambycid fauna of Orissa. Though they indicated the distribution of some of these species in West Bengal, recent State 'Accepted June, 1999 :Entomology Laboratory, Department of Zoology, University of Calcutta, 35, Ballygunge Circular Road, Calcutta 700019, West Bengal, India. Fauna Series 3: Fauna of West Bengal Pt 6A, 6B (Insecta : Coleoptera), 1995-96 Z.S.I. did not include Cerambycidae. Our survey of Buxa forest, presently known as Buxa Tiger Reserve ( Jalpaigun, West Bengal) during 1994-97 revealed the existence of 12 species of longhorn beetles (Cerambycinae; Prioninae) belonging to 1 1 genera. Raychaudhuri (1996) had reported 10 species belonging to 10 genera of the same subfamilies from the forest. Besides, we have several species of lamiids in our collection. We now present the taxonomic details of the beetles belonging to the subfamilies Cerambycinae and Prioninae. Incidentally, Basak and Biswas (1993) have not presented any taxonomic key or detailed description of the 32 listed species. Such a taxonomic treatise is necessary for India, as several decades have elapsed since the publication of Gahan (1906). This paper details the morphology of each species, together with keys, even of the higher categories. All the species have been illustrated. Ceresium leucosticticum White, C. rufum Lameere, Hoplocerambyx spinicornis Newman, Macrotoma spinosa (Fabricius), Tetraommatus filiformis Perroud, Thranius simplex Gahan and Xoanodera regularis Gahan appear to be new records from the state of West Bengal. All the reported species are at present in the collection of Entomology Laboratory, Department of Zoology, University of Calcutta. 74 JOURNAL, BOMBAY NATURAL HISTORY SOCIETY, 97(1), APR. 2000 LONGICORN BEETLES OF BUXA TIGER RESERVE Material and Methods Insect samples have been collected and preserved for further study following the recommendations of Zoological Survey of India, Calcutta (Ghosh and Sengupta, 1982). Abbreviations used: BG - Bhutanghat — B.T.R. - Buxa Tiger Reserve DM - Damanpur EL - Length of elytra EW * Humeral width of elytra FA - Apical width of frons FL - Length of frons A. HW - Maximum width of head JY - Jayanti a. LT - Light trap NL - Newland NM - Nimati al PA - Apical width of pronotum PB - Basal width of pronotum PK - Phaskhawa — PL - Length of pronotum PN - Panbari PW - Maximum width of pronotum — RB - Rajabhatkhawa RM - Raimatang SB - South Bholka — SR - South Raydak TG - Tashigaon B. Taxonomy Key to Subfamilies, Tribes and Genera 1. Prothorax marginate at sides, sometimes entire, more frequently dentate or spinose; fore coxae strongly transverse; antennae usually inserted close to mandibular base; mesonotum without al . stridulatory area (except in Philus ); vein Cu2 usually present; vein Al with a large subelliptical cell Prioninae A. Episterna of metathorax with posteriorly converging sides, narrowly truncate or obtusely pointed at apex; intercoxal process of prosternum arched; lateral margins of prothorax unarmed or with 1-3 spines or teeth; antennal joints spinose; 1st antennal joint short Megopidini, Megopis Serville Episterna of metathorax parallel-sided for greater part of their length, broadly truncate behind; intercoxal process of prosternum flat and horizontal; lateral margins of prothorax crenulate, denticulate or spinulose; antennal joints not spinose, if at all with short spines; 1 st antennal joint long or moderately long .... Macrotomini, Macrotoma Serville Prothorax emarginate at sides; fore coxae rarely strongly transverse; antennae inserted at some distance from base of mandibles; mesonotum generally with stridulatory area; veins Cu2 and branch of Cul usually absent; vein Al mostly without any cell Cerambycinae Intercoxal process of prosternum not or weakly dilated at apex a Ligula corneous; antennae never ciliated but may have long pubescence; vein Cu2 absent Oemini Head flat between antennae; 1st coxae contiguous; antennae never spinose or dentate Tetraommatus Perroud Head raised forming a ridge, broadly concave between antennae; 1st coxae separate; antennae dentate Xystfocera Serville Ligula membranous; antennae ciliated; either vein Cu2 or posterior branch of Cul absent .. Hesperophanini, Stromatium Serville Intercoxal process of prosternum distinctly dilated at apex B Acetabula of fore coxae closed or nearly closed posteriorly, rarely angulated on outer side .... i Metasternum with scent-pores; acetabula of middle coxae extended to epimera Callichromini, Aiiubis Thomson Metasternum without scent-pores; acetabula of middle coxae open to epimera Cerambycini Pronotum transversely irregularly wrinkled with broken ridges; elytra with a spine at sutural apex; 1 st joint of hind tarsus nearly as long as the next two united Hoplocerambyx Thomson Pronotum without ridge, instead either transversely grooved near base and apex or with variable number of sharp, straight, longitudinal costae; elytra without spine at sutural apex; 1st joint of hind tarsus shorter than next two united Xoanodera Pascoe JOURNAL, BOMBAY NATURAL HISTORY SOCIETY, 97(1), APR. 2000 75 LONG1CORN BEETLES OF B UXA TIGER RESER VE Fig. 1 : Anubis inermis (White): A. Whole body, B. Antenna, C. Hind leg — Acetabula of fore coxae open posteriorly, angulated on outer side ii ii Eyes coarsely faceted Callidiopsini, Ceresium Newman — Eyes finely faceted iii iii. Elytra dehiscent posteriorly, acuminate at apex; front coxae prominent, contiguous; prothorax parallel-sided Thranini, Thranius Pascoe — Elytra neither dehiscent nor acuminate at apex; front coxae not prominent, globular and not contiguous; prothorax with sides weakly to distinctly rounded Clytini, Xylotrechus Chevrolat Subfamily 1 : Cerambycinae Tribe: Callichromini Genus: Anubis Thomson Anubis , Thomson 1864, Syst. Ceramb. : 177. Type-species: Anubis clavicornis Fabricius Anubis inermis (White) (Fig. 1) Polyzonus inermis White 1853, Cat. Coleopt. B.M., Longic.: 171. Male: Head, antennal segment I, pro- notum, scutellum, elytra at base and apex chalybeate blue, pronotal disc and elytra violaceous, elytra with a pair of yellow transverse bands, one just above the middle and the other just below the middle, antennae and legs blue- black; body beneath clothed with silvery grey pubescence. Head at base narrower than pronotum, much narrowed beyond eyes, densely and coarsely punctate; vertex flat; frons midlongi- tudinally sulcate between the antennae; clypeus broad, flat, transverse; HW/PA 1.11; FA/FL 0.46; 76 JOURNAL, BOMBAY NATURAL HISTORY SOCIETY, 97(1). APR. 2000 L ON G I CORN BEETLES OF B UXA TIGER RESER VE mandibles robust, weakly curved, strongly dentate; genae long, midlongitudinally faintly raised, sloped on either side; eyes deeply emarginate, extending beyond antennal support. Antennae 1 1 -segmented, a little longer than body, gradually swollen apically, segment III longest, last segment longer than the penultimate, obtuse at apex. Pronotum ovate cylindrical, longer than broad, lateral margin rounded, basally nearly straight, apically a little constricted, densely and coarsely punctate; PL/ PA 1.52, PL/PW 1.0, PB/PA 1.0, PB/EW 3.09. Scutellum small, triangular and obtuse, finely and rather densely punctate. Elytra long, parallel-sided, rounded at apex, finely and very densely punctate, those near base large and distinct, a pair of costae evidently running from the middle of the base almost to apex; EL/EW 18.18; metasternum midlongitudinally sulcate and free of pubescence; abdominal venter at sides with more dense pubescence. Legs moderately long, femora pedunculate, gradually subclavate, densely punctate, joint 1 of hind tarsi longer than 2+3. Body length: 13-17 mm. Material examined: 1 male, PK, B.T.R., Jalpaiguri, West Bengal, 9.V.1994. Distribution: india: Sikkim, West Bengal; Laos; Malaysia; Myanmar; Pakistan; South China; Thailand (Gahan, 1906; Gressitt and Rondon, 1970). Tribe: Callidiopsini Genus: Ceresium Newman Ceresium Newman, 1842. Entomologist, i: 322. Type-species: Ceresium raripilum Newman KEY TO SPECIES 1. Elytra with yellow-white pubescence, 1st pair oblique, near scutellar apex, 2nd round, transverse, marginal, 3rd oblique, close to suture and 4th comma-shaped, transverse, marginal near apex; head without any pubescence near base; body brown-black; antennae twice as long as body leucosticticum White — Elytra without any such pubescence; head with yellow-white pubescence between eyes near base; body reddish brown; antennae a little shorter than body rufum Lameere Ceresium leucosticticum White (Fig. 2) Ceresium leucosticticum White, 1855, Cat. Col. B.M. Longic. 2: 245. Male: Brown black, with elytra centro- medially reddish, pronotum laterally with 2 pairs of yellow white pubescence, 1st pair near apex, other pair almost basal, scutellum with similar pubescence, elytra also with similar pubescence arranged thus: 1st pair near scutellar apex, rather oblique, broad distally, 2nd pair at basal 1/3, transverse, circular, placed marginally, 3rd almost at midlength, near the suture, oblique, directed towards apex, 4th near the turning of elytra, transverse, marginal, comma-shaped, directed towards apex, eyes at inner margin with semilunar band of similar pubescence; antennae reddish brown, with faint pubescence; legs reddish brown with femora apically darker; body ventrally red brown to dark brown, with yellow white pubescence laterally. Head a little narrower than pronotum, densely punctate, concave between antennae; frons midlongitudinally sulcate; vertex sloped towards eyes, anteriorly truncate; clypeus transverse, band- like; HW/PA 1.16; FA/FL 0.9; eyes emarginate. Antennae 1 1 -segmented, slender, twice as long as body, 1st joint closely punctate, little longer than 3rd, nearly equal to 4th, 5th and following segments longer, 10th twice as long as 11th. Pronotum elongately rectangular, longer than wide, marginally rounded, medially broad, densely and coarsely punctate, clothed with short hairs; PL/PA 1.15, PL/PW 1.15, PB/PA 0.93, PB/EW 1.40. Scutellum small, obtuse. Elytra parallel-sided, naiTOwed just before the truncate apex, densely JOURNAL, BOMBAY NATURAL HISTORY SOCIETY, 97(1), APR. 2000 77 LONGICORN BEETLES OF BUXA TIGER RESERVE Fig. 2: Ceresium leucosticticum White: A. Whole body, B. Antenna, C. Hind leg punctate, clothed with short hairs; EL/EW 5.35; prostemum truncate, punctate; metastemum plate- like, midlongitudinally with a black streak. Legs moderately long, femora basally pedunculate, apically clavate, hind legs extending much beyond the abdomen. Body length: 12 mm. Material examined: 2 males, RB(LT), B.T.R., Jalpaiguri, West Bengal, 22. v. 1995, 25. v. 1995. Distribution: india : Assam (Gahan, 1906), West Bengal; Indonesia; Laos; Myanmar; South China; Thailand (Gahan, 1906; Gressitt and Rondon, 1970). Ceresium rufum Lameere (Fig. 3) Ceresium rufum Lameere, 1890, Ann. Soc. Ent. Beige, 34, C.R. : cc 11. Male: Reddish brown, densely pubescent, head with 2 white semilunar bands of pubescence, pronotum with such pubescence submarginally, anterior ones round, posterior ones rather elongate and longitudinal, extending a little on the basal margin, scutellum with similar pubescence, elytra without any such, antennae reddish brown, 9th joint onwards much darker, legs reddish brown, body ventrally red brown with white pubescence laterally. Head narrower than pronotum, anteriorly narrowed, densely punctate; frons concave, midlongitudinally sulcate, anteriorly subquadrate; vertex sloped towards eyes, anteriorly truncate; clypeus transverse, band-like, truncate; HW/PA 1.12, FA/FL 0.69; eyes emarginate. Antennae 1 1 -segmented, slender, a little shorter than body; 4th joint much shorter than any of the succeeding joints, hairy beneath. Pronotum elongately rectangular, longer than wide, marginally rounded, broad medially; pronotal disc with dense, coarse, transverse rugosities, clothed with short hairs; PL/PA 1 .25, PL/PW 1.00, PB/PA 1.04, PB/EW 2.06. Scutellum small and obtuse. Elytra parallel- sided, narrowed just before the truncate apex, strongly punctate, those towards apex feeble and scanty, clothed with short, dense hairs; EL/EW 8.31; prosternum truncate, punctate; metasternum plate-like, midlongitudinally with a black streak. Legs moderately long, clothed with rather long pubescence, femora basally pedunculate, apically clavate, hind legs extending much beyond the abdomen. 78 JOURNAL, BOMBAY NATURAL HISTORY SOCIETY, 97(1), APR. 2000 L ONG1CORN BEE TEES OF BOX A TIGER RESER VE Fig. 3: Ceresium rufum Lameere: A. Whole body, B. Antenna, C. Hind leg Body length: 13 mm. Material examined: 1 male, SB(LT), B.T.R., Jalpaiguri, West Bengal, 21.V.1997. Distribution: india: Kunbur (Gahan, 1906), West Bengal; Myanmar (Beeson, 1961). Tribe: Cerambycini Genus: Hoplocerambyx Thomson Hoplocerambyx Thomson, 1 864, Syst. Ceramb.: 229. Type-species: Hammciticherus spinicornis Newman Hoplocerambyx spinicornis (Newman) (Fig. 4) Hammciticherus spinicornis Newman, 1842, Entomologist, 1: 245. Male: Pitch brown, ventrally more reddish; head, pronotum, antennae, legs and underside with fine grey pubescence, elytra more densely covered with red-ochraceous silky pubescence, faintly banded light and dark. Head strongly exserted, apically finely punctate, basally wrinkled, with the vertex deeply grooved medially, extending between the eyes, continuing as a shallow groove between antennal supports, carinate on either side; frons oblique with a fovea on each side; clypeus apically sinuate, basal submedian area bi-tuberculate, sloping towards the frontal fovea with a few long, grey hairs at the corners; genae long; slightly shorter than width of pronotum; HW/PA 1.22; FA/FL 0.50; mandibles longer than in female, straight at base; eyes deeply emarginate. not extending beyond the antennal supports; gula with 3 strong transverse ridges. Antennae 11- segmented, 1/5 to 1/3 longer than body, faintly pubescent, 1st to 7th segments sparsely but strongly punctate, 3rd segment onwards spinose, 8th segment onwards gradually shortened and almost weakly so on the last segment, flattened or slightly canaliculate above. Pronotum a little longer than broad, constricted in front, rounded at the sides between the anterior constriction and the base; the disc with a slightly raised oblong space in the middle, the rest of the surface with deep, irregular, transverse wrinkles, with the JOURNAL, BOMBAY NATURAL HISTORY SOCIETY, 97(1), APR. 2000 79 LONGICORN BEETLES OF BUXA TIGER RESERVE Fig. 4: Hoplocerambyx spinicorn is Newman: A. Whole body, B. Antenna, C. Hind leg ridges more or less broken and convolute towards the sides, at base with 2 transverse ridges continuing up to the margin; PL/PA 1.22, PL/ PW 0.79, PB/PA 1.14, PB/EW 1.34. Scutellum pitch brown, small and broadly triangular, with grey pubescence. Elytra with a slight elevation close to the suture at about 1/4 of their length; each elytron convex, narrow posteriorly, obliquely truncate at apex, with a spine at suture and a feeble tooth at outer angle, the surface (where rubbed bare of pubescence) with 2 kinds of punctures, some minute and very dense, others larger and less numerous, suture just below the scutellum reflexed; EL/EW 5.39; prosternum very sparsely scattered with punctures, moderately pubescent, with hmd margin of epipleural process moderately arcuate. Venters of meso- and metathoracic segments hardly punctate, clothed with silky grey pubescence, metasternum midlongitudinally sulcate, abdominal venter with fine silky pubescence. Legs moderately long and stout; femora slightly compressed, hind pair scarcely reaching the elytral apex, gradually swollen apically and widest at apical 3/5; hmd tibia long and slender; hind tarsi 1 as long as 2+3; claw-bearing joint of 80 JOURNAL, BOMBAY NATURAL HISTORY SOCIETY, 97(1), APR. 2000 L ON G I CORN BEETLES OF B UXA TIGER RESER VE the tarsi long and paronychium nearly always distinctly visible between the claws. Body length: 32-51 mm. Material examined: 2 males, PN, B.T.R., 20. v. 1995; 1 male, RB, B.T.R., 20.V.1995; 1 female, DM, B.T.R., 23.V.1995; 2 males, JY(LT), B.T.R., 24. v. 1995, 25.V.1995; 1 male, RB(LT), 16. ix. 1996; 1 male, SB, B.T.R., 20.V.1997; 1 male, NM, B.T.R., 21. v. 1997. All from Jalpaiguri, W. Bengal Distribution: india: Assam, Uttar Pradesh (Gahan, 1906), Gujarat, Karnataka, Maharashtra, Rajasthan (Beeson, 1961), Bihar, Madhya Pradesh, Orissa (Basak and Biswas, 1993), West Bengal; Afghanistan; Indonesia; Laos; Malaysia; Myanmar; Nepal; The Philippines; Singapore; Sunda Island (Gahan, 1906; Gressitt and Rondon, 1970). Genus: Xoanodera Pascoe Xoanodera Pascoe, 1857, Ent. Soc. (2) iv : 92. Type-species: Xoanodera trigona Pascoe Xoanodera regularis Gahan (Fig. 5) Xoanodera regularis Gahan, 1890, A.M.N.H. (6) V : 52. Male : Dark brown, head, pronotum and elytra (greater part) with dense yellowish-brown pubescence, elytra at base with a ring-like dark brown band encircling the scutellum and a lateral area from the shoulders extending a little beyond the middle dark brown, devoid of dense pubescence; the narrow border between submarginal carina and outer margin sparsely pubescent. Head and 1st antennal joint closely JOURNAL BOMBAY NATURAL HISTORY SOCIETY. 97(1). APR. 2000 81 LONGICORN BEETLES OF BUKA TIGER RESER VE mgulose-punctate. Antennae reddish brown, with a faint covering of grey pubescence. Pronotum strongly and irregularly ridged, scantily clothed with patches of tawny pubescence, laterally with a dense, yellowish brown pubescence. Thoracic and abdominal sternites rather densely covered with greyish white pubescence, legs less so. Head shorter and narrower than pronotum, broadest across eyes, anteriorly sloped, medially raised, posteriorly weakly sloped to a little beyond the eyes, medially with a deep longitudinal sulcus, 2 such on either side of the median, or just behind the antennal socket; vertex densely punctate, and entirely covered by pubescence; frons medially lobed, enclosed by a deep circular sulcus, anterolaterally broadly produced, truncate; clypeus broadly rectangular; HW/PA 1.20; FA/FL 0.50; mandibles broad at base, curved, bluntly pointed at apex, with lateromedian depression, outer margin reflexed up to a little beyond the middle; eyes large, deeply emarginate, extending almost to the frontal sulcus, closely approximated above, with a narrow space in between. Antennae 11- segmented, a little shorter than body, densely covered by minute pubescence; 5th to 10th joints sharply edged in front and acutely angulated at apex, 1 1th shorter than 10th. Pronotum as long as broad, sides unevenly rounded, apex with 1 and base with 2 transverse grooves; PL/PA 1 .44, PL/PW 0.90, PB/PA 1.17, PB/EW 1.66. Scutellum obtuse, with dense pubescence. Elytra parallel-sided, narrowed near apex, posterior submarginal carina with apex truncate and feebly bidentate; EL/EW 6.75; prosternum raised between coxae, sharply deflexed posteriorly, and dilated at the end to meet the epimera, acetabula of front coxae not angulated outwards. Legs moderately long, femora carinate on each side near their lower portions; 1 st joint of hind tarsus shorter than the next 2 united. Body length: 20-21 mm. Material examined: 2 males, SR, Jalpaiguri, West Bengal, 5.iv.l993. Distribution: India: North India?, West Bengal; Laos; Myanmar (Gahan, 1906; Gressitt and Rondon, 1970). Tribe: Clytini Genus: Xylotrechus Chevrolat Xylotrechus Chevrolat, 1860, Ann. Soc. Ent. Fr. : 456. Type-species: Xylotrechus sartorii Chevrolat Xylotrechus srnei (Lap. et Gory) (Fig.: 6) Clytus smei Lap. et Gory, 1841, Hist. Nat. et Iconogr. des Ins. Coleopt. : 37. Male: Black, head and most of pronotum with olive green and yellowish pubescence, such pubescence on elytra forming bands and spots; pronotal disc with 3 black spots, one centrally near base, the other two very near the middle, but a little marginal; elytral bands and spots are as follows: 1) elongately circular yellow band, with the outer margin rather narrow, a little discontinuous near the scutellar apex enclosing (2) a transverse marginal yellow broad band extending to the shoulder hump and narrowing towards the suture, (3) midposteriorly with a transverse yellow band at sutural margin, broad, narrowed towards the margin, (4) apical yellow band broad at apex and narrowed in front with the margin oblique; venter with bands or spots of whitish pubescence. Head a little narrower than pronotum, anteriorly sloped; vertex flat with a median longitudinal carina bifurcating anteriorly; frons with 4 carinae, outer ones strongly curved inwardly, median ones nearly parallel-sided, anteriorly united; clypeus transverse, ridged; HW/PA 1.28; FA/FL 0.58. Antennae shorter than half the body, 1st joint equal to 3rd, 3rd to 5th subequal, 6th to 10th gradually shorter. Pronotum nearly squarish, a little longer than wide, with lateral margins rounded, broadest just below the middle, medially raised; PL/PA 1 .35, 82 JOURNAL, BOMBAY NATURAL HISTORY SOCIETY, 97(1), APR. 2000 LONGICORN BEETLES OF B UXA TIGER RESER VE Fig. 6: Xylotrechus smei (Lap. et Gory): PL/PW 0.90, PB/PA 1.0, PB/EW 2.33. Scutellum small, broad, semilunar, densely clothed with white pubescence. Elytra weakly narrowed at apex and truncate apically; EL/EW 8.66; prosternum produced and truncate, metastemum plate-like, midlongitudinally with one black sulcus, basally and apically with transverse bands of white pubescence. Legs moderate, femora thick, hind pair of legs extending a little beyond abdomen, 1st joint of hind tarsus about twice as long as the next two joints united. Body length: 15 mm. Material examined: 1 male, SR, B.T.R., Jalpaiguri, West Bengal, 5.iv.l993; 1 male, RB(LT), B.T.R., Jalpaiguri, West Bengal, 20. v. 1995. Distribution: india: Assam, Orissa, West Bengal, North, West, Central and South India (Gahan, 1906), Bhutan; Myanmar; Sri Lanka (Gahan, 1906; Beeson, 1961; Basak and Biswas, 1993). . Whole body, B. Antenna, C. Hind leg Tribe: Hesperophanini Genus: Stromatium Serville Stromatium Serville, 1834, Ann. Soc. Ent. Fr.3: 80. Type-species: Callidium barbatum Fabricius Stromatium barbatum (Fabricius) (Fig. 7) Callidium barbatum Fabricius, 1775, Syst. Ent. : 189. Male: Red brown to a little darker; faintly covered with orange brown pubescence, 1 st joint of antennae brown black, rest red brown, apical segments a little darker. Head at base narrower than pronotum, densely and rather coarsely punctate, longitudinally sulcate between the antennae; clypeus short, transversely depressed, anteclypeus leathery; HW/PA 1.19, FA/FL 0.69; mandible short, oblique; eyes rather deeply emarginate, with large lower lobe, extending JOURNAL, BOMBAY NATURAL HISTORY SOCIETY. 97(1). APR. 2000 LONGICORN BEETLES OF B UXA TIGER RESER VE Fig. 7: Stromatium barbatum (Fabricius): A. Whole body, B. Antenna, C. Hind leg anteriorly almost up to the genal edge; antennal tubercles posteriorly raised and bluntly pointed. Antennae 1 1 -segmented, about 1/3 longer than body, with long silky pubescence beneath, 3rd joint longest, 4th slightly shorter than 5th. Pronotum broader than long, subquadrate, with numerous strong coarse punctures; the disc with 5 slightly raised, less distinct tubercles, 2 placed anteriorly, 1 behind middle, and 2 near base, straighter and each marked with a large depression; PL/PA 1.0, PL/PW 0.74, PB/PA 1.0, PB/EW 1.85. Scutellum broadly triangular, with a midlongitudinal depression forming 2 weakly raised lobes on either side. Elytra nearly parallel- sided, narrowed and truncate at apex, coarsely and very densely punctured, each with 2 distinct dorsal and 1 lateral costae, a short sutural tooth at apex; EL/EW 6.57; prosternum weakly sloped at apex. Venters of meso- and metathorax truncate, covered with pubescence, metathoracic plate with median longitudinal black streak; abdominal venters punctate, laterally with rather dense pubescence, medially weakly so. Legs moderately long, femora compressed, fore tibiae very broad a little below the base and gradually narrowed outwards, the middle and hind pairs gradually widened up to the middle; the hind pair nearly reaching elytral apex; 1st joint of the hind tarsus subequal to 2+3, last tarsus with distinct paronychium. Body length: 21-23 mm. Material examined: 1 female, RB(LT), 84 JOURNAL, BOMBAY NATURAL HISTORY SOCIETY, 97(1), APR. 2000 LONGICORN BEETLES OF BUXA TIGER RESER VE B.T.R., 20. v. 1995; 1 male, RM, B.T.R., 30. v. 1996; 1 male, NL, B.T.R., 17.V.1997; 1 male, SB(LT), B.T.R., 19.V.1997. All from Jalpaiguri, West Bengal. Distribution: india: All over; Africa; Bangladesh; Islands of Reunion; Rodriquiz & Seychelles; Pakistan; Malagasy Rep.; Mauritius; Myanmar; North America; Sri Lanka (Gahan, 1906; Beeson, 1961 Khan and Maiti, 1983); England (Beeson, 1961). Tribe: Oemini Genus: Tetraommatus Perroud Tetraommatus Perroud, 1855, Ann. Soc. Linn. Lyon (2) ii:390. Type-species: Tetraommatus filiformis Perroud Tetraommatus filiformis Perroud (Fig. 8) Tetraommatus filiformis Perroud, 1855, Ann. Soc. Linn. Lyon (2) h : 391. Male: Head, pronotum reddish brown; elytra brown; antennae yellow brown, legs yellow. Head narrower than pronotum, narrowed at both ends, broadest medially, flat, raised between the antennal sockets, densely and coarsely punctate, sparsely hairy; clypeus transverse, band-like; HW/PA 1.40; FA/FL 0.75; mandibles dark brown, robust, strongly curved, apically broad and truncate; eyes large, deeply emarginate; gula indicated, apically narrowed. Antennae 1 1-segmented, as long as body, segment III onwards subequal, each at least twice of segment I, sparsely setose. Pronotum subcylindrical, basally broad, anteriorly narrowed, lateral margin sharply rounded towards apex, constricted near base, transversely sulcate striate near middle, densely and finely punctate, sparsely setose; PL/PA 1.40, PL/PW 0.87, PB/PA 1 .53, PB/EW 1 .76. Scutellum short, broad and obtuse. Elytra parallel-sided, sharply curved towards apex, apically blunt, punctate- striate, sparsely setose; EL/EW 5.70; prosternum between fore coxae short, metasternum medially raised, midlongitudinally with a black sulcus. Legs moderately long, intercoxal part of prosternum very short, front coxae contiguous. Fig. 8: Tetraommatus filiformis Perroud: A. Whole body, B. Antenna, C. Hind leg JOURNAL BOMBAY NATURAL HISTORY SOCIETY. 97(1). APR. 2000 85 LONGICORN BEETLES OF B UXA TIGER RESER VE with acetabula angulate outwardly and open posteriorly, middle coxae almost contiguous, femora rather long, pedunculate, apically clavate and darker, basally curved, laterally compressed towards apex; tibia basally dark; tarsi long and narrow; 1st joint subequal to the following 3 joints. Body length: 7-9 mm. Material examined: 1 male, BG, B.T.R., 25. v. 1995; 3 males, TG, B.T.R., 26.V.1996; 2 males, RM, B.T.R., 30. v. 1996. All from Jalpaiguri, West Bengal. Distribution: india: Pondicherry (Gahan, 1906; Beeson, 1961), West Bengal; Sri Lanka (Gahan, 1906 ; Beeson, 1961). Genus: Xystrocera Serville Xystrocera Serville, 1834, Ann. Soc. Ent. Fr. 3: 69. Type-species: Xystrocera globosa (Olivier) Xystrocera globosa (Olivier) (Fig. 9) Cerambyx globosa Olivier, 1795, Entomologist, 4 (67) : 27. Male: Reddish brown; pronotum with green metallic bands: along the anterior and posterior margins, midlongitudinally but narrow, and laterally running oblique joining the fore and hind bands; elytra testaceous yellow, with green metallic bands: the median longitudinal extending obliquely from base, over the shoulder almost to the tip, the outer running from base and at apex turning along the apical margin; head with 2 such rounded spots on either side of the median sulcus of the vertex. Head at base narrower than pronotum, vertical in front, raised, forming ridges, broadly concave from side to side, between the antennae, densely punctate; antennal supports emarginate in front, acutely pointed on the inner side; vertex densely punctate, midlongitudinally sulcate. 86 JOURNAL, BOMBAY NATURAL HISTORY SOCIETY, 97(1). APR. 2000 LONGICORN BEETLES OFBUXA TIGER RESER VE continuing to the clypeus, laterally weakly emarginate; clypeus transverse, strongly ridged, anteclypeus membranous, postclypeus separated from the front by a transverse groove; HW/PA 1.50; FA/FL 0.83; mandible basally straight, apically turned at right angles, knobbed at point of turning; eyes large, deeply emarginate, the lower lobes extended close to the mandibular edge of the genae. Antennae 1 1 -segmented, about 1/3 to twice as long as the body, fust 4 joints strongly warty, margins with blunt spines, these in distal segments greatly reduced and finally obsolete; 1st joint asperate at apex with spines, 3rd to 5th strongly asperate, with the apices thickened and dentate beneath; 3rd joint thicker and about 1/4 shorter than the 4th. Pronotum anteromedially convex, following the transverse depression (-s/*), otherwise weakly depressed medially, anterolaterally depressed on both sides, marginally rounded, base with rounded lobe at the middle, entirely strongly warty; PL/PA 1 .06, PL/PW 0.85, PB/PA 0.69, PB/ EW 3.41. Scute Hum tongue-shaped. Elytra long, anteriorly broad, posteriorly narrowed, apex rounded, densely and strongly punctate; each with 3 slightly raised longitudinal striae - 2 dorsal and 1 lateral EL/EW 19.54; prostemum with transverse striate metallic glossy band, close to the front margin, the rest and the sides of the lower part of prothorax form a convexly raised, very minutely and densely punctate dull red area; mesostemum moderately broad, narrowed posteriorly and truncate at tip, metastemum plate-like, with a black median streak. Legs long, fore legs shorter; femora fusiform-clavate, compressed, pedunculate at base, hind femora long; tibiae compressed. Body length: 23-26 mm. Material examined: 2 males, RB(LT), B.T.R., Jalpaiguri, West Bengal, 26. v. 1995. Distribution: India: Assam, Karnataka, Maharashtra, Tamil Nadu, West Bengal; Celebes; China; Egypt; Hawaiian Islands; Indonesia; Japan; Korea; Laos; Malaysia; Malagasy Rep.; Mauritius; Myanmar; Philippines; Pacific Island; Taiwan; Thailand; Sri Lanka (Gahan, 1906; Beeson, 1961; Gressitt and Rondon, 1970; Khan and Maiti, 1983). Tribe: Thraniini Genus: Thranius Pascoe Thranius Pascoe, 1859, Trans. Ent. Soc. (2) v : 22. Type-species: Thranius gibbosus Pascoe Thranius simplex Gahan (Fig. 10) Thranius simplex Gahan, 1 894, Ann. Mus. Civ. Genov., 34 : 15. Male : Dark brown; head, thorax, abdomen, legs, antennae brown black; antenniferous tubercles, pronotum, scutellum reddish brown; clypeus anteriorly and maxillary palpi yellow. Head narrower than pronotum, wide transversely, strongly sloped anteriorly, frons flat, subquadrate, midlongitudinally sulcate between the eyes, punctate, vertex transversely depressed below the eyes, clypeus transverse, rectangular, punctate; HW/PA 1.28; FA/FL 0.66; eyes rather transverse, long, weakly emarginate, with upper lobe short, not extending behind the antenniferous tubercles, lower lobe rather prominent inwardly. Antennae 1 1 -segmented, shorter than body, joints cylindrical, 1st joint closely and rather finely punctate, apex rather pale. Pronotum parallel-sided, squarish, basal margin straight, anterior margin weakly concave, lateral margin weakly rounded, medially a little broad, midlongitudinally sulcate, strongly gibbose anteriorly, densely punctate; PA/PL 1.14; PL/PW 0.80, PB/PA 1.28, PB/EW 2.25. Scutellum small, obtuse, scantily punctured. Elytra elongate, almost flat above, deflexed at the sides, narrowed up to the middle, the surface densely punctate, with the front edges of the punctures slightly raised; EL/EW 8.37; prosternum punctate; metathoracic plate with a median longitudinal black streak, its episterna very broad in front, nan owed almost to a point posteriorly. Legs moderately long, femora clavate, with the 1st tarsal joint of hind legs a little longer than 2+3 united. JOURNAL. BOMBAY NATURAL HISTORY SOCIETY. 97(1). APR. 2000 87 LONGICORN BEETLES OF B UXA TIGER RESER VE Fig. 10: Thranius simplex Gahan: A. Whole body, B. Antenna, C. Hind leg Body length: 12 mm. Material examined: 1 male, SB, B.T.R., Jalpaiguri, West Bengal, 20.V.1997. Distribution: india: Manipur (Gahan, 1906), West Bengal; Bhutan; Myanmar (Gahan, 1906). Subfamily 2: Pnoninae Tribe: Megopidini Genus: Megopis Serville Megopis Serville, 1832, Ann. Soc. Ent. Fr. i : 162. Type-species: Megopis muticci Serville Megopis (A ego soma) bowringi (Gahan) (Fig. 11) Aegosoma bowringi Gahan, 1894, A.M.N.H. 14 (6): 226. Male: Reddish brown; densely clothed with short faint brown pubescence, elytral costae free. Head narrower than pronotum, elongate behind, eyes more or less inclined in front, densely warty and with deeply distinct midlongitudinal black sulcus; vertex flat; frons anteriorly sloped, posteriorly weakly concave; clypeus transverse; HW/PA 0.87; FA/FL 0.59; mandibles short, oblique, toothless; eyes narrowly emarginate in front; gula short, basally broad, anteriorly narrow, either side marked by black ridge. Antennae shorter than body, basal segments densely punctate, apical 3 segments coarsely wrinkled, 1st joint short and stout. 3rd joint longest, subequal to 4+5. Pronotum broadly transverse, wider than long, its warty basal and apical margins nearly straight, lateral margin medially weakly produced, antero-lateral comers weakly produced, rounded, strongly reflexed: PL/ PA 0.66, PL/PW 0.59, PB/PA 0^83, PB/EW 2.0. Scutellum nearly globose, densely warty. Elytra broader than pronotum, nearly parallel-sided for the greater part of their length, slightly narrowed posteriorly, rounded at apex, with sutural teeth; EL/EW 8.09; prostemum raised, sloped, on either side extending beyond fore coxae; mesosternum sulcate, midlongitudinally blackish, metasternum broad, plate-like, midlongitudinally with a deeply distinct black sulcus; abdomen ventrally a little paler, densely punctate, segmental joints brown- black, transverse, band-like, clothed with pale brown hairs. Legs moderately long, the hind pair 88 JOURNAL, BOMBAY NATURAL HISTORY SOCIETY, 97(1), APR. 2000 LONGICORN BEETLES OFBUXA TIGER RESER VE Fig. 1 1: Megopis ( Aegosoma ) bowringi (Gahan): A. Whole body, B. Antenna, C. Hind leg longest, femora laterally compressed, with their dorsal and ventral borders nearly parallel, tarsi rather narrow, with the claw joint at least as long as 1+2. Body length: 20-23 mm. Material examined: 1 male, RB, B.T.R., 22. v. 1995; 1 male, SB(LT), B.T.R, 19.V.1997. Both Jalpaiguri, West Bengal Distribution: india: Arunachal Pradesh, West Bengal; Bangladesh; Myanmar (Gahan, 1906). Tribe: Macro tomini Genus: Macrotoma Serville Macrotoma Serville, 1832, Ann. Soc. Ent. Fr. 2: 264. Type-species: Prionus serripes Fabricius Macrotoma (. Zooblax ) spinosa Fabricius (Fig. 12) Prionus spinosus Fabricius, 1787, Mailt. Ins. 1 : 130. Male: Red brown, elytra rusty brown towards base and yellowish towards apex, antennae with three basal segments brown-black, legs reddish, venter glossy red. Head elongate behind eyes, coarsely punctate between eyes, closely and finely granulate behind, vertex impressed with a median groove; frons punctate, midlongi- tudinally sulcate due to bulging antennal tubercles, anteriorly vertical and truncate; clypeus depressed, limited above by an impression, weakly punctate; shorter than width of pronotum; HW/PA 0.61; FA/FL 0.76; mandibles vertical, straight at base, incurved at tip, each with 2 teeth on inner edge, punctate; eyes not deeply emarginate on front; venter warty. Antennae 1 1 -segmented, reaching basal 2/3 of elytra, 1st joint apically broad, basally pedunculate, twice as long as broad, closely and coarsely punctate, 3rd segment more than twice JOURNAL, BOMBAY NATURAL HISTORY SOCIETY, 97(1), APR. 2000 89 LONGICORN BEETLES OF BUXA TIGER RESERVE Fig. 12: Macrotoma ( Zooblax ) spinosa (Fabricius): A. Whole body, B. Antenna, C. Hind leg as long as 1 st, 4th onwards shorter, 1 1 th much longer, spinose beneath and along the front margin, the spines rather short, 4th weakly spinose. Pronotum rather strongly de flexed at sides just before the middle, very closely and finely punctate and opaque, with 2 triangular spaces before the middle, a small spot external to each of these, a narrow transverse band near base, a median streak from the middle and an oblique band from each end of the basal band, all more or less strongly lustrous, lateral edges armed with a series of short spines and teeth, basally broad, apically narrowed; PL/PA 0.80, PL/PW 0.53, PB/PA 1.50, PB/EW 2.14. Scutellum long, tongue-shaped. Elytra much longer than broad, rounded at apex, usually dentate at suture, rugulose-punctate and very finely granulose, the granules more distinct and the surface rough towards base, especially on the slightly elevated part near scutellum, each with 4 longitudinal striae; EL/EW 6.00; prostemum sloped on either side, mesosternum at apex slightly clubbed, metasternum plate-like, medially sulcate with a longitudinal black streak. Legs long, spinose beneath; fore femora and tibiae asperate with short sharp spines beneath, those on mid and hind legs reduced and punctate; middle and hind femora sparsely punctate, armed with a few spines beneath, 1 st joint of front tarsus a little shorter than 2+3. 90 JOURNAL, BOMBAY NATURAL HISTORY SOCIETY, 97(1), APR. 2000 L ONGICORN BEETLES OF B UXA TIGER RESER VE Body length: 56 mm. Material examined: 1 male, JY(LT), B.T.R., Jalpaiguri, West Bengal, 25.V.1996. Distribution: india: Bihar, Karnataka (Gahan, 1906), West Bengal; Arabia; Laos; Sri Lanka; (Gahan, 1906; Gressitt and Rondon, 1970). R E F E Basak, P.K. & S. Biswas (1993): Insecta: Coleoptera: Cerambycidae. Zool. Surv. India: State Fauna Series, 1 : Fauna of Orissa, (Part 4): 185-195. Beeson, C.F.C. (1961): The Ecology and Control of Forest Insects of India and the Neighbouring Countries. Govt, of India Publication, Delhi, pp. 767 (Reprint). Gahan, C.J. (1906): Fauna of British India. Coleoptera. Vol. 1 (Cerambycidae), Taylor and Francis, London, pp. 329. Ghosh, A.K. & T. Sengupta ( 1 982): Handbook on Insect Collection, Preservation and Study (Ed. Director), Zool. Surv. India, Calcutta, pp. 64. Acknowledgements We thank the authorities of Buxa Tiger Reserve for facilities and the Head of the Department of Zoology, University of Calcutta for kind permission to carry out the work. E N C E S Gressitt, J.L. & J. A. Rondon (1970): Cerambycidsof Laos (Disteniidae. Prioninae, Philinae, Aseminae. Lepturinae, Cerambycinae). Pacific Insects Monograph, 24: 1-314 Khan, T.N. & P.K. Maiti (1983): Studies on the biotaxonomy, biology and ecology of some longicom beetle borers (Coleoptera: Cerambycidae) of the islands of Andaman, India. Rec. zool. Surv. India. Misc. Pubi. Occ. Paper, No. 45, 1-100. Raychaudhuri, D. (1996); Longhorn beetles (Cerambycidae : Coleoptera) of Buxa Tiger Reserve, Jalpaiguri, West Bengal. Insect Environment 2(3): 81 . JOURNAL, BOMBAY NATURAL HISTORY SOCIETY, 97(1), APR. 2000 91 FISHES OF THE CYPRINID GENUS SEMIPLOTUS BLEEKER 1859, WITH DESCRIPTION OF A NEW SPECIES FROM MANIPUR, INDIA1 Waikhom Vishwanath and Laishram Kosygin2 {With one text-figure and one plate) Key words: Cyprinid fish, Semiplotus, new species, Manipur The cyprinid fishes of the genus Semiplotus Bleeker are distributed in northern India, Myanmar and Nepal. Four species (including a new one) of the genus are recognised. They are: S. semiplotus (McClelland), S. modestus Day, S. cirrhosus Chaudhuri and S. manipurensis sp. nov. This paper describes the new species from the rivers and streams draining Ukhrul dist. of Manipur (Chindwin drainage), India. Semiplotus manipurensis differs from S. semiplotus and S. cirrhosus in having a broader body, fewer branched dorsal rays and several horny tubercles scattered randomly on the snout. It differs from S. modestus in having a broader body and an unserrated last dorsal spine. Semiplotus cirrhosus is considered a valid species. A key to identification of species of the genus Semiplotus is provided. Introduction Bleeker (1859) established the genus Semiplotus to accommodate Cy prinus semiplotus McClelland, 1839 (type locality: River Brahmaputra, Assam). Day (1870) described another species, S. modestus from Akyab in Myanmar and distinguished it from S. semiplotus by the serrated last dorsal spine. Later, Chaudhuri (1919) described S. cirrhosus based on a single specimen collected from Putao of Myanmar, and distinguished it from the former two species mainly by the presence of two pairs of maxillary barbels and absence of knob at the symphysis of the lower jaw. However, Hora (1973) treated S. cirrhosus as a synonym of S. semiplotus. Jayaram (1981) included only S. semiplotus and S. modestus in the genus. The distribution of the genus is restricted to the Himalayan foothills of Nepal, north and northeast India and Myanmar (Fig. 1). On the basis of its jaw anatomy, Howes (1982) put Semiplotus under the genus Cyprinion Heckel, 1843. Talwar and Jhingran (1991) recognised Semiplotus as a subgenus of Cyprinion without justification. However, Banarescu and Herzig (1995) recognised Semiplotus as a distinct genus, as it has more 'Accepted January, 1999 department of Life Sciences, Manipur University, Canchipur 795 003, Manipur, India. branched dorsal fin rays. No detailed revisional work on this genus has been conducted, and very little is known about the fishes of this genus. This is partly due to the difficulty in obtaining specimens. A brief revision of the genus Semiplotus is made here. Material and Methods The new species was collected by cast net. Type specimens are deposited in the Manipur University Museum of Fishes (M-UMF) and National Science Museum, Tokyo (NSMT). Type and other specimens of S. cirrhosus, S. modestus and S. semiplotus in Zoological Survey of India, Calcutta were re-examined. Measurements and counts follow Jayaram (1981). Body proportions are expressed as percentage of standard length (SL) and head length (HL). Total number of vertebrae was counted from radiographs and dissected specimens. Transverse scales were counted as scales between lateral line and dorsal fin origin (including mid-dorsal scale)/lateral line scale/ scales between lateral line and pelvic fin origin. Semiplotus Bleeker. 1 859 Semiplotus Bleeker, 1859, Nat. Tijdschr. Neder. -Indie. 20: 424 (type species Cyprinus semiplotus McClelland, 1839); Banarescu & 92 JOURNAL, BOMBAY NATURAL HISTORY SOCIETY 97(1 ) APR. 2000 NEW AND KNOWN FISHES OF THE CYPRINID GENUS SEMI P LOTUS Fig. 1: Drainages of Nepal, northern and eastern parts of India and Myanmar showing the distribution of known species of Semiplotus. Herzig-Straschil, 1995, Ann. Naturhist. Mus. Wien., 97 B: 411 (status discussed). Diagnosis: A genus of Cyprinidae with the following combination of characters: body large and deep (depth 35.4-41.3% SL); head short, as long as high at occiput (height 93.3-1 16.1% HL); snout broad, blunt with open pores or tubercles; mouth inferior, wide (width 45.3-65.2% HL) with exposed cornified mandibular cutting edge; dentary with a broad deflected labial surface; maxillary barbel rudimentary; long dorsal fin with 20-25 branched rays; anal fin with 7-9 branched rays; lateral line scales 27-36; lower jaw with a knob at symphysis. Distribution: india, Ganga-Brahmaputra, Kaladan and Chindwin drainages, Nepal and Myanmar. Remarks: Banarescu and Herzig (1995) differentiated Semiplotus from Cyprinion on the basis of (i) more branched dorsal fin rays (20-25 vs. 9-17), (ii) fewer branched anal fin rays (5 vs. 7) and (iii) no barbels. The first character holds true. However, the characters (ii) and (iii) differ from our observations. All the Semiplotus specimens studied by us have a pair of small maxillary barbels and 7-9 branched anal fin l ays. From the literature it is also observed that Semiplotus has more pelvic rays (8-9 vs. 7), fewer scales on lateral line (27-36 vs. 33-45), and a deeper body than Cyprinion. Key to the species of genus Semiplotus Bleeker la Last simple dorsal ray serrated posteriorly; branched dorsal rays 20-21 S. modestus 1 b Last simple dorsal ray not serrated posteriorly; branched dorsal rays 20-25 2 2a Tubercles on snout randomly distributed on each side of tip of snout; branched dorsal rays 20-23 S. mcinipurensis JOURNAL. BOMBAY NATURAL HISTORY SOCIETY. 97(1). APR. 2000 93 NEW AND KNOWN FISHES OF THE CYPRINID GENUS SEM IPLOTUS 2b Tubercles/open pores on snout arranged in a transverse row; branched dorsal rays 23-25 ... 3 3a Open pores on snout 4; lateral transverse scales 7/1/4 S. cirrhosus 3b Open pores on snout 10-12; lateral transverse scales 6/1/4 S. semip lotus Semiplotus cirrhosus Chaudhuri, 1919 Semiplotus cirrhosus Chaudhuri, 1919, Rec. Indian Mus. 16(4): 280 pi. 22 figs 3, 3a (type locality: Putao plains, Burma); Hora, 1973, Rec. Indian Mus., 39(1): 46 (part). Material examined: ZSI F 9747/1 holotype, 41.0 mm SL, Myanmar: Putao plains near Tibetan frontier, coll. Murray Stuart, ? . ii . 1 9 1 8 Diagnosis: A species of Semiplotus with large eye (diameter 36.2% HL); predorsal length 47.6% SL; a row of 4 open pores (2 on each side) on the snout; the last simple dorsal fin ray not serrated; 25 branched dorsal fin rays; 8 branched pelvic fin rays; 9 branched anal fin rays; a small knob at the symphysis of lower jaw. Description: Dorsal rays iii, 25; pectoral rays 15; pelvic rays i, 8; anal rays ii, 9; lateral line scales 33; scales above lateral line to origin of dorsal fin 7; scales below lateral line to origin of pelvic fin 4; predorsal scales 13. Head and body laterally compressed. Snout broad, obtuse, with a row of 4 open pores (2 on each side). Maxillary barbels well developed, extending to below anterior margin of orbit. Eye large, almost in the middle of head. Caudal peduncle deep. Dorsal fin origin slightly nearer snout tip than caudal fin base. Last simple dorsal ray not serrated. Pectoral fin almost reaching pelvic fin origin. Caudal fin forked. Colour: Head and body silvery with black dorsal surface. Ventral surface dull white. Distribution: Myanmar: Putao plains (Irrawady drainage). Remarks: Chaudhuri (1919) described S. cirrhosus and differentiated it from other Semiplotus by the presence of two small maxillary barbels and the absence of a knob at the symphysis of the lower jaw. Hora (1937) treated S. cirrhosus as a synonym of S. semiplotus after he found that all other specimens of the genus in ZSI possessed small maxillary barbels. It has not been possible to examine more specimens from Myanmar. However, the holotype of S. cirrhosus in ZSI (F9747/1) has been examined. The species differs from S. semiplotus as it has fewer pores on the snout [4 (2 on each side) vs. 10-12 (5-6 on each side)]; a longer head (length 26.9% SL vs. 21.9-23.7); larger eye (diameter 36.2% HL vs. 20.8-30.0); longer predorsal length (47.6% SL vs. 39.5-44.2); one more scale row between dorsal fin origin and lateral line (7 vs. 6) and fewer branched pelvic fin rays (8 vs. 9). The anal fin of the holotype is damaged. But Chaudhuri (1919) reported that it had two simple and nine branched rays. Thus, it also differs from 5. semiplotus as it has more branched anal rays (9 vs. 7). Thus S. cirrhosus is treated here as a separate species. Semiplotus manipurensis sp. nov. (Plate 1 Figs. 1, 2a) Material examined: Holotype: MUMF 2049, 83.5 mm SL, India: Chall ou river at Thetsi, near Jessami, Manipur (Chindwin basin), 94° 35’ E, 25° 38’ N, about 1,270 m above msl, coll. L. Kosygin, 2.vi.l994. Paratypes: NSMT-P 52636. 1 ex., 85.0 mm SL, same data as holotype; MUMF 2011, 2045- 2048,2051-2055,2145,2146, 12 ex., 55.3-126.0 mm SL, same data as holotype; MUMF 2236- 2240, 5 ex., 42.9-57.5 mm SL, India: Chall ou river, Chingai, Manipur, 94° 3 V E, 25° 18’ N, 130 km northeast of Imphal, 30.iv.1995; MUMF 2250, 2251, 2 ex., 53.3-185.0 mm SL, India: Wanze stream, Khamsom, Manipur, (Chindwin basin), 116 km northeast of Imphal. 94° 32’ E, 25° 12’ N, coll. L. Kosygin. 7.vii.l995. Diagnosis: A species of Semiplotus with a broad body (width 17.3-22.1% SL); last dorsal spine not serrated; 20-23 branched dorsal fin 94 JOURNAL BOMBAY NATURAL HISTORY SOCIETY. 97(1) APR. 2000 NEW AND KNOWN FISHES OF THE CYPRINID GENUS SEMIPLOTUS Vishwanath, Waikhom et al: Semiplotus manipurensis sp. nov. Plate 1 Fig. 1 : Semiplotus manipurensis sp. nov (holotype, MUMF- 2049, 83.5 mm SL). Scale bar indicates 10 mm a b Fig. 2: Front view of snout showing arrangement of tubercles/open pores: a. S. manipurensis (MUMF-2251, 185.0 mm SL); b. S. semiplotus (ZSIF-2662/2, 181.0 mm SL) JOURNAL, BOMBAY NATURAL HISTORY SOCIETY, 97(1), APR. 1999 95 1 NEW AND KNOWN FISHES OF THE CYPRINID GENUS SEM I PL OTUS rays; 9 branched pelvic fin rays; 12-13 predorsal scales; dorsal fin base length 34.0-39.7% SL; 32- 36 lateral line scales; 7 scale rows between dorsal fin origin and the lateral line; many horny tubercles distributed randomly on each side of snout tip, extending posteriorly to the region below the anterior margin of orbit. Description: Dorsal rays iv, 20-23 (last ray branched at base); pectoral rays 15-16; pelvic fin rays i, 9; anal fin rays ii-iii, 7-8 (last ray branched at base); principal caudal fin rays 10 + 9; lateral line scales 32-36; scales above lateral line to origin of dorsal fin 7; scales below lateral line to origin of pelvic fin 4; predorsal scales 12- 13; total vertebrae 36. Body short, deep and compressed. Dorsal profile arched from tip of snout to dorsal fin origin and then gently sloping down to caudal fin base. Dorsal profile more convex than ventral. Abdomen edge rounded. Head short and small compared to body depth, almost as long as high at occiput. Snout thick, prominent, broad, obtuse, overhanging the mouth. Snout with horny tubercles distributed randomly on each side, extending to the region below the anterior margin of orbit. Tubercles larger and more prominent towards tip of snout, smaller and less prominent posteriorly. Tubercles not well developed in small specimens (<56.0 mm SL). Number and size of tubercles increasing with total length. Eye large, not visible from below, placed almost in middle of head. Nostrils close to each other, closer to eye than to tip of snout. Mouth wide, transverse, inferior, lower jaw with a knob at symphysis, and an exposed cornified cutting edge. Small maxillary pair of barbels, more prominent in smaller specimens, hardly visible in larger specimens as they are concealed in groove between maxilla and snout. Scales moderate to large, those on chest and abdomen smaller than those of other parts of body. Lateral line complete. Dorsal fin origin nearer to snout tip than to caudal fin base, extending from a little ahead of pelvic fin to above anal fin base. Last simple dorsal ray strong, osseous and not serrated in large specimens. In small specimens (<130 mm SL), distal third of spine slightly serrated posteriorly. Height of dorsal almost equal to head length. Pectoral fin shorter than head, not reaching pelvic fin origin, latter not reaching anal fin origin. Caudal fin deeply forked with a slightly longer upper lobe. Colour: Body silvery white, slaty grey dorsally. All fins tinged orange with dusky edges. Distribution: india: Chall ou river and Wanze stream (Chindwin drainage), Ukhrul District, Manipur. Etymology: The species is named after the state of Manipur. Habitat: Moderate to fast flowing hill streams with rocky beds. Smaller specimens inhabit shallow and fast flowing water, while larger ones inhabit deeper waters where water current is comparatively slow. Remarks: Semiplotus manipurensis differs from S. semiplotus in its wider body (width at dorsal fin origin 17.3-22.1% SL vs. 1 1.8-16.7), fewer branched dorsal fin rays (20-23 vs. 23-25) and randomly distributed tubercles on either side of the tip of snout vs. a transverse row of open pores on the snout including its tip [all the specimens of S. semiplotus in ZS1 and the freshly collected specimen (MUMF 2307) from the Brahmaputra river at Dibrugarh, Assam have open pores on snout, while the 2 1 specimens of S. manipurensis have tubercles on snout]; shorter dorsal fin base (34.0-39.7% SL vs. 40.9-44.6); more scales in lateral line (32-36 vs. 27-33) and one more scale row between the origin of dorsal fin and lateral line (7 vs. 6). The new species is also distinct from S. cirrhosus as it has a wider head (63.3-74.2% HL vs. 58.8); wider body (width at dorsal fin origin 17.3-22.1% SL vs. 11.1); fewer branched dorsal rays (20-23 vs. 25); smaller eye (diameter 20.0-31.8% HL vs. 36.2); shorter predorsal length (40.8-45.7% SL vs. 47.6); one more branched pelvic fin ray (9 vs. 8): fewer branched anal fin rays (7-8 vs. 9) and many randomly in JOURNAL. BOMBAY NATURAL HISTORY SOCIETY. 97(1). APR. 2000 NEW AND KNOWN FISHES OF THE C.YPRINID GENUS SEM IPLOTUS distributed tubercles on snout (vs. a transverse row of 4 open pores across the snout). Semiplotus manipurensis is distinct from S. modestus as it has fewer predorsal scales (12- 13 vs. 14-15); broader body (width at dorsal fin origin 17.3-22.1% SL vs. 9.9%); more branched pelvic rays (9 vs. 8) and last dorsal spine not serrated posteriorly (vs. serrated). Semiplotus modestus Day, 1870 Semiplotus modestus Day, 1870, Proc. Zool. Soc. Lond.: 101 (type locality: Akyab, Burma); Barman, 1988, J. Bombay nat. Hist. Soc. 85(1): 210 (Koladyne R., Mizoram). Cyprinion modestum : Howes, 1982, Bull. Brit. Mus. nat. Hist. (Zool), 42(4): 331 (status discussed). Material examined: ZSI 2343, 1 ex., (syntype), 85.4 mm SL, Myanmar: hill ranges near Akyab, coll. F. Day, no date. Diagnosis: A species of Semiplotus with last dorsal spine osseous and serrated posteriorly; 20-2 1 branched dorsal fin rays; 8 branched pelvic fin rays; 14-15 predorsal scales; 32-34 lateral line scales; and several open pores on either side of snout. Description: Dorsal fin rays iv, 20-21; pectoral fin rays 15; pelvic fin rays i, 8; anal fin rays iii, 7; principal caudal fin rays 10 + 9; lateral line scales 32-34; scales above lateral line to origin of dorsal fin 7; scales below lateral line to origin of pelvic fin 4; predorsal scales 14-15. Body deep, laterally compressed. Head short with concave dorsal profile, as long as high at occiput. Snout short, obtuse, overhanging the mouth with several open pores on either side. Maxilla extending below the middle of orbit. Eye large, longer than snout. Dorsal fin origin nearer snout tip than caudal base. Last simple dorsal ray serrated posteriorly. Pectoral fin extends to pelvic fin origin, latter to anal fin. Caudal fin forked, lower lobe slightly longer than upper. Colour: Silvery grey with black dorsal surface. Pelvic and anal fins orange. Distribution: india: Kaladan river (Koladyne river as per Barman, 1 988), Mizoram; Myanmar: Akyab. Remarks: The species is quite distinct from other members of the genus Semiplotus as it has a posteriorly serrated last dorsal spine. Semiplotus semiplotus (McClelland, 1839) (Plate 1 Fig. 2b) Cyprinus semiplotus McClelland, 1839, Asiatic Researchers, 19(2): 274, 346, pi. 37 fig. 2 (type locality: River Brahmaputra, upper Assam, India). Semiplotus mcclellandi: Day, 1878, Fishes of India: 550 (description). Semiplotus semiplotus: Hora, 1937, Rec. Indian Mus., 39:45 (part). Cyprinion semiplotum: Howes, 1982, Bull. Brit. Mus. nat. Hist. (Zool), 42(4): 331, figs la- c (Jaw structure studied, status discussed). Material examined: MUMF 2307, 1 ex., 131.4 mm SL, India: Brahmaputra river, Dibrugarh, Assam, coll. L. Kosygin, 22.x. 1995; ZSI F 2861/2 1 ex., 162.0 mm SL, India: Darjeeling Himalayas, coll. G.E. Shaw & E.O. Shebbeare, 28.iii.1937 ZSI F 2662/2, 3 exs. 89.7- 181.0 mm SL, India: Tista drainage, S.L. Hora, ?.xi.l938. Diagnosis: A species of Semiplotus with last simple dorsal fin ray not serrated; 23-25 branched dorsal fin rays; a transverse row of 10- 12 open pores (5-6 on each side) across the snout posteriorly directed toward middle of orbit. Description: Dorsal fin rays iv, 23-25; pectoral fin rays 15-16; pelvic fin rays i, 9; anal fin rays ii, 7 (last ray branched at base); principal caudal fin rays 10 + 9; lateral line scales 27-33; scales above lateral line to origin of dorsal fin 6; scales below lateral line to origin of pelvic fin 4; predorsal scales 11-12. Head and body deep, laterally compressed, with convex dorsal profile. Snout blunt with a very distinctive transverse row of 10-12 (5-6 on each side) open pores across it. 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Eye moderate, almost in middle of head. Mouth wide, inferior, lower jaw with a horny layer. Barbels a small maxillary pair, more prominent in smaller specimens. Dorsal fin high, with long base. Last dorsal simple ray strong, osseous, not serrated in large specimens but slightly serrated in distal half in juveniles. Pectoral fin equal to head, almost reaching pelvic fin origin. Pelvic fin shorter than pectoral, not reaching anal fin. Caudal fin forked. Colour: Dull silvery with black dorsal surface. Pectoral, pelvic and anal fins orange. Distribution: india: Arunachal Pradesh, Assam (Brahmaputra drainage), north Bengal; Nepal: Terai (Ganga drainage). Remarks: According to Day (1878), the species is often termed Rajah-mas (King fish) in upper Assam, as it was asserted that when captured it had to be taken to the Rajahs for their own consumption. He also remarked on the statement of McClelland that the fish attained at least two feet in length and was reckoned the most delicious in Assam. Menon (1989) included S. semiplotus in the list of endangered freshwater fishes of India. The underutilised hill stream fishes of Nepal were listed by Shreshtha (1997), who included this species, and suggested the possibility of developing recreational fishery of these fishes in Nepal. Discussion Most workers (Bleeker, 1859; Gunther, 1868; Day, 1878; Jayaram, 1981; Barman, 1988) erroneously considered that Semiplotus lacks barbels. However, Hora (1937) examined all the specimens of Semiplotus in ZSI and a specimen from Nepal collected by Col. Bailey, and concluded that the presence of small maxillary barbels is a constant feature of the genus. He further remarked that in young specimens barbels are longer and project outside the groove, whereas in half-grown and adult specimens they are more or less concealed, though it is not very difficult to make them out. This statement of Hora {op. cit) holds true for the present study, as all the specimens of Semiplotus examined (including the type specimens of S. manipurensis ) have a small pair of maxillary barbels. Thus the presence of a small pair of maxillary barbels is a distinct character of the genus Semiplotus. Interesting observations have been made in the ichthyogeography of Semiplotus species which are endemic in Southeast Asia. McClelland (1839) originally described S. semiplotus from the Brahmaputra river, upper Assam. Day (1878) put the fish under S. mcclellandi and reported that it inhabited the rivers of Assam, especially in the upper portion but was also found as low as Goalpara and in Myanmar. Gunther (1868) on the other hand mentioned only Assam as the place of its distribution. Mukerji (1933) included this species in the list of fishes of Mali Hka river, upper Myanmar without giving a systematic account. As there is no specimen of the fish collected by either F. Day or D.D. Mukerji in ZSI (although they are supposed to be there), it is difficult to establish the correct identity of the species and its distribution in Myanmar. Hora (1937) reported this fish from the Nepal terai which is drained by tributaries of the Ganga. Thus, S. semiplotus is perhaps present only in the Ganga-Brahmaputra drainage. On the other hand S. cirrhosus and S. manipurensis share the Clnndwin-Irrawaddy drainage, which is entirely separate from the Brahmaputra drainage (Chaudhuri, 1919). Further, distribution of S. modestus is totally isolated from other species of the genus. The species is distributed in Akyab of Myanmar and parts of Mizoram (India) which are drained by the Kaladan drainage which enters the Bay of Bengal directly. Kaladan drainage is separated from the Barak- Brahmaputra drainage of India by the Chittagong hill tract. The region is also separated from the Chindwin-Irrawaddy drainage of Myanmar by the north-south JOURNAL. BOMBAY NATURAL HISTORY SOCIETY. 97(1). APR. 2000 101 NEW AND KNOWN FISHES OF THE CYPRINID CMOS'S EM IP LOTUS extension of the Arakan Yoma hill range. From the above statements it is clear that Semiplotus is distributed in north India, Myanmar and Nepal, with restricted distribution in different drainages. A detailed study of the geological history of the region may give a true picture of the phylogeny of these fishes. Refer Banarescu, P.M. & B. Herzig-Straschjl(1995): A revision of the species of the Cyprinion mcicrostomus - group (Pisces: Cyprinidae). Ann. Naturhist. Mus. Wien. 97 B 41 1-420. Barman, R.P. (1988): First record of the King-fish, Semiplotus modestus Day, 1870 (Pisces: Cyprinidae) from India. Bombay nat. Hist. Soc. 85(1): 210. Bleeker, P. (1859): Conspectus systematics cyprinorum. Nat. Tijdschr. Neder. -Indie, 20: 421-441 . Chaudhuri, B.L. (1919): Report on a small collection offish from Putao (Hkamti Long) on the northern frontier of Burma. Rec. Indian Mus., 16(4): 271-282. Day, F. ( 1 870): The fishes of India; being a natural history of the fishes known to inhabit the seas and freshwaters of India, Burma and Ceylon, William Dowson and Co., London, pp. 778. Gunther, A. (1 868): Catalogue of the fishes in the British Museum, John Edward Gray, London, 7, pp 512. Heckel, J.J. (1843): Abbildungen and Beschreibungen der Fische Syriens. In: Russegger, J. Reisen in Europa, Asien und Afrika Bd. 1, T. 2. Stuttgart, Schweizerbart’sche Verlags-buchhandlung. 991 - 1099. Hora, S.L. (1937): On a collection of fish from Nepal. Rec. Indian Mus. 39(1): 43-46. Howes, G.J. ( 1 982): Anatomy and evolution of the jaws in the semiplotine carps with a review of the genus Acknowledgements We thank Dr. J.R.B. Alfred, Director, ZSI and Mr. T.K. Sen, Head, Fish Section, ZSI, for their help in examining types of Semiplotus. We also thank Dr. Keiichi Matsuura, Chief Curator, National Science Museum, Tokyo, for his help in registering a type of the new species. NCES Cyprinion Heckel, 1843 (Teleostei: Cyprinidae). Bull. Brit. Mus. nat. Hist. (Zool), 42(4): 299-335. Jayaram, K.C. (1981): Freshwater fishes of India, Pakistan, Bangladesh, Burma and Sri Lanka — a handbook. Zoological Survey of India, Calcutta. 475 pp. Kottelat, M. (1989): Zoogeography of the fishes from Indo-Chinese inland waters with an annotated checklist. Bull. Zoologisch Museum. Univ. Amsterdam. 12(1): 1-56. Menon, A.G.K. (1989): Conservation of the ichthyofauna of India. In: Jhingran A.G. and V. V. Sugunan, (Eds). Conservation and management of Inland capture fisheries resources of India. Inland Fisheries Society of India, Barrackpore: 25-33. McClelland, J. (1839): Indian Cyprinidae. Asiatic Researches, 19(2): 274, 346. Mukerji, D.D. (1933): Report on Burmese fishes collected by Lt.-Col. R.W. Burton from the tributary streams of the Mali Haka River of the Myitkyina district (upper Burma). J. Bombay nat. Hist. Soc. 36(4): 812-831. Shreshtha, T.K. (1997): Sustained development of fisheries resources of Himalayan waters of Nepal. J. Freshwater Biol. 9(1): 47-56. Talwar, P.K. & A.G. Jhingran (1991): Inland Fishes of India and adjacent countries, /, Oxford and IBH Publ. Co. Pvt Ltd., New Delhi, 541 pp. 102 JOURNAL, BOMBAY NATURAL HISTORY SOCIETY. 97(1) APR. 2000 FOOD AND FEEDING HABITS OF INDIAN BARBETS, MEGALAIMA SPP.1 Hafiz SA. Yahya2 ( With three text-figures) Key words: Barbets, congeneric, sympatric, food, feeding method, ecological isolation, coexistence, conservation. A comparative study on the ecology and biology of Indian barbets ( Megalaima spp.) was carried out between 1 977 and 1 980 in different parts of the country. Megalaima viridis and M. rubricapilla malabarica were studied more intensively at Thekkady (Kerala), while M. virens, M. zeylanica, M. lineata, M. asiatica, M. franklinii and M. haemacephala were studied at other places. The findings on the feeding behaviour of the barbets are discussed in this paper. Data on food items, fruiting seasons, abundance of fruiting trees, feeding method and extent of ecological isolation in food habits of coexisting species are discussed. The barbets are predominantly frugivorous, but during the breeding season all species feed their young with insects. Among the congeneric sympatric M. viridis and M. rubricapilla at Thekkady, the former was found to be more insectivorous, helping considerably in checking the deadly teak defoliator Hyblaea puera. Contrary to reports of M. zeylanica and M. viridis being minor pests on coffee, they were found to be quite helpful to coffee plants in picking up the coffee stem borer, Xylotrechus quadripes. Barbets also help in seed dispersal and pollination of scores of trees, and thus play an important role in maintaining the rich biodiversity of the country, and they deserve conservation priorities. Introduction The name barbet is derived from the French Barbu (=bearded) which is suggested by the presence of nasal and rictal bristles. They are closely related to Old World honeyguides (Indicatoridae) and the New World puff birds (Bucconidae). The barbet family Capitonidae has a pantropical distribution. Ripley (1961) reported 10 species from the Indian subcontinent under the single genus Megalaima. According to Simmons (1970), food supply plays an important role in determining the breeding biology, dispersion pattern and social system of a species through natural selection. In this paper, apart from mentioning the main food items, fruiting seasons and abundance of fruiting 'Accepted June, 1 999 ^Centre of Wildlife & Ornithology, Aligarh Muslim University, Aligarh 202 002, Uttar Pradesh, India. trees at Thekkady, the food and feeding methods of coexisting M. viridis and M. rubricapilla are described to ascertain the extent of isolation in food habits. Food habits of M. zeylanica and M. haemacephala are also discussed briefly. The impact of food habits of M. viridis on coffee plantations was assessed and has been published elsewhere (Yahya 1982). Barbets do not drink water regularly, but they were often recorded drinking water and bathing from the rain filled natural tree holes. Drinking and bathing behaviour have been described elsewhere (Yahya 1991). The study was carried out mainly in the Periyar Tiger Reserve (9° 30' N lat. and 77° 10' E long.) Kerala, consisting of evergreen, semi- evergreen, shola, moist-deciduous and savanna forests. Details of the study area have been published earlier (Ali 1935, Yahya 1980, 1988, 1989, Vijayan 1984, and Robertson and Jackson 1992). Comparative studies were made at several other locations. JOURNAL, BOMBAY NATURAL HISTORY SOCIETY, 97(1), APR. 2000 103 FOOD AND FEEDING HABITS OF INDIAN BARBETS On reconnaissance, it was found that moist deciduous forest was favoured most by the barbets (Yahya 1989). Therefore, Thekkady - a small area of moist deciduous forest, 3 km in length and an average of 0.5 km wide, was selected for intensive study. A road of about 4 km passes through the middle of the forest connecting the reserve to the nearby town Kumily. There are several buildings in this area, besides a picnic spot and a caravan park. Methods Barbets were observed in their natural condition for about three years to study various aspects of their ecology and biology. Data was collected on food and feeding habits between April 1978 and April 1979. Each day was divided into three 4-hour shifts; 0600 to 1000 hrs, 1000 to 1400 hrs and 1400 to 1800 hrs. Observations were made on alternate shifts. On two days in each month, barbets were followed for the whole day. Fruiting of trees was recorded each month by trekking through different routes in the study area at least every fifth day. The data collected included food items, heights at which the birds fed, number of birds feeding at that time and any antagonistic behaviour. Barbets are mostly arboreal birds and only on five occasions did I note M. viridis searching for food on the ground. Hence, the vertical height distribution of feeding zone was divided into three broad levels: Primary level 1 to 4 m. Secondary level 4 to 8 m and Tertiary level above 8 m. In the beginning, I tried to distinguish different canopies at which the birds fed, but this was not done later as both species were found exploiting the canopy equally. The total numbers of each species of barbet recorded feeding on different fruiting trees and hunting insects were considered during the final analysis. As barbets hunt in the brighter hours of the day and in exposed areas, it was possible to identify such prey as cicadas, leafhoppers, ants, termites, butterflies, spiders, beetles and caterpillars. But barbets were seen to be primarily frugivores, and easy to observe visually, therefore no specimen was collected for stomach analysis. The data gathered from April 1978 to April 1979 are analysed here. During this period, a total of 3,346 M. viridis and 1,889 M. l'ubricapilla were recorded feeding. Fruiting season and relative abundance of FRUIT TREES IN THE INTENSIVE STUDY AREA Fruiting/flowering seasons of the principal trees/shrubs on which barbets were found feeding/sipping and relative abundance of fruiting trees in the intensive study area are shown in Tables 1 and 2 respectively. Table 1 RELATIVE ABUNDANCE OF DIFFERENT SPECIES OF FRUITING/FLOWERING TREES VISITED BY BARBETS FOR FOOD IN THE STUDY AREA (3 km x Vi km) Plant species Relative abundance <5 5 to 10 10 to 15 >15 Actinodaphne hookeri X Bischofw javanica X Bridelia retusa X Bombax ceiba X Careya arborea X Erythrina sp. X Eucalyptus sp. X Evodea lunuankenda X Ficus gibbosa X F. infectoria X F. insignis X F. mysorensis X F. retusa X F. tsiela X Grewia tiliaefolia X Lantana camara X Leea indica X Machilus macrantha X Macaranga sp. X Olea dioica X Santalum album X Scolopia crenata X Solanum indicum X Spathodea campanulata X Syzygium cumin i X Ziziphus sp. X 104 JOURNAL , BOMBAY NATURAL HISTORY SOCIETY. 97(1). APR. 2000 FOOD AND FEEDING HABITS OF INDIAN BARBETS Table 2 FRUITING/FLOWERING SEASONS* OF PRINCIPAL TREES AND SHRUBS ON WHICH BARBETS FEED Plant species Jan Feb Mar Apr May Jun Jul Aug Oct Nov Dec Actinodaphne hookeri X X X X Bischofia javanica X X Bridelia retusa X X X Bombax ceiba X X X X X Careya arborea X X Erythrina sp. X X X X X Eucalyptus sp. X X X X Evodea lunuankenda X X X Ficus gibbosa X X X X X X F. infectoria X X X F. insignis X X X F. mysorensis X X X X X X X X X F. retusa X X X X X X X X X X F. tsiela X X X X X X X X X X X Grewia tiliaefolia X X X X Lantana camara X X X X X X X X X X X Leea indica X X X X X X X Machilus macrantha X X X Macaranga sp. X X X X X X Olea dioica X X Santalum album X X X Scolopia crenata X X Solanum indicum X X X X X X X X X X X X Spathodea campanulata X X X X X Syzygium cumini X X *As recorded between April 1 978 to July 1 979; no data for September 1 978. Almost all the trees except some Ficus fruit annually at Thekkady. Though the fruiting period varies from species to species and at times from one individual to another, there appear to be two peak periods of fruiting, April-June and November-December. However, during April- June 1979, comparatively few species of trees were recorded fruiting. This could be due to lower rainfall in the previous year, as the fruiting period of the same tree may vary from year to year due to rainfall and other climatic factors. During April-June Actinodaphne hookeri, Ficus gibbosa, F, tsiela, Grewia tiliciefolia, Machilus macrantha, Macaranga sp., Olea dioica, Santalum album, Scolopia crenata and Syzygium cumini were the main fruiting trees. During November-December, different species of Ficus were the main fruiting trees. Some other tree species also start flowering. From the flowers of Erythrina indica, Bombax ceiba and Spathodea sp., only M. viridis was seen sipping nectar. Among these, Erythrina flowers for an extended period of 5 months, mainly October to February, Bombax flowers from November to February and Spathodea mainly during June to August, though some trees were found flowering as late as November. Bischofia javanica fruits from November to January, whereas, Bridelia retusa fruits from August to November. Fruits of Lantana camara and Solatium indicum comprise the regular food of M. viridis. These plants fruit almost throughout the year. Leea indica, on which only M. viridis feeds, fruits for a long period of 8 months (May to December), some trees with a few fruits are found in other months also (Table 2). JOURNAL, BOMBAY NATURAL HISTORY SOCIETY, 97(1), APR. 2000 105 FOOD A ND FEEDING HA BITS OF INDIA N BA RBETS The fruit abundance in this region from April to June appears to be a reciprocal adaptation with the breeding season of local birds. Most of the resident birds breed during this period (Yahya 1988) and thus the chances of seed dispersal are maximum. Ficus trees provide the maximum quantity and variety of food to barbets. Ficus tsiela and F. retusci are more versatile and one or other of these species may be found fruiting throughout the year. However, no fruit was recorded on F. retusa in July-August. Ficus mysorensis and F. insignis were recorded fruiting during the rainy months, whereas F. gibbosa commonly fruits during drier months. F. hispida and F. glomerata, which were found fruiting invariably throughout the study area (the former at Thekkady and the latter at Lowercamp, Tamil Nadu) were never eaten by either species. At Sanjay Gandhi National Park, Mumbai M. zeylanica, were observed at times pecking at the ripe receptacles of F. glomerata , but never successfully, as the fruit fell down before the bird could pluck it. This could be due to the very weak peduncle of the ripe receptacle. However, near Churchgate, Mumbai, I found M. haemacephala pecking bit by bit on the semi-ripe receptacle of F. glomerata, but on no occasion did I find any barbet feeding on F. hispida). Comparatively few species of trees fruit during February and March at Thekkady. This could be due to the deciduous nature of the dominant species. During this period, almost all the trees shed their leaves, the rain is comparatively meagre, and most of the trees prepare for the forthcoming fruiting season. According to Champion and Seth (1968) the seasonal distribution of rainfall has a far- reaching influence on the nature of vegetation. Results and Discussion The ratio of consumption of animal and plant matter by M. viridis and M. rubricapilla is almost similar in every month (Fig. 1 & 2), except during the nesting period (March- July) for M. viridis which then consumes a larger quantity of animal matter. This could be due to the marked difference between the nestlings’ food in the two species (Yahya 1980, 1988). % plant & animal matter I % plant matter L 1 % animal matter Fig. 1: Monthly feeding pattern of M. viridis 106 JOURNAL, BOMBAY NATURAL HISTORY SOCIETY, 97(1), APR. 2000 FOOD AND FEEDING HABITS OF INDIAN BARBETS Apr May Jun Jul Aug Sep Oct Nov Dec Jan Feb Mar Apr 1978 I 1979 ■H % plant matter t I % animal matter Fig. 2: Monthly feeding pattern of M. rubricapilla Plant matter consumed by M. rubricapilla was restricted to fruits, and was about 8% more than that of M. viridis. However, the latter is a more versatile vegetarian, feeding on a wider range of vegetable matter. While M. viridis often feeds on the nectar of various flowers, M. rubricapilla was never found to do so. Though predominantly frugivores, both M. viridis and M. rubricapilla also feed on a large amount of animal matter, the former consuming about 10% more than the latter (Table 3). Animal food items consumed by M. viridis were larger in size and more diverse than those of M. rubricapilla. M. viridis feeds on earthworms (seen only twice) butterflies, dragonflies, mantids, cicadas, beetles, spiders, termites and caterpillars, whereas M. rubricapilla restricts itself to smaller caterpillars, borer larvae, termites and ants. Though the food preferences of these congeneric species are distinguishable, their feeding niches and food often overlap. Vegetame tood of M. viridis and M. rubricapilla As shown in Table 4, M. viridis and M. rubricapilla both show a preference for certain fruits in each month, but many fruits favoured by one species are frequently taken by the other also. Before analysing the data for a possible explanation of how these two congeneric sympatric species manage to coexist in the same habitat, a broad outline of their month-wise food items and preferences is given briefly. During January-February when only a Table 3 PERCENTAGE OF BARBETS FEEDING ON PLANT/ANIMAL MATTER Species No. of individuals Fruit/nectar Insects M. viridis (n = 3346) 2352 - 70.29% 994-29.71% M. rubricapilla (n = 1 889) 1485-87.61% 404-21.39% limited number of trees are fruiting, M. viridis very frequently forages on shrubs, while M. rubricapilla restricts itself to certain Ficus species. The common trees, on which the feeding of both species considerably overlaps during this period, are Ficus mysorensis , F. retusa, F. gibbosa, F. infectoria and F. tsiela. Among these, M. rubricapilla shows a much higher preference for/7, gibbosa, F. tsiala arid F. retusa, JOURNAL, BOMBAY NATURAL HISTORY SOCIETY, 97(1), APR. 2000 107 Tablk 4 PERCENTAGE OF M. VIRIDIS AND M. RUBRIC APILLA FEEDING ON FRUITS/NECTAR FOOD AND FEEDING HA BITS OF INDIAN BA RBETS Cl. < 04 t — t — O' T; O h OO On ^ O' — • o) 04 m ON r- ON tu NO o', oi o~, o 04 04 ON NO 04 o NO, 04 NO oc No! O) ioi O P no d co ^ OO OO OO 04 O Nt - r, it, - Tt — P P co 04 04 — O' N C d «o) • oo d NO OO 1 ON NO ON »n p — - no, — 04 — NO) 04 ioi O- ON — o ON 00 04 NO — < OO 4/S 04 — 04 04 O', CJ <D Q 1 o CO CO t — 1 1 I NO CO OO CO 04 in O CO 04 - O Tt O O 04 o OO CO © O' 40, lO, — OO CO CO — i d CO p- co d CO 04 CO CO CO — 04 04 — 04 O OO ON ON 04 OO p — ' 04 no. 1 i o. p p F. i r. > o z NO) ' 04 04 NO — o NT) ON CO Tf 04 in oo oo —nOoooo- — d d oi co oo — o- no) no, 0 — < CO NO 04 01 irl o - - 04 — Tj-ONO — r- P — ' OO - o ci o o — 04 — 04 04 NO, 010 O'. 1 p 04) 3 < CO P NO) CO O O', 04 NO) ON ON ON — — 04 vO oo — ON OO oo o — 04 0 4 © 1 NO, O' oo O' ON 04 OO NO O' O' NO) NO O'; NO 04 Nt O NO p — NO, o^ O' »oi p 04 ON O', — NO ON 1 04 1 O — ; ON — ; p no NO NO) Nr) Tl- NO O' O', 04 04 O' P tp NO ON O' P NO, NT) NO O' O 04 04 NO) OO O) oi oi oi cc 04 04 p 04 OO O) 04 NO, d »o] o'] O', 04 oc oi O' — as o4 04 O^ O' NO 04 04 NT) NO, _ ' O'. p O P as OO iTi O'. oo oo O', NT) P ok oi oi 04 * o. < oo — 00 p 01 p; p QQ .2 PS c OQ 04 N 6 — P O' NO O' P 04 O F oo ci - 04 04 Q <3 a? ■s Q a U 53 p kJ kj ’5 .00 Lr, cp k; g as k; .5 p k-i ■2 Ju 3 g § $ p p <4, 108 JOURNAL. BOMBAY NATURAL HISTORY SOCIETY. 97(1). APR. 2000 Table 4(contd.) * PERCENTAGE OF M. VIRIDJS AND M. RUBRICAPILLA FEEDING ON FRUITS/NECTAR FOOD AND FEEDING HABITS OF INDIAN BARBETS o <35 Os m> o ml EL i ■ in i 1 i m i , , , , so m < O mi m mi Os Os so m- c3 m, , , ■n- i i n — m — ; oo SO — i sd mi — ■ SO oo Os m~ Os X> ■ SO SO i • Os 1 1 1 i , 1 m mi m m o mi oo [Jh sd mi mi mi — m~ oo C , , , , , i i i sO 1 ''t sO a m m. r- *— > mi r- O m- oo o c-~ 1 OS OO a> 00 Os O Q mi — : d r- m- > m in ■ , , 1 m o o so p o Z O so r- m- bO , , ' Os , t 1 so o 3 r- o °o m> < mi o oo so t-~ oo , ! j , i i i 1 m 1 os m m* Os 3 o 00 m, m- mi o as m- r- C , , , , , i , o 1 o i i i , ' os m 3 m m mi m. — d SO OO , , , 1 m~ ' Os m m i i i , , 1 so Os t 1 — i in c? O os os m, m, oo £ mi mi mi mi mi d mi — '< 00 so m- , 1 m t — t~- i — r*~> i> 1 t"~ m, t 1 ^ 1 -rj- oo EL in m m r-; P so p ml o m> mi < mi SO mi mi mi mi mi mi m mi so r- .3 3 3 ■3 3 cL 3 1 cL C/3 .§ 3 3 g 3 3 g" 3 5 S c O oo W) C3 Loranthus s g .3 -5 1 a 3 5 a ‘o Olea dioica "a S 3 2 s cn £ 3 1 cn •S 3 3 .3 3 V "3 -3 3 CO- Syzygium cl Viscum sp. Ziziphus sp. C ’C ^ & <u a E 3 C P « t— a. JOURNAL, BOMBAY NATURAL HISTORY SOCIETY, 97(1), APR. 2000 109 *Notes: Upper figures in each column are for M. viridis and lower for M. rubricapilla. Percentage is calculated from the total number of birds observed feeding both on animal and plant materials. Remaining percentage is formed by birds feeding on animal matter. FOOD AND FEEDING HABITS OF INDIAN BARBETS 1 . Actinodaphne hookeri, 2. Ficus gibbosa, 3. F. retusa , 4. F. tsiela , 5. Grewia tiliaefolia, 6. Ficus mysorensis, 7. F. insignis. Fig. 3: Percent difference in common fruits consumed by a: M. viridis , b: M. rubricapilla whereas, both feed almost equally on F. infectorici. Comparatively, M. viridis prefers receptacles of F. mysorensis. In addition to feeding together with M. rubricapilla on Ficus trees, M. viridis frequently feeds on the fruits of Bischofia javanica, Leea indica, Lantana camara and Solanum indicum. During March, F. retusa and F. tsiela are the mam trees on which both species feed, M. rubricapilla far more than M. viridis. In April, several more species start fruiting. M. viridis sho\vs a greater preference for Actinodaphne hookeri which fruits from April to July. In addition to berries of shrubs, M. viridis feeds exclusively on Machilus macrantha. F. gibbosa and F. tsiela fruit during April, for whose receptacles M. rubricapilla always shows greater preference (Fig. 3a & b). From May to August, Grewia tiliaefolia, F. retusa and F. tsiela are the main fruiting trees on which both M. viridis and M. rubricapilla feed. During this period, both barbets show a higher preference for Grewia than for other fruits, though as usual Ficus trees are also visited freely and M. rubricapilla feeds more on Ficus receptacles than M. viridis. In addition, M. viridis also feeds on the fruits of Macaranga, Lantana, Solanum and on nectar of Erythrina; rarely also on nectar of Spathodea campanulata. The only fruits on which M. rubricapilla feeds exclusively are of Eucalyptus. During May to August, young fruits of Eucalyptus are quite often eaten by M. rubricapilla, mainly in the morning hours. M. viridis does not feed on Eucalyptus fruits, probably because this plant is recently (about 20 years earlier) introduced in the area. M. viridis shows higher preference for the fruits of Olea dioica, Scolopia crenata and Syzygium cum ini, which appear from April to May. Sandalwood drupes are exclusively eaten by M. viridis during April-May. During November-December M. viridis shows a higher preference for the figs of F. insignis, whereas M. rubricapilla feeds more frequently on F. gibbosa, F. retusa and F. tsiela ; both show almost equal preference for F. infectoria. Only M. viridis feeds on fruits of Solanum, Lantana and Leea indica, and nectar of Bombax ceiba and Erythrina. Very rarely, both feed on Evodea lunuankenda, Loranthus and Vis cum berries. Therefore, although both the species no JOURNAL, BOMBAY NATURAL HISTORY SOCIETY, 97(1), APR. 2000 FOOD AND FEEDING HABITS OF INDIAN BARBETS overlap on certain fruiting trees, almost every month M. viridis feeds exclusively on certain other fruits (such as Bridelia retusa, Carey a arborea, Lantana camara, Leea indica), thus reducing the extent of food competition. Possible reasons for food preference The primary reason for food preference in M. viridis and M. rubricapilla appears to be the size of the fruit. However, selection of food may also depend on various other factors such as colour, taste, nutritive value, and even on smell, as suggested by several workers. Figs of F. tsiela , F. retusa , and F. gibbosa are preferred by M. rubricapilla and those of F. mysorensis and F. insignis by M. viridis. The figs of the former group are about one-fourth the size of the latter two. M. rubricapilla shows less preference for the fruits of A. hookeri , G. tiliaefolia , Olea dioica , Scolopia crenata and Syzygiutn cumin i, which are larger than its favourite figs. The frequency of feeding in relation to fruit size is shown in Table 5. M. rubricapilla shows markedly higher preference for smaller fruits, though M. viridis also feeds on them. Low preference for larger fruits by M. rubricapilla can be correlated with its smaller beak. Correlation between the size of the food and beak has also been reported in the Galapagos ground finches by Lack (1971), and in British finches by Newton (1967). Vijayan (1975) also found that the whitebrowed bulbul ( Pycnonotus luteolus) with its slightly larger beak prefers bigger sized fruits than the coexisting redvented bulbul (P. cafer ). Though not analysed statistically, M. Table 5 PERCENTAGE OF M. VIRIDIS AND M. RUBRICAPILLA FEEDING ON FRUITS OF DIFFERENT SIZE Average size of the fruit <8 mm 8- 1 6 mm > 1 6 mm M. viridis (n = 2352) 37.45% 34.31% 28.24% M. rubricapilla (n = 1 485) 70.09% 1 9.90% 1 0.01 % zeylanica and M. haemacephala in San jay Gandhi National Park, Borivli, Mumbai (SGNP) and at Lowercamp, appeared to show remarkable food preference according to size; the former preferring figs of F. bengalensis and F mysorensis, whereas the latter always congregated in greater numbers on F. gibbosa, F. infectoria and F. religiosa. M. zeylanica was often found sipping nectar on Butea monosperma at SGNP, but M. haemacephala was never seen doing so. At Ranikhet (Uttar Pradesh) M. virens was recorded gulping pear blossom ( Pyrus sinensis ) conveniently owing to its large beak. Animal food of M. viridis and M. rubricapilla Insects of different groups comprise the main animal food of M\ viridis and M. rubricapilla. A month-wise record of animal food taken by these two species is shown in Tables 3 and 4 respectively. They usually hunt insects while following mixed hunting parties. However, during the breeding season both search for insects individually or in pairs. Quite often, both the barbet species were found capturing winged termites by short ‘flycatching’ sallies after light rain during March- April. Thesekhunts normally take place in groups; one such group of 30 M. rubricapilla was recorded hunting winged termites for 30 minutes at Thanikuddy area. All the birds were perched on a Terminal ia paniculata tree and caught the termites in the air one by one as they emerged from the ground. While the barbets were catching termites at about 16m height, swallows were also catching the termites much higher than the barbets, while red- whiskered bulbuls Pycnonotus jocosus were diving after them from bushes nearby. During April-May the teak defoliator, Hyblaea puera, swarm on young teak leaves and both barbets congregate in large numbers to feed on these caterpillars along with other birds. Except for this caterpillar, no swarming of any particular species was noticed during the study period at Thekkady. A Phalangid species was JOURNAL, BOMBAY NATURAL HISTORY SOCIETY, 97(1), APR. 2000 FOOD A ND FEEDING HABITS OF INDIAN BA RBETS found swarming in hundreds on some shrubs and tree trunks throughout the year, but no bird was seen to feed on them. Formation of Mixed Hunting Parties (MHP) At Thekkady, the formati on of large MHPs is a common avian activity. Generally bright hours of the day (Table 6) and comparatively open areas are selected for foiming a MHP. In the non- breeding season, both AT. viridis and M. rubricapilla commonly hunt with MHPs. A MHP sometimes follows a longer route, but normally limits itself to a circumference of c. 250 m or so. A ‘wave of birds’ as described by McClure (1972) was always noticed while observing the MHP. The difference between the flocks of insectivores and those assembled in a fruiting tree is that the insectivores’ ‘wave’ moves through the forest, while the frugivores confine themselves to a specific tree (McClure, 1972). Table 6 PERCENTAGE OF M. VIRIDIS AND M. RUBRICAPILLA FEEDING AT DIFFERENT HOURS OF THE DAY WITH MIXED HUNTING PARTY A VERAGE OF 1 1 MONTHS APRIL 1 978 TO APRIL 1 979 6 to 10 10 to 14 1 4 to 1 8 Total No. of hr. hr. hr. birds observed M. viridis 09.75 75.00 15.25 682*/994** M. rubricapilla 15.00 78.15 06.84 205*/404** * Number of birds seen hunting with MHP ** Total number of birds seen feeding on animal mater Position of barbets in MHP Normally, 10-12 bird species comprise a single MHP, but sometimes as many as 25 species were recorded, the commonest and perhaps the ‘nucleus’ of the party being drongos. The common species forming a MHP were usually the racket-tailed drongo ( Dicrurus paradiseus ), grey drongo (Z). leucophcieus ), bronzed drongo ( D . aeneus ), goldenbacked woodpecker {Din opium benghalense), goldenbacked threetoed woodpecker (D. javanense), common and southern tree pies ( Dendrocitta vagabundci, D. leucogastra ), common woodshrike (Tephrodornis virgatus ), jungle and hill mynas (Acridotheris fuscus, Gracula religiosa) minivets (Pericrocotus flammeus, P. cinnamomeus ), tits {Parus major, P. xanthogenys ) velvet-fronted nuthach (Sitta frontalis ) and various species of flycatchers. Barbets are opportunist members of the party, joining a passing MHP and hunting actively with the rest. AT. viridis being far more active than AT. rubricapilla exploits the maximum feeding zone. While ‘flowing’ with the wave, AT. viridis makes short sallies, glides down after insects or even lands on the ground, whereas AT. rubricapilla never descends below the secondary level. However, both peck on dry and dead tree trunks like woodpeckers, and at times on dry leaves, and pick up caterpillars. Intraspecific aggression between AT. viridis and AT. rubricapilla was not as common in a MHP as noted on fruit trees. This could be due to the marked difference in their feeding zones and larger feeding areas. On a fruit tree, especially when fruit is scarce, there is more rivalry and aggression — fight and chase — while in a MHP the food resource is always scattered. However, intraspecific aggression among AT. viridis itself is not uncommon. Aggression among other groups of birds in a MHP is also not as common as among a feeding flock of frugivores in a fruiting tree. However, racket-tailed drongos always try to dominate and chase other birds, even snatching morsels from them, as I have witnessed on several occasions. Competition for Food and Coexistence From the foregoing account, it appears that AT. viridis and AT. rubricapilla do not compete severely for food. However, they do overlap on certain fruiting trees or when hunting in a mixed hunting party of insectivores. As discussed below, 1 12 JOURNAL, BOMBAY NATURAL HISTORY SOCIETY, 97(1), APR. 2000 FOOD A ND FEEDING HA BITS OF INDIA N BA RBE TS the food competition is further reduced owing to their different feeding behaviour and overall dimensions. Feeding habitat Utilisation of different parts of the vegetation differs greatly between M. viridis and M. vubriccipiUa. The feeding zone is clearly distinguishable when both feed in a single microhabitat. During the study period, whether feeding on fruit or hunting insects, individually or with MHP, 82% M. rubricapilla were recorded feeding on the tertiary level, whereas only 51% M. viridis fed at that level. M. rubricapilla was seldom recorded descending below the secondary level (Table 7), while M. viridis frequently fed at the primary level or at times even on the ground, M. rubricapilla never does so. Table 7 PERCENTAGE OF M. VIRIDIS AND M. RUBRICAPILLA FEEDING AT DIFFERENT LEVELS M. viridis (n = 3346) M. rubricapilla (n = 1889) Ground 0.15% - Primary level, 1 to 4 m 18.42% - Secondary level, 4 to 8m 29.93% 17.06% Tertiary level, above 8m 51.50% 82.94% Ecological isolation by feeding heights has been reported in English titmice Parus major by Hartley (1953) and Gibb (1954). Vijayan (1975) suggested that the difference in the feeding zone is distinguishable in coexisting Pycnonotus cafer and P. luteolus at Point Calimere (Tamil Nadu) and plays a major role in isolating them ecologically. At Lowercamp, M. zeylanica , M. viridis and M. haemacephala were sometimes observed hunting together with a MHP. On those occasions, the feeding zones of the three species were always markedly different; M. zeylanica hunted at the topmost level, M. haemacephala mostly at secondary level, whereas M. viridis fed at the primary and secondary levels. Method of feeding The feeding methods of M. viridis and M. rubricapilla differ considerably, especially on larger fruits like the receptacles of Ficus mysorensis and F. bengalensis and other similar sized fruits. While M. viridis swallows the entire fruit, M. rubricapilla feeds by pecking and eating it bit by bit. The difference in feeding method is obviously due to the differences in their beak size. While M. viridis swallows larger fruits easily, M. rubricapilla cannot do so, and has to spend more time and energy on the same fruit. At Lowercamp, occasionally M. zeylanica , M. viridis , M. haemacephala and sometimes M. rubricapilla, were recorded feeding together on F. bengalensis and F. mysorensis. The feeding method of the two larger and two smaller 'pairs' was noted to be different: M. zeylanica and M. viridis with larger beaks normally swallowed the entire receptacles, whereas the other two (with almost equal beak size) fed by pecking at them bit by bit. Such a difference in method of feeding was recorded in unequal sized congeneric sympatric M. asiatica and M. haemacephala, and M lineata and M. haemacephala respectively in Calcutta Botanical Garden and in Valmiki Tiger Reserve (Bihar). Even while hunting insects individually or with MHP, M. viridis frequently catches cicadas, butterflies and such larger insects, whereas M. rubricapilla restricts itself to ants, small flies and termites. Feeding cycle Barbets are voracious feeders and can be seen feeding throughout the day. However, intensity of feeding activity varies during different hours of the day (Table 8). Both M. viridis and M. rubricapilla show maximum feeding activity during morning hours. M. viridis is comparatively less active around noon and more active in the afternoon. The difference in feeding cycle appears to be due to the differences in their roosting hours (Yahya 1987). On an 113 JOURNAL. BOMBAY NATURAL HISTORY SOCIETY. 97(1). APR. 2000 FOOD AND FEEDING HABITS OF INDIAN BARBETS average, M. rubricapilla roosts one hour longer than M. viridis, and hence is probably more active in the noon hours also, while M. viridis takes rest. After some rest M. viridis becomes more active and hence spends more time in feeding, while in the later afternoon M. rubricapilla spends more time in preparing to roost. Table 8 PERCENTAGE OF M. VIRIDIS AND A/. RUBRICAPILLA FEEDING AT DIFFERENT HOURS OF THE DAY ON DIFFERENT FRUIT TREES 6 to 10 10 to 14 14 to 18 lus hrs hrs M. viridis (n = 2325) 47.40 22.50 30.09 M. rubricapilla (n = 1485) 48.00 28.33 23.67 Similar results were obtained while observing M. zeylanica and M. haemacephala at Sanjay Gandhi National Park. M. zeylanica showed less feeding activity during noon hours whereas M. haemacephala was quite active during that period. M. haemacephala also roosts almost one hour longer than M. zeylanica (Yahya 1 987). Skutch (1944) also found the prong-billed barbet most active in the morning hours and least active at noon, when it rested for 1 to 2 hours. Aggression at feeding sites Intraspecific aggression is much more pronounced in M. viridis than in M. rubricapilla. The former is far more aggressive towards other species of birds as well. While feeding with frugivorous flocks, M. viridis chases almost all birds except the koel Eudynamys scolopacea. The koel was found to be the most dominant species and no other bird dared to fight it back. Intraspecific aggression at feeding sites might play some role in isolating two congeneric sympatric species and thus help in successful coexistence. Grubh (1979) concludes that intraspecific aggression at food plays an important role in successful coexistence of the Eurasian griffon Gyps fulvus, whitebacked vulture G. bengalensis and longbilled vulture G. indicus in Gir Forest: while the whitebacked is comparatively peaceful at feeding sites, the 1 14 other two spend considerable time quarrelling with their own kind, thereby indirectly permitting the weaker whitebacked to feed. Morphological adaptations for feeding In physical dimensions M. viridis and M. rubricapilla are different. The larger beak of viridis enables it to swallow larger fruits and insects, which rubricapilla cannot do. This could help them in reducing food competition and successful coexistence. Zacharias (1978) states that owing to the difference in overall size, the larger jungle babbler Turdoides striatus mostly feeds on larger insects while hunting together with whiteheaded babblers T. affinis. Another point which supports the view that the overall size difference in barbets may play an important role in their successful coexistence is the common occurrence side by side of two species of different sizes. During my study I found M. viridis and M. rubricapilla occurring together at Thekkady; M. zeylanica and M. haemacephala coexisting at Sanjay Gandhi National Park, Hazaribagh National Park and at the Betla Tiger Reserve; M. lineata and M. haemacephala coexisting in Valmiki Tiger Reserve and Corbett National Park, and M. asiatica and M. haemacephala in Calcutta City. All these coexisting congeneric species have the same remarkable differences in size. Hinde (1959) suggested that the morphological differences between coexisting species are not merely adapted to feeding methods, but largely determine them. The degree of dominance while feeding may also vary according to the body size as reported by Grubh ( 1 979) among different species of griffon vultures - the largest (Eurasian griffon) was found to be the most dominant and the smallest (whitebacked) the least. Conclusion Though fruits of different species of plants constitute the main food of barbets, both M. viridis and M. rubricapilla feed on insects to JOURNAL, BOMBAY NATURAL HISTORY SOCIETY, 97(1), APR. 2000 FOOD A ND FEEDING HA BITS OF INDIA N BA RBETS a considerable extent. The former consumes about 30% animal matter, whereas the latter consumes about 20%. M. viridis consumes more insects during the breeding season than M. rubricapilla. Only M. viridis sips nectar from flowers. Both species often hunt together with mixed hunting parties of insectivores; M. viridis always joins the party in larger numbers and for longer periods. During March- April, after light showers, both the species hunt winged termites in groups; sometimes this single-species group may consist of as many as 30 individuals. Food competition between the coexisting M. viridis and M. rubricapilla is not severe, for they normally procure food from different feeding zones. Intraspecific aggression among M. viridis is markedly more while feeding either on a fruit tree or with a mixed hunting party of insectivores which reduces its competition for food with M. rubricapilla to some extent. Another factor responsible for ecological isolation in feeding behaviour is the varying heights from which they exploit food: while M. viridis feeds at primary, secondary and tertiary levels, and at times lands even on the ground, M. rubricapilla restricts itself to the secondary and tertiary levels. The study also supports Huxley’s (1942) postulation that “big size difference between congeneric species of birds are means of ecological isolation”. Based on the data collected in the present study, it could be added that since food is the primary requirement of an animal, for the successful coexistence of two closely related species in a single habitat, divergent morphological adaptations in relation to feeding habits are an outcome of the process of natural selection. Refer Ali, Salim, (1935): The ornithology of Travancore and Cochin (with notes by Hugh Whistler) Part 1 . Bombay nat. Hist. Soc. 37: 814-843. Champion, H.G. & S.K. Seth (1968): A revised survey of the forest types of India. Govt, of India Press, New Delhi. Gibb, John. (1954): Feeding ecology of tits, with notes on Barbets are economically important and play a significant role in controlling various harmful insects, in cross-pollination and seed dispersal of trees. Though they are presently common in many places, their conservation priorities should be anticipated by wildlife biologists and managers to maintain sustainable populations of different species. Acknowledgements I am greatly indebted to the late Dr. Salim Ali for his guidance, constant interest and constructive criticism at various stages of the study. I am grateful to the Bombay Natural History Society for supporting the study under Salim Ali — Loke Wan Tho Ornithological Research Fund. Mr. J.C. Daniel, Mr. S.A. Hussain, Dr. Robert Gmbh and Dr. V.S. Vijayan kindly visited the study area and provided valuable suggestions. Dr. Mohammed Ali Raza Khan and Dr. Priya Davidar were always source of encouragement throughout the study. I thank Mr. K.K. Nair, Ex. CCF (Kerala) for permission for field study, and Mr. S.N. Asari, Field Director of Periyar Tiger Reserve and his subordinate officers for help. Dr. P.S. Easa, Dr. M. Balakrishnan, Dr. Lalitha Vijayan, Dr. K.K. Ramachandran and other members of Kerala Forest Research Institute were very cooperative while working at Thekkady. I thank Babu and Kumaran (local assistants) for their tireless help during the fieldwork. I am obliged to my colleagues Drs. Asad R. Rahmani, Salim Javed and Satish Kumar for peer reviewing the draft and helping in preparing the diagrams. ENC ES Tree creeper and Goldcrest. Ibis 96: 5 1 3-543. Grubh. R.B. (1979): Competition and Co-existence in Griffon Vultures. J. Bombay nat. Hist. Soc. 75: 810- 814. Hartley, P.H.T ( 1 953): An ecological study of the feeding habits of the English Titmice. J. Anim. Ecol. 22: 261-288. JOURNAL, BOMBAY NATURAL HISTORY SOCIETY. 97(1), APR. 2000 115 FOOD A ND FEEDING HA BITS OF INDIA N BA RBE TS Hinde. R.S. (1959): Behaviour and speciation in birds and lower vertebrates. Bird Rev. 34: 85-128. Huxley, J.S. (1942): Evolution: The Modern Synthesis. George Allen & Unwin Ltd. London. Lack, D. (1971): Ecological Isolation in Birds. Blackwell Scientific Publications. Oxford and Edinburgh. McClure, E.H. (1972): Two Tropical forests and their birds. J. Bombay nat. Hist. Soc. 71: 517-535. Newton, I. (1967a): The adaptive radiation and feeding ecology of some British Finches. Ibis 109: 33-78. Rjpley, S.D. (1961): A Synopsis of the Birds of India and Pakistan. Bombay Natural History Society, Bombay. Robertson, A. & M.C.A. Jackson (1992): Birds of Periyar — An aid to Birdwatching in the Periyar Sanctuary, Kerala. Tourism & Wildlife Society of India. Simmons, K.E.L. (1970): Ecological determinations of breeding adaptation and social behaviour in two fish-eating birds. Social behaviour in Birds & Mammals. Edited by Crook. H.J. The Whitefriars Press Ltd., London & Tonbridge. Skutch, A.F. ( 1 944): The life-history of the Prong-billed Barbet. Ank 61: 61-87. Vijayan, L. (1 984): Comparative biology ofDrongos with special reference to ecological isolation. Ph.D. thesis, Dept, of Field Ornithology, Bombay Natural History Society, Bombay. Vijayan, V.S. (1975): Ecological isolation in Bulbuls (Family Pycnonotidae. Class Aves) with special reference to Pycnonotus cafercafer (Linnaeus) and P. luteolus( Lesson) at Point Cali mere. Tamil Nadu. Ph.D. Thesis. University of Bombay. Yahya, H.S.A. (1980): A comparative study of ecology and biology of Barbets. Megalaima spp. (Capitonidae: Piciformes) with special reference to Megalaima viridis (Boddaert) and M. rubricapilla malabarica (Blyth) at Periyar Tiger Reserve. Kerala. Ph.D. thesis, University of Bombay. Yahya, H.S.A. (1982): Observation on the feeding behaviour of barbets, Megalaima spp. in Coffee estates of South India. J. Coffee Research 12(3): 72-76. Yahya, H.S.A. (1987): Roosting behaviour of barbets, Megalaima spp. In: Recent Trends in Ethology Ed. M. Balakrishnan & K.K. Alexander Ethological Society of India, Bangalore, pp 101-106. Yahya, H.S.A. (1988): Breeding biology of barbets, Megalaima spp. with special reference to M. viridis and M. rubricapilla malabarica at Periyar Tiger Reserve, Kerala. J. Bombay nat. Hist. Soc. 85(3): 493-511. Yahya, H.S.A. ( 1 989): Habitat preference of birds in the Periyar Tiger Reserve, Kerala. Indian Journal of Forestry 12(4): 288-295. Yahya, H.S.A. (1991 ): Drinking and bathing behaviour of barbets, Megalaima spp. J. Bombay nat. Hist. Soc. 88(3): 454-455. Zacharias. V.J. (1978): Ecology and biology of certain species of Indian Babblers ( Turdoides spp.) in Malabar, Ph.D. Thesis, University of Calicut, Kerala. ■ H ■ 116 JOURNAL. BOMBAY NATURAL HISTORY SOCIETY. 97(1). APR. 2000 NEW DESCRIPTIONS SPINY EELS OF THE GENUS MACROGNATHUS LACEPEDE FROM MANIPUR, WITH DESCRIPTION OF A NEW SPECIES' L. Arunkumar and H. Tombi Singh2 ( With four text-figures ) Key words: Macrognathus morehensis sp. nov., Yu drainage system, Manipur. The paper gives a systematic account of two species of Macrognathus, hitherto known from Manipur, viz. M. aral (Bloch & Schneider) and M. pancaius Hamilton-Buchanan, which are distributed in the Barak drainage system and in the hill streams of Manipur respectively. A new species M. morehensis occurring in Manipur has been described here. It is found in the southeastern corner of this state and the adjoining areas of Myanmar, drained by the Yu drainage system, known as the Chindwin of Meaner. The diagnostic feature of M. morehensis is the combination of the following distinctive characters: 1 1 to 16 dorsal spines, 20 to 25 broad black transverse bars, 12 to 14 black spots that form imperfect ocelli at the base of dorsal fin rays, 6 black oval spots at the base of dorsal spines, 10 to 13 black spots at the base of anal fin rays, 5 to 7 oblique striations of black dots arranged in longitudinal parallel rows at the dorsal and anal fin rays, 7 to 10 black lines of striations formed by the dots at the caudal fin, a single ocellus at base of caudal fin, 8 to 11 rostral tooth-plates, and 76 vertebrae. Introduction Manipur is an isolated hill state in the northeast corner of India having three drainage systems: the Barak, Manipur, and Yu drainage systems draining the western, central and eastern water bodies respectively (Fig. 4). The Barak drainage system is connected with the Barak- Brahmaputra river system of India, whereas the Manipur drainage and the Yu drainage systems are connected with the Chindwin river system of Myanmar. Hora (1921) described a new species of spiny eel, Mastacembelus manipurensis from Khurda (Khordak) stream of Manipur and Rhynchob della dhcinashorii from Dhanashori stream, about a mile from Dimapur, Assam. Menon (1954), while reporting on the fishes known from Manipur, listed two spiny eels, viz. M. armatus and M. manipurensis. Later, Menon (1974) considered M. manipurensis and ‘Accepted August, 1997. department of Life Sciences, Manipur University, Canchipur795 003, Imp'hal, Manipur, India. R. dhanashorii as synonyms of M. armatus and Macrognathus aculeatus respectively. Presently, three species, viz. M. aral (Bloch & Schneider), M. guentheri (Day) and M. pancaius Hamilton- Buchanan have been reported from Indian waters (Talwar and Jhingran, 1991). No further report is available on the spiny eels of the genus Macrognathus of Manipur. Recently, several specimens of Macrognathus were obtained from the Lokchao river and the Maklang river of the Yu drainage system of this state near Moreh, which is known as Chindwin of Meaner. From this collection, a new species, Macrognathus morehensis , is described here. Material and Methods Fishes were collected using different types of nets, grooping, dewatering of shallow water pockets and with the help of local fishermen. Some fishes were also purchased from Moreh Bazar, Chandel dist. , Manipur, near the Indo- Myanmar border. In the field, their local names and fresh colours were noted.The fishes were JOURNAL , BOMBAY NATURAL HISTORY SOCIETY, 97(1), APR. 2000 NE W DESCRIP TIONS then preserved in 10% formaline and brought to the Fishery Laboratory of Manipur University. The fishes were identified with reference to Day (1889), Hamilton-Buchanan (1822), Roberts (1980, 1986) and, Talwar and Jhingran (1991). The specimens were deposited in the Manipur University Museum of Fishes (MUMF). Registration numbers are given below. Results Macrognathus aral (Bloch & Schneider) (Fig. 1) Fig. 1: Macrognathus aval (Bloch and Schneider) Rhynchobdella dhcinashorii Flora, 1921. Rec. Indian Mus., XXII : 205, PI. IX fig. 2 (sp. nov.). Macrognathus aral Roberts, 1980. Copeia, 3: 385-391, fig. 1 2b (Revision). Macrognathus aval Talwar & Jhingran, 1991 Inland Fish. India & Adjacent Countries , 2\ 1026 (Distribution extended). Manipuri name: Ngaril/Jirigi ngaril pokch aob i/Ngari l yangmitpanbi. Material examined: 3 exs. Uncat. MUMF. 1 from Jiri River; 120 mm total length; 7. viii. 1 995, 2 ex. Makru stream; 124 to 135 mm total length; 2.ix.l985, coll. M.G. Sharma, 1 ex. MUMF 20 1/1 A, Jiri River, 205 mm total length, 16.x. 1992, coll. L.A. Distribution: Manipur: Barak drainage system. Remarks: Formerly reported as M. aculeatus and distributed strictly in the western sides of this state, drained by the Barak drainage system of the Brahmaputra system in India. It is easily distinguished from M. aculeatus by the lack of 14 to 17 oblique dark bars on the body and smaller number of rostral tooth-plates (18 to 21 vs. 38 to 55). Roberts (1980) stated that M. aculeatus was known from the southern half of the Malay Peninsula: several of the principal rivers of Sumatra; the Kapaus river of Borneo and northern Java as far east as the Brantas river. The specimens (M. aral) from the Barak drainage of Manipur are similar to M. siamensis (Roberts 1980) in the presence of ocelli at the base of dorsal fin rays, but can be easily distinguished by the lack of ocelli at caudal fin, smaller number of rostral tooth-plates ( 1 8 to 2 1 vs. 7 to 14), and total number of vertebrae (71 vs. 75). Macrognathus pancalus Hamilton-Buchanan (Fig. 2) Fig. 2: Macrognathus pancalius Hamilton-Buchanan Macrognathus pancalus Hamilton- Buchanan, 1822. Fish Ganges, 30, 364. pi. XXII, fig. 7. Mastacembelus pancalus Sufi, 1956 Bull. Raffles Mus., 27: 93-146 (Revision). Macrognathus pancalus Talwar & Jhingran, 1991 Inland Fish. India & Adjacent Countries , 2: 1027-1028, Fig. 292. Manipuri name: Ngaril/Ching-ngaril- macha. Material examined: 3 exs. MUMF 202/ 3A, 1 ex. Jiri River; 1 1 1 mm total length; 1 3 .xii. 1 990, 1 ex. Litan stream at the root of Thoubal river; 132 mm total length; 15.xi.1991 and 1 ex. Maklang river; 124 mm total length; 8. xii. 1992, coll. L.A. Distribution: Manipur: Hill streams and 118 JOURNAL, BOMBAY NATURAL HISTORY SOCIETY 97(1) APR. 2000 NE W DESCRIP TIONS rivers of the Barak drainage, the upper and lower regions of Manipur drainage and the Yu drainage system. Remarks: It is the smallest among spiny eels and mainly found in hill streams. A distinct streak of longitudinal spots runs along the lateral line from the eye to the base of caudal fin in the present specimen, with 65 to 66 vertebrae. Roberts ( 1986) mentioned that it belongs to the second group of Macrognathus which lack rostral tooth-plates. Macrognathus morehensis sp. nov. (Fig. 3) Fig. 3: Macrognathus morehensis sp. nov., MUMF 203/8 A Paratype, 147 mm TL, Manipur: Yu drainage system. Holotype: MUMF 203/1 A, fromMaklang river near Moreh Bazar, Chandel district, 155 mm total length 17.x. 1992. Coll. L.A. Paratypes. MUMF 203/8A, 3, from Lokchao river near Moreh Bazar. 5 from Moreh Bazar; near Indo-Myanmar border, 80 to 147 mm total length. 2 1.x. 1992. Coll. L.A. Manipuri name: Ngaril/Ngamoi-tup/ Ngamu-tup/Tou-ngaril Diagnosis: A Macrognathus has the distinctive combination of the following characters: (i) 1 1 to 16 dorsal fin spines, (ii) 20 to 25 black broad transverse bars on the body, (iii) 8 to 11 rostral tooth-plates, (iv) 12 to 14 black spots that are imperfect ocelli at the base of dorsal fin rays, (v) 10 to 13 distinct black spots at the base of anal fin rays, (vi) 6 black oval spots at the base of dorsal spines, (vii) 5 to 7 oblique striations of black dots arranged in parallel longitudinal rows at the dorsal and anal fin rays, (viii) 7 to 10 black lines of striations formed by dots at the caudal fin, (ix) a single ocellus at the base of caudal fin and (x) 76 vertebrae. Description: Br. 3-5, D. 11-16/39-51, P. 15-20, A. 3/40-54, C. 11-14. Body slightly compressed. Rostrum slightly rounded. Pre- orbital and pre-opercular spines absent. Head long and pointed. Mouth inferior, cleft of mouth narrow. Ventral side of snout transversely striated by 2 to 4 black bars. Eyes not visible from ventral side and covered by a thin membrane. Eye diameter more or less same as the interorbital distance. Lips thin. No gill rakers. Caudal fin distinctly separated from the dorsal and anal fins. Scales are minute. The third anal spine is very near the origin of anal soft fin rays and difficult to identify, since it is buried inside the skin. Proportional measurements of holotype and paratypes (the latter in parenthesis): Depth of body 10.97 (11.11-12.93), height of head at eye 4.51 (4.08-6.25), height of head at occiput 6.45 (6.25-7.69), length of head at occiput 12.90 (11.11-15.47), length of head at the end of lateral operculum 17.42 (17.36-22.22) and length of caudal fin 7.09 (6.94-9.52) in the percentage of total length respectively. Depth of body 1 1 .80 (1 1 .94-13.79), height of head at eye 4.86 (4.41-6.84), height of head at occiput 6.94 (6.72-8.33), length of head at the end of lateral operculum 18.76 (18.65- 24.33), length of pectoral fin 6.94 (7.14-8.62), predorsal length at the origin of dorsal fin spine of 43.85 (43.29-46.72), and predorsal length at the origin of dorsal fin soft rays 63.29 (62.1 1- 69.93) in the percentage of standard length respectively. Diameter of eye 33.33 (33.33-35.59), interorbital distance 33.33 (33.33-35.95) and width of mouth 22.22 (20.00-25.00) in percentage of length of snout respectively. Colour: Body light yellowish to ashy. In young stages (81-120 mm total length), the dorsal fin soft rays, anal fin soft rays and caudal JOURNAL, BOMBAY NATURAL HISTORY SOCIETY, 97(1), APR. 2000 119 NEW DESCRIPTIONS Fig. 4: Map of Manipur showing distribution of Macrognathus fins are distinctly red with black striations and pin dotted transverse bars along their entire length. Abdomen yellowish white. Habitats: M. morehensis has a habit of actively burrowing in the swampy and vegetated stream bed. The inhabitants of Kwatha village near Moreb, of the Indo-Myanmar border call it ‘ Tou-ngariV , according to the habitat of the fish. The fish hides under pebbles, stones, gravel, sandy beds of clear torrential stream. It makes small pits and holes in the swampy arid vegetated beds of stagnant streams and waterbodies. The fish is associated with Amblypharyngodon molci , Aspidoparia morar , Badis badis , Chanda nama , Chela laubuca , Colisa fasciata , Danio aequipinnatus , Esomus dancricns , Garni gravelyi , 120 JOURNAL, BOMBAY NATURAL HISTORY SOCIETY. 97(1) APR. 2000 NEW DESCRIPTIONS G. lissorhynchus , G. rupecola, Glyptothorax pectinoptems , G platypogonoides , G. trilineatus, Mystus bleekeri , M. cavasius , Nemacheilus vinciguerrae and Parluciosoma dciniconius. Its distribution extends upto Tumu of Myanmar. Remarks: M. morehensis is easily distinguished from M. guentheri (Day) and M. pancalus Hamilton-Buchanan by the presence of rostral tooth-plates. It is also easily distinguished from M. aral (Bloch & Schneider) by the presence of less rostral tooth-plates (8-11 vs. 14-28), dorsal fin spines (11-16 vs. 16-23), total number of vertebrae (76 vs. 7 1 ), coloration (indistinct i.e. imperfect ocelli vs. distinct ocelli at base of the dorsal soft fin rays, and an ocellus at base of caudal fin vs. absent) and specific distribution, viz. Yu drainage system vs. Barak drainage system of Manipur. Etymology: The specific name is derived from Moreh, the type locality of the fish. Discussion: Roberts (1980, 1986) stated that M. aculeatus was known strictly from the southern half of the Malay Peninsula. He synonymised and referred all the formerly well known species of M. aculeatus which are distributed in India to M. aval. M. aculeatus had not been found in Myanmar or in the Indian subcontinent, but was found in Thailand at Surat Thani, Chiao Lam and the Tapi River basin. M. morehensis can be easily differentiated from M. caudiocellatus, M. circumcinctus , M. semiocellatus and M. zebrinus by the absence of preopercular and preorbital spines, and presence of rostral tooth-plates. M. morehensis differs from M. aral in having a smaller number of rostral tooth-plates (8-11 vs. 14-28), fewer dorsal fin spines (11-16 vs. 1 6-23), more vertebrae (76 vs 7 1 ) and pattern of bands (20 to 25 transverse dark bars vs. 2 pale longitudinal stripes along its entire length). M. morehensis differs from M. aculeatus in having fewer rostral tooth-plates (8-11 vs. 29- 55) and numbers of oblique transverse bars on the body (20-25 vs. 14-17). M morehensis differs from M. meklongensis in having a smaller number of dorsal fin rays (39- 51 vs. 50-54), pectoral fin rays (15-20 vs. 22-23), caudal fin rays (11-14 vs. 16-19), coloration of black spots at dorsal fin base (12-14 distinct large spots which are imperfect ocelli vs. no ocelli or 10-12 faint small ocelli), rim of anterior nostril with finger-like projections (absent vs. 6) and transverse bars on the body (20-25 vs. absence of transverse bars). M. morehensis differs from M. siamensis in the presence of fine dark striations in the caudal fin (7-10 fine black striations formed by dots vs. absence of striations), ocelli in the dorsal fin base (12-14 black spots which are imperfect ocelli vs. with series of large and distinct form of ocelli), fine oblique striations in the dorsal soft fin-rays (5-7 vs. absent) and distribution (Manipur vs. Thailand and Kampuchea). M. morehensis has a restricted distribution in Moreh, near the Indo-Myanmar border, Chandel Dist., Manipur at lower portion of Lokchao river, Pumpum stream of Kwatha, Lairok Maru, lower portion of Maklang river, which belong to the Yu drainage- system of Manipur, extend to Myanmar and join the Chindwin river. The fish was also collected from the adjoining areas of Manipur-Myanmar border of the Yu river system, which is known as the Chindwin of Meaner. According to Kottelat (1989) and Zakana- Ismail (1994), M. aral, M. caudiocellatus and M. zebrinus are the Salween elements of fishes. Kottelat (loc. cit.) described M. aculeatus as the Malay peninsular element of fish and M. circumcinctus, M. meklongensis, M. semiocellatus and M. siamensis as the Thailand elements of fishes. According to Zakaria-Ismail (loc. cit.) M. aculeatus, M. caudiocellatus, M. circumcinctus, M. meklongensis, M. semiocellatus and M. siamensis belong to the Indo-Chinese elements of fishes. M. pancalus is the true Indian element of fish. Hence M. morehensis is a distinct species with JOURNAL BOMBAY NATURAL HISTORY SOCIETY. 97(1), APR. 2000 121 NEW DESCRIPTIONS meristic, morphometry, anatomy (vertebrae), colour pattern and specific distributional areas, i.e. the Chindwin of Meaner as the defining features. ACKNOWLEDGEM ENTS The first author thanks Dr. W. Vishwanath Refe Day, F. (1889): The Fauna of British India, including Ceylon and Burma, Fishes, 2, 509. Taylor and Francis, London. Hamilton-Buchanan, F. (1822): An account of the fishes found in the river Ganges and its tributaries, vii, 1- 405, 39 pis. Edinburgh and London. Hora, S.L. ( 1 92 1 ): Fish and fisheries of Manipur with some observations on those of Naga Hills. Rec. Indian Mus. 22(3): 165-214. Kottelat, M. (1989): Zoogeography of the fishes from Indo-Chinese inland waters with an annotated checklist. Bull. Zool. Mus. 12: 1-55. Menon, A.G.K. (1954): Further observations on the fish fauna of Manipur state. Rec. Indian Mus. 25(1): 21-26. Menon, A.G.K. (1974): A check-list of fishes of the Himalayan and the Indo-Gangetic plains. Inland of Manipur University, Dr. Maurice Kottelat of Switzerland (CMK), Dr. Kelvin K.P. Lim, and Dr. Peter K.L. Ng of National University, Singapore and Dr. Mohd. Zakaria Ismail of University of Malaya for help. We also thank UGC, Special Assistance Programme, Life Sciences Department, Manipur University for financial assistance. E n c e s Fisheries Society of India. Special Publications. 1. pp 136. Roberts, T.R. ( 1 980): A revision of the Asian Mastacembelid fish genus Macrognathus. Copeia 3: 385-391 . Roberts, T.R. (1986): Systematic review of the Mastacembelidae or Spiny eels of Burma and Thailand, with description of two new species of Macrognathus. Jap. J. Ichthyol. 33(2): 95-109. Sufi, S.F.K. ( 1 956): Revision of the Oriental fishes of the family Mastacembelidae. Bull. Raffles Mus. 27: 93- 146. Talwar, P.K. & A.G. Jhingran: Inland Fishes of India and Adjacent Countries. 2, Oxford & IBH Publishing Co. Pvt. Ltd., Calcutta. 543-1 1 58. Zakaria-Ismail, M. (1 994): Zoogeography and biodiversity of the freshwater fishes of Southeast Asia. Hydrobiologia. 285: 41-48. 122 JOURNAL. BOMBAY NATURAL HISTORY SOCIETY. 97(1) APR. 2000 THREE NEW GENERA OF WHITEFUES MOHANASUNDARAMIELLA, SHANTHINIAE AND VA SANTHARAJIELLA (ALEYRODIDAE : HOMOPTERA) FROM INDIA1 P. Manidurai Manoharan David- ( With three text-figures) Key words : Whiteflies, Aleyrodidae, Homoptera, Mohanasundaramiella , Shanthiniae, V cisantharajiella In a survey conducted in the southern districts of Tamil Nadu and the adjoining forests of Kerala during 1991-1994, 84 species of aleyrodids under 34 genera were collected and studied. Of the 34 genera three, viz., Mohanasundaramiella, Shanthiniae and Vasantharajiella were found to be new and Genus Mohanasundaramiella gen. nov. Type-species: Mohanasundaramiella rubiae sp. nov. Pupal case oval in outline, pale with no wax secretion; margin lobulate to serratulate; marginal setae present; tracheal pores and combs absent; submargin characteristically separated from dorsal disc by a well defined furrow; submarginal transverse ridges and furrows conspicuous; subdorsal fold-like suture extending between meso-metathoracic suture and third-fourth abdominal suture; submarginal setae present; cephalic, mesothoracic, metathoracic, first abdominal, eighth abdominal and caudal setae present; first abdominal setae located late rad of subdorsal fold- like suture. Vasiform orifice subcordate, operculum filling orifice, lingula tip exposed but included. Caudal furrow and ridges absent. Tracheal folds discernible. Diagnosis. This genus is strikingly different from the known genera of Aleyrodini in having a well defined submarginal furrow that distinguishes submargin from dorsal disc. It resembles some species of Crenidorsum Russell in the furrow in inner subdorsal area of 'Accepted August, 1 999 department of Agricultural Entomology, Agricultural College & Research Institute, Killikulam, Vallanad 628 252, Tamil Nadu, India. are described and illustrated. cephalothorax and abdomen, but differs from them in the presence of first abdominal setae and conspicuous submargin. Other distinguishing characters include presence of fewer than 19-21 pairs of dorsal setae that separate it from Aleuromarginatus Corbett, presence of first abdominal setae on subdorsum that are absent in Aleyrodes Latreille, presence of submedian meso- and metathoracic setae that are lacking in Aleurocybotus Quaintance & Baker, oval shape of pupal case that is typically elongate, parallel-sided and slightly square anteriorly and posteriorly in Aleurotulus Quaintance & Baker, and presence of minute submarginal setae that do not occur in Aleurotrachelus Quaintance & Baker. Etymology: This genus is named in honour of Dr. M. Mohanasundaram, Professor of Agricultural Entomology, Tamil Nadu Agricultural University, Coimbatore, the renowned acarologist, who taught the author the science of taxonomy, and suggested this study. Mohanasundaramiella rubiae gen. et sp. nov. (Fig. 1) Pupal case: Oval in shape. 1.03-1.05 mm long and 0.87-0.89 mm wide, widest across abdominal segment III. Pale white with no wax secretion. Living on either surface of leaves. JOURNAL, BOMBAY NATURAL HISTORY SOCIETY, 97(1), APR. 2000 123 NEW DESCRIPTIONS Fig. 1 : MohanasundaramieUa rubiae gen. et sp. nov. : A. pupal case; B. prothoracic leg and antenna; C. margin; D. vasiform orifice; E. disc pore and porette Margin; Lobulate to serratulate, about 14 teeth occupying 0.1 mm length of margin; teeth each longer than wide; margin slightly indented at cephalic and caudal ends medially and at thoracic tracheal pore areas. Tracheal pores and combs wanting. Anterior marginal setae 18.6 pm long, posterior marginal setae 38.0 pm long. Dorsum: Dorsal disc separated from submargin by a distinct furrow. Submargin rather uniformly wide. Well defined transverse ridges 124 JOURNAL, BOMBAY NATURAL HISTORY SOCIETY. 97(1) APR. 2000 NE W DESCRIPTIONS and furrows running mesad from margin to outer subdorsum; adjoining 2-3 furrows more sclerotized, alternating every 3-4 less prominent ridges and furrows. Minute submarginal setae m 8 pairs, 6 on cephalothorax and 2 on abdomen, 6.2-12.4 pm long. Dorsal disc with numerous disc pores and porettes, one row in submarginal fold, one each at the darker transverse furrows. Transverse moulting suture reaching margin, curved caudad from its midpoint, terminating slightly anterior to the first-second abdominal suture at the longitudinal subdorsal fold-like suture which characteristically extends between meso-metathoracic suture and third-fourth abdominal suture. Meso-metathoracic suture much pronounced. Abdominal segmentation distinct. Abdominal segments IV- VI with a weak rachis, their sutures running lateral into subdorsum. Median length of abdominal segment VII slightly shorter than that of VI. Submedian cephalic setae long, tapered with a prominent base, 86.8 pm long. Two pairs of submedian thoracic setae present: one pair on mesothorax 18.6 pm long, the other on metathorax 34.1 pm long. First abdominal setae transpositioned on subdorsum laterad of longitudinal fold-like suture, 6.2 pm long. Eighth abdominal setae 65. 1 pm to at least 114.7 pm long, their bases anterocephalad of vasiform orifice. Caudal setae 28.0-74.4 pm long, located on submargin. Submedian abdominal depressions weak. Vasiform orifice subcordate with a broader rim, 46.5 pm long, 62.0 pm wide, posterior margin notched inside. Operculum of identical shape, filling three-fourths of orifice, 34.1 pm long and 46.5 pm wide. Lingula tip setose, exposed but included. Caudal furrow and caudal ridges absent. Venter: Thoracic tracheal folds faint, caudal fold distinct. Anterior thoracic and posterior abdominal spiracles evident. Adhesive sacs present. Antennae reaching anterior thoracic spiracles, their tips with a finger-like projection. Ventral abdominal setae 37.2 pm long. Host: Morindci sp. (Rubiaceae) Holotype: One pupal case mounted on slide, on Morindci sp., india: Kerala: Walayar forest, 24.ix.1992, coll. M. Mohanasundaram (No. 58 A.I.). Paratypes: Six pupal cases on slides, same data as holotype. Etymology: Species name derived from Rubiaceae, the host plant family. Remarks. Two out of the seven pupal cases are parasitised. Genus Shanthiniae gen. nov. Type-species: Shanthiniae sheryli sp. nov. Pupal case rather uniquely polyhedral in outline with lateral evaginations at six places on either side; margin very finely crenulate and crenate-looking, margin at pore area cleft; marginal setae present; thoracic tracheal pores well defined, inset with a single not very conspicuous tooth; dorsum ornamental with spots, reticulations; transverse moulting suture reaching margin; bases of cephalic, first abdominal and eighth abdominal setae distinct, caudal setae not discernible; abdominal segments VI and VII equally long, porettes with typically sclerotized rims; vasiform orifice cordate, with posterior margin toothed, operculum filling the orifice, lingula concealed; caudal furrow and ridges distinct; thoracic and caudal tracheal folds defined. Diagnosis: Pupal case outline of this genus is extraordinarily unique in being a 12-sided polyhedron with six corners on either side. This shape is not seen in any other whitefly species or genera of the world. However, Shanthiniae is related to Dialeurodes Cockerell, Dialeurolonga Dozier, and Dialeuronomanda Quaintance & Baker in the distinct structure of thoracic tracheal pore with tooth and in the presence of a comb of teeth in the inner margin of vasiform orifice, especially posteriorly. It shows affinity to Dialeurolonga in lacking subdorsal or JOURNAL BOMBAY NATURAL HISTORY SOCIETY. 97(1), APR. 2000 125 NEW DESCRIPTIONS submarginal row of setae but can be distinguished by the absence of small papillae in a row on the submarginal area. Its abdominal segments I-IV are subequal in length, similar to those in Dialeurodes, but are distinctive in the absence of a row of subdorsal setae. In both, Shanthiniae and Dialeuronomada, medium length of abdominal segment VII is shorter than that of VIII. Presence of a row of 12 marginal setae and a peripheral row of characteristic papillae on the submarginal area readily separate the latter from the former. The new genus differs from all the above three genera in the characteristic sclerotic pattern on the dorsum, in the transverse moulting suture typically reaching margin, in the location of vasiform orifice not greater than its length from posterior body margin, and in the absence of stipples in the tracheal folds as well. Etymology: This genus is named after the author’s wife Mrs. Shanthini David. Shanthiniae sheryli gen. et sp. nov. (Fig. 2) Pupal case: Typically polyhedral, with six corners on either side. 0.68 mm long and 0.56 mm wide, widest across abdominal segment I. Black in colour with no wax secretion. Margin: Very finely crenulate and crenate-looking, with about 4 crenulations in 0.1 mm length of margin; crenulations each much wider than long, their apices subconical to rounded or rather straight. Margin at pore area cleft. Thoracic tracheal pores well defined, inset with a single tooth-like projection. Caudal tracheal pore area not inset. Anterior marginal setae 9.3 pm long, posterior marginal setae 12.4 pm long. Dorsum: Ornamented with spots, reticula- tions and minute tubercles. Submargin narrowly marked by a weak furrow. Transverse ridges and furrows running mesad from margin and submargin to anastamose in the dorsal disc area, giving a leopard skin-like appearance to the dorsum. Marginal furrows each alternated with 2-4 submarginal ones. Subdorsum granulated. Submedian area on cephalothorax and median area on abdominal segments densely spotted. Longitudinal and transverse moulting sutures reaching margin, the ends of the latter opposite meso-metathoracic suture. Base of cephalic, first and eighth abdominal setae distinct, setae very minute; bases of eighth abdominal setae located laterad of top of vasiform orifice. Caudal setae not discernible. Segmenta-tion distinct in submedian area; sutures each with anterior and posterior branches, their ends anastomosing with subdorsal reticulation. Median length of abdo- minal segments subequal, median length gradually decreasing from abdominal segment I- VII; that of VII as long as that of VI and shorter than VIII. Disc pores and porettes present on dorsai disc; porettes dark-rimmed and characteristically sclerotized laterad. Vasiform orifice cordate, located about its length from posterior body margin, its sides prominent; its inner margin with teeth, especially posteriorly; 37.2 pm long and 34.1 pm wide. Operculum cordate, nearly filling the orifice; 24.8 pm long and 27.9 pm wide. Lingula concealed. Caudal furrow well defined. Caudal ridges distinct. Venter: Thoracic and caudal tracheal folds well defined. Ventral abdominal setae 9.3 pm long. All four pairs of spiracles evident, anterior thoracic spiracles larger than others. Setae or spines on legs not discernible. Antennae reaching the base of prothoracic legs. Adhesive sacs not discernible. Rostrum distinctly segmented, setae at base absent. Host: An unidentified plant. Holotype: A pupal case mounted on slide, on an unidentified plant, India: Tamil Nadu: Karaiyar Dam (Papanasam), 14.iv.1993. Coll: P.M.M. David (No. 180.A.). Etymology: This species is named after the author’s son D. Sheryl who often accompanied him during the survey. 126 JOURNAL , BOMBAY NATURAL HISTORY SOCIETY. 97(1) APR. 2000 NEW DESCRIPTIONS Fig. 2: Shanthiniae sheiyli gen. et sp. nov. : A. pupal case; B. thoracic tracheal comb; C. section of margin and submargin; D. disc pore and porette; E. vasiform orifice; F. dorsal markings. Genus Vasantharajiella gen. nov. Type-species: Vasantharajiella kalakadensis sp. nov. Pupal case oval, jet black in colour; margin lobulate; tracheal combs distinct; submargin separated from the dorsal disc by a distinct furrow not interrupted even at caudal region; marginal setae absent; submarginal setae present; first abdominal setae absent; transverse JOURNAL, BOMBAY NATURAL HISTORY SOCIETY. 97(1), APR. 2000 127 NEW DESCRIPTIONS moulting suture reaching submarginal furrow; median area of abdominal segments tuberculate; rachis on cephalothorax and abdomen present; vasiform orifice much smaller compared with its body size, subcordate, wider than long; operculum of similar shape, filling half the orifice, concealing lingula. Diagnosis: The genus differs from such genera of Aleurolodini as Aleuropapillatus Regu & David, Aleurolobus Quaintance & Baker, Africaleurodes Dozier, and Asterochiton Masked in the presence of complete submargin all around the case, without interruption even in the caudal region. Though it resembles the rest of the genera in the presence of the complete submargin, it is clearly distinguished from them by several characters. It is distinct from Crescentaleyrodes David & Jesudasan in the presence of thoracic and caudal tracheal combs and in the absence of a row of crescent-shaped pores in the submargin. Absence of first abdominal setae and a pouch-like structure at the thoracic and caudal tracheal comb area separate Vasantharajiella from Rositaleyrodes Meganathan & David. Oriental eyr odes David possesses very long hair-like submarginal setae, comparatively shorter median length of abdominal segment VII, much larger (80 x 75 pm) elevated vasiform orifice located only about its length from the posterior body margin. On the other hand, Vasantharajiella has minute submarginal setae, well defined rachis on cephalothorax and abdomen, equally long abdominal segment VII and, compared to the body size, much smaller (24.8 x 34.1 pm), non- elevated, vasiform orifice located about 10 times its length from posterior body margin. The genus also differs from all these genera in lacking anterior and posterior marginal setae. Etymology: Named in honour of Dr. B. Vasantharaj David, Director, Jai Research Foundation, Valvada, Gujarat, as a mark of respect. Vasantharajiella kalakadensis gen. et sp. nov. (Fig. 3) Pupal case: Jet black in colour, surrounded by a thick fringe of white waxy filaments; powdery wax deposits on dorsal sutures and submarginal lines. Living on the upper surface of leaves. 1.82-1.86 mm long and 1.46-1.54 mm wide; widest across abdominal segments II-IV. Margin: Lobulate, 10-11 lobulations mO.l mm width of margin; teeth each as long as wide. Margin at tracheal pore area slightly indented. Tracheal combs distinct; about 6 teeth at pore area larger than other teeth, with incisions in between teeth deeper than those in between other teeth. Anterior and posterior marginal setae not discernible in available specimens. Dorsum: Submargin characteristic; complete without any interruption even at caudal region; separated from dorsal disc by a well defined furrow gradually widening caudad; approximately ‘A the width of dorsal disc across the greatest width of body. Submarginal ridges and furrows distinct. A row of faint papillae-like markings evident. Submarginal setae in 1 3 pairs arranged in 2 rows: 4 pairs in outer submargin (2 pairs at cephalic end, one just posterior to thoracic tracheal furrow, one opposite abdominal segment I) and 9 pairs in inner submargin (3 pairs anteriad of and 6 pairs posteriad of thoracic tracheal furrow) 12.4-24.8 pm long, their bases porous, setae tapered, apices acute. Cephalic setae 15.5 pm long. First abdominal setae absent. Eighth abdominal setae 9.3-15.5 pm long, located laterad of top of vasiform orifice on a conspicuous ridge, their apices pointing towards orifice. Caudal setae 18.6 pm long, located on outer submargin anterior to the lateral tooth of tracheal comb. Longitudinal transverse moulting suture reaching margin. Transverse moulting suture curved caudad from its midpoint, recurved cephalad, terminating at submarginal furrow opposite meso-metathoracic suture. Segmenta- tion well defined in submedian area. Median length of abdominal segments I-VI equal and of 128 JOURNAL, BOMBAY NATURAL HISTORY SOCIETY. 97(1) APR. 2000 NE W DESCRIPTIONS Fig. 3: Vasantharajiella kalakkadensis gen. et sp. nov.: A. pupal case; B. thoracic tracheal comb; C. margin; D. disc pore and porette; E. vasiform orifice; F. submarginal seta VII shorter than that of other segments. Median area of abdominal segments I- VIII finely tubereulate. Rachis on cephalothorax and abdomen characteristic; promesothoracic suture bifurcated, each curved cephalad and caudad, recurving mesad with chain-like designs all along in subdorsal area; meso-metathoracic suture bifurcated. A somewhat transverse rachis laterad of meso-metathoracic suture; abdominal rachis III- VII each bifurcated, finger-like in inner subdorsum with chain-like designs along sutures; minute striations inside abdominal rachis. Disc JOURNAL. BOMBAY NATURAL HISTORY SOCIETY. 97(1). APR. 2000 129 NE W DESCRIPTIONS pores and porettes in a row on central submargin, in groups of 2-3 in a row each on outer subdorsum and on submedian area. Submedian depressions distinct on thoracic and abdominal segments. Pockets on abdominal segment VII inconspicuous. Vasiform orifice very small, subcordate, located about twice its length from posterior suture and 1 0 times its length from body margin; wider than long, 24.8 pm long and 34. 1 pm wide; its top straight, less sclerotized; sides prominent. Operculum of similar shape; longer than wide, 12.4 pm long and 27.9 pm wide, filling about half the orifice. Lingula concealed. Caudal furrow indicated by shallow depressions. Caudal ridges distinct anterior to submarginal furrow and laterad of vasiform orifice. Venter: Tassellated. Thoracic tracheal fold marked up to submarginal line, then indicated by minute stipples; caudal fold distinct; stipples up to central submargin. All spiracles evident. Ventral abdominal setae 46.5 pm long, 31.0 pm apart. A minute seta at base of mesothoracic legs, their tips with a finger-like projection. Host: An unidentified woody climber. Holotype: Pupal case mounted on slide, on an undetermined climber, rNDiA; Tamil Nadu: Kalakad forest, 5. i. 1993. Coll: P.M.M. David (No. 208.4). Paratypes: 1 1 specimens on slides bearing the same collection data as of holotype. Etymology: Derived from the collection site, Kalakad forest. Types Depository: The holotypes are deposited with the Division of Entomology, Indian Agricultural Research Institute, Pusa Campus, New Delhi. The paratypes of M. rubicie and V. kalakadensis are available with the Centre for Advanced Studies in Agricultural Entomology, TNAU, Coimbatore; with Dr. B.V. David, Director, Jai Research Foundation, Vapi; and with the Department of Entomology, Natural History Museum, London. Acknowledgements The author is grateful to Dr. M. Mohanasundaram, Professor of Agricultural Entomology (Retd.), Tamil Nadu Agricultural University, Coimbatore, for suggesting the problem and for guidance throughout the investigations, and to Dr. B. Vasantharaj David, Director, Jai Research Foundation, Valvada, Gujarat, for confirming the identity of the species, valuable suggestions and providing reprints. 130 JOURNAL BOMBAY NATURAL HISTORY SOCIETY 97(1) APR. 2000 L YSIONOTUS PALINENSIS — A NEW SPECIES OF GESNERIACEAE FROM ARUNACHAL PRADESH, INDIA1 G.D. Pal2 ( With one text-figure) Key words: Lysionotus palinensis, new species, Arunachal Pradesh. During plant exploration in the district of Lower Subansiri, Arunachal Pradesh, an interesting species of the genus Lysionotus D. Don was collected. A critical study, based on the regional herbarium specimens of allied species and literature on species of Lysionotus D. Don, has proved it to be quite distinct from all known species and is described. Line drawings are provided. Lysionotus palinensis G.D. Pal sp. nov. (Figs. : A-D) Lysionotus serrato D. Don affinis, sed differt foliis lanceolatis ad lineari-lanceolatis, ad marginum spinulosis crenato-serratis coriaceis, pedunculis 3-7 cm longis, calieis lobis lanceolatis, 0. 9-1.0 x 0. 1-0.2 cm, cuspidato- acuminatis, 4-6 nervatis. Typus: Holotypus lectus a G.D. Pal ad locum Arunachal, Inferior Subansiri district, Palin c. 1400 m, dia 9.ix.l983, subnumero 400, ex positus in CAL. Shrubby herbs, 30-40 cm tall, usually unbranched, rooting at the lower nodes. Stems terete or faintly ridged, pubescent above. Leaves ternate, sometimes basal leaves opposite, lanceolate to narrowly lanceolate, 3-7 x 0. 5-2.0 cm, cuneate or rounded at base, acuminate, spinulous crenate-serrate at margin, glabrous, coriaceous; young leaves hairy on nerves underneath, pale gren underneath; lateral nerves 4-6 pairs; petioles 0.2-0. 3 cm long, hairy. Inflorescence laxly cymose many flowered; peduncles 3-7 cm long, terete, wiry, glabrous or sparsely pubescent; bracts ovate-lanceolate, 0.5- 0.6 x 0.3-0.35 cm, acuminate, 3-nerved; pedicels 0.5-1 .0 cm long, wiry, glabrous. Flowers bluish- purple; calyx lobes lanceolate, 0.9- 1.0 x 0.1 -0.2 'Accepted December, 1998. :Botanica1 Survey of India, Arunachal Field Station ltanagar791 111, Arunachal Pradesh, India. cm, caudate-acuminate, 4-6 nerved, purple; corolla tubular, broad at middle, 4. 0-4. 5 x 0.8- 1.0 cm, distinctly nerved within, reticulate at throat; upper lip 0.8 cm longer than lower lip. 3-lobed; middle lobes oblong to sub orbicular, about 0.8 x 0.6 cm, obtuse; lower lip shorter, truncate; stamens 2, fertile; filaments 0.8-1 .0 cm long, flattened; anthers connivent; ovary oblong, 0.2 cm long; styles slender, 2. 0-2. 3 cm long, unevenly thickened. Capsules not seen. FI.: August-September. Remarks: Grows in moist shaded places of subtropical primary forests on humus rich soil associated with Impatiens, Begonia spp. Distribution: India: Arunachal Pradesh, Lower Subansiri dist . , Palin c. 1400 m, 9.ix.l983; G.D. Pal 400 (Holotype- CAL) Note: The new species is closely allied to L. servants D. Don, but can be differentiated by: leaves lanceolate to linear-lanceolate; spinulous crenate-serrate at margin, coriaceous; peduncles 3-7 cm long; calyx-lobes lanceolate, 0.9- 1.0 x 0.1 -0.2 cm, cuspidate-acuminate, 4-6 nerved. Acknowledgements I am grateful to the Director, Botanical Survey of India, Calcutta for facilities. I also thank Dr N.C. Majumder, ex Scientist SE, Botanical Survey of India, Calcutta for the la tin diagnosis of the taxon and Dr G.S. Giri, Scientist SE, Central National Herbarium, Howrah for sketches. JOURNAL , BOMBAY NATURAL HISTORY SOCIETY, 97(1), APR. 2000 131 NEW DESCRIPTIONS Figl. A-D: Lysionotus palinensis sp. nov., A. Habit; B. Corolla split open; C. Anther; D.Gynaecium with calyx. 132 JOURNAL. BOMBAY NATURAL HISTORY SOCIETY. 97(1) APR. 2000 REVIEWS 1. BIRDS OF NEPAL: FIELD ECOLOGY, NATURAL HISTORY AND CONSERVATION, by Tej Kumar Shrestha. Published by Mrs. Bimal Shrestha, Kathmandu, Nepal. 2000. Pp 287, (21 x 14 cm). Price not mentioned. Prof. Tej Kumar Shrestha of Nepal is a prolific writer on natural history. As the jacket of his latest book birds of nepal claims, he is the author of the book the spiny babbler: an endemic bird of nepal, and other “eleven outstanding books on wildlife and natural resources”. However, his books (I have read three) lack scientific accuracy and the high standard expected from a professor with a D.Sc. degree. While the quality of printing of the present book is much better than his earlier books, the language is pedantic and needs proper editing. There are too many editorial mistakes to be enumerated in this brief review. The book claims to be a photographic field guide. There is a chapter on bird photography, (Bird-watching and Field Craft, pp. 30-38) which shows Prof. Shrestha with his various cameras, but many of the pictures are taken in zoos (e.g. plates 10-13, 17-22, 46-47, 52-53, 69), some pictures are of trapped birds (e.g. coot, Baillon’s crake and Indian moorhen, plate 89, painted snipe, plate 93), and some even of mounted specimens (plates 31-32). The flight pictures on plates 33, 34 and 35 are quite interesting, and some close-ups are sharp. I particularly liked the picture of the jungle myna (plate 36). Another interesting picture is that of a male magpie robin (not foster mother as claimed in the caption) feeding a juvenile cuckoo (plate 70). More than 250 species have been depicted in 144 colour plates and many in black & white, but identification pointers are not given in the captions. Moreover, arrangement of bird pictures is arbitrary, so to find a particular species is not easy. The most ill-conceived aspect of this book is the checklist. The author enigmatically starts his checklist with the spiny babbler Turdoides nipcilensis, a species on which he has done his Ph.D. Since the publication of Sibley & Monroe’s new classification based on DNA finger printing, there is already confusion in the classification of birds. Prof. Shrestha’s book will further add to this confusion. Strangely, he has not given any valid reason for adopting his own classification. The book covers a wide spectrum of subjects from ‘Ornithography of Nepal’ to ‘Birds in Nepalese Literature’. Despite its many drawbacks, it is an interesting book for ornithologists and conservationists of the Indian subcontinent because birds face the same conservation problems all over South Asia. The type of trapping methods described by Prof. Shrestha in his book are also used in India. Similarly, the reasons for trapping birds (and other wildlife) and the modus operandi of trappers/traders in Nepal are the same as in India. Nepal is becoming the main conduit for smuggling of birds to European and Middle East markets. If this book creates awareness, this special third millennium edition would serve its stated puipose “to conserve environment and the health of people in the approaching millennium”. ■ ASAD R. RAHMANI 2. BIOGEOGRAPHY OF THE REPTILES OF SOUTH ASIA by Indraneil Das, Kreiger Publishing Co., Malabar, Florida, 1996, pp. 87 + xxxvi colour plates, (24.5 x 16.6 cm). Price not mentioned. As the title suggests, this book is a The practice of displaying colour plates at compilation of different aspects of geographical the very beginning of the book, though not distribution of reptiles of the South Asian region, common, catches the reader’s attention. All the JOURNAL, BOMBAY NATURAL HISTORY SOCIETY, 97(1), APR. 2000 133 REVIEWS plates are of exemplary quality. The Montane trinket snake ( Elaphe Helena montico Haris), on the cover is excellent. The author, at the outset, acknowledges the treatises of Smith (1931; 1935; 1943) (P. 1) to be ultimate sources of identifying the Subcontinent’s reptilian fauna. He also admits drawing on the biogeographic analysis of individual countries by various workers as sources for data regarding distribution, yet does not ‘necessarily’ agree with them on the ‘interpretation’ of their findings. The author has devoted an entire chapter to apprise the reader of the methods used in his analysis. He also points out that the status of nomenclature and species reallocation is in great turmoil and uncertainty. Thus, it is safe to assume that a more exhaustive study comprising of valid names, distribution and species reallocation is to follow. The descriptions of the physiographic zones within the South Asian region, first from the physical and biological point of view and then solely on the basis of faunal characteristics like biodiversity and endemicity, are very informative and give a detailed view of the South Asian region. The results obtained and subsequent discussions on (a) Biodiversity and Endemicity (b) Faunal characteristics of physiographic zones (c) Patterns and correlates of diversity (d) Affinities between physiographic zones (e) Affinities with extralimital fauna (f) Barriers and speciation (g) Disjunct distribution of taxa, are written in a lucid style. A map is used in Chapter 3 to depict the physiographic zones. A tree-diagram effectively explains the affinities between physiographic zones. Graphs have been used to interpret or express certain data. The presence of a map and graphs depicting the physiographic zones makes it easier to visualise the explanation given. These aids make the initial chapters very informative and interesting. The analysis is complete, but a $ the author himself points out, the checklist can be the basis of a more exhaustive study owing to the instability of the taxonomic arrangements and interpretation. ■ MEGHANA GAVAND 3 . MOSSES OF KHANDALA AND MAHAB ALESHWAR IN THE WESTERN GHATS (INDIA) by G.T. Debhade, Published by A.S. Dalvi, Thane, 1998. Pp. iv + 193, (25 x 18.5 cm). Hardbound price Rs. 800/-, $ (US) 40. The mosses and in fact the entire group of Bryophytes are neglected, though they have an important bearing on the evolutionary history of the plant kingdom. The very fact that the bryophytes represent the transitional zone between the amphibian and the terrestrial habit, and that they may, perhaps, be the base in the origin and the organisation of the reproductive machinery of other plant groups from pteriodophytes to angiosperms, explains the importance of the Bryophyta in comparative morphology, ecology and phylogeny. Inspite of the academic importance and even economic value, the group remains neglected and it is in this context that this publication becomes significant. The work, though confined to a certain region of the Western Ghats, is of value in the identification of many mosses in the entire Western Ghats. The author has painstakingly made field collections and described them, giving information on characters of diagnostic value, with particular reference to the capsule and even the spores at times. Altogether, 87 species under 48 genera and 27 families have been covered. The general introduction, the review of previous work and the scope of the present work, together with the table containing salient 134 JOURNAL. BOMBAY NATURAL HISTORY SOCIETY. 97(1) APR. 2000 REVIEWS information on the concerned taxa, give valuable information on phytogeography and ecology of the mosses in South India. The author has given information on the physiographic, edaphic and environmental conditions of the areas covered by the present study, followed by an account of methodology of collection, preservation and identification, including preparation of permanent slides of the whole plants and their parts, including spores. The morphological analysis, both mega and micro- characters, provides a picture of the thoroughness and depth of research that the author has carried out. The descriptions are followed by a general analysis of various aspects of the study which, perhaps, is the best part of the publication. The information on habitat shows the location- specific occurrence of mosses on calcareous, lateritic, or peaty soils, and such information is of immense environmental importance for using mosses as bio-indicators. Further, information on moss sociology is in fact a reflection of biodiversity combinations in various micro- ecosystems. The account on geographical distribution gives very useful data on the abundance or rarity of various taxa. The finding of a large number of endemic species in the restricted area of Khandala and Mahabaleshwar should receive conservative attention to help save these species from extinction. The concluding sentence “As we go south, more and more humid species begin to appear and as. one goes northwards, drier species are met with” holds good for the