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JOURNAL

of the

Bombay Natural History

Society

Vol. 81, No. 1

Editors : J. C. Daniel, P. V. Bole & A. N. D. Nanavati

APRIL 1984

Rs. 45

m

T’-

NOTICE TO CONTRIBUTORS

Contributors of scientific articles are requested to assist the editors by observ- ing the following instructions:

1. Papers which have at the same time been offered for publication to other

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♦ i

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8. Reference to literature in the text should be made by quoting the author’s name and year of publication, thus: (Banerji 1958).

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Hombill House, Editors,

Shaheed Bhagat Singh Road, Journal of the Bombay

Bombay 400 023. Natural History Society .

VOLUME 81(1) : APRIL 1984

Date of Publication : 9-8-1984

CONTENTS

Page

Some aspects of the Biology and Ecology of Narcondam Hornbjll ( Rhyticeros narcondami). By S. A. Hussain. ( With one coloured & five Black-and-W hite plates and five text- figures) • • 1

Seasonal variation in the population of Acrida exaltata Walk, at Aligarh.

By Shamshad Ali. ( With seven text-figures ) .. 19

Larval culture of the Hermit Crab Clibanarius aequabilis var. merguiensis De Man (Decapoda, Anomura, Diogenidae) reared in the Laboratory. By Venkatray N. Nayak. ( With seven text-figures ) . . 29

Tourist activity and behaviour of the Leopard Panthera pardus fusca (Meyer,

1794) in the Ruhuna National Park, Sri Lanka. By M. R. Chambers, Charles Santiapillai and N. Ishvvaran. {With two text-figures ) . . 42

Some observations of scarce birds in Nepal. By N. J. Redman, F. Lambert and

R. Grimmett . . 49

Spawning of some important cqldwater fish of tfie Garhwal Himalaya. By

S. P. Badola and H. R. Singh. {With a plate) . . 54

Reproduction biology of the Soft-furred Field Rat, Rattus meltada pallidior (Ryley, 1914) in the Rajasthan desert. By B. D. Rana and Ishwar Prakash.

{With three text-figures) . . 59

Flowering plants around the holy shrine of Kedarnath, Uttar Pradesh. By

J. K. Semwal. {With a text-figure) . . 7!

Observations of the reproductive biology of the Indian Chameleon, Chamaeleo

zeylanicus (Laurenti). By L. A. K. Singh, L. N. Acharjyo and H. R. Bustard 86

New records and hosts of Aphid parasitoides (Hymenoptera: Aphidiida) from

Kashmir, India. By R. C. Bhagat . . 93

Host plants of the Fruit Flies (Diptera: Tephritidae) of the Indian sub-

continent, exclusive of the sub-family Dacinae. By Mohammad Zaka-ur-Rab 99

Observations cn the length-weight relationship of the fish Rasbora daniconius

(Ham.-Buch.). By V, Y. Thakre, and S. S. Bapat. {With two text-figures) .. 105

Influence of atmospheric temperature and humidity on the variations in seasonal abundance and phenology of Micrcnecta striata Fieber. By T. C. Banerjee, A. S. Mondal and T. K. Nayek. {With seven text-figures) .. 110

Further contribution to the Flora of Buxa Forest Division, Jalpaiguri District

(West Bengal). By J. K. Sikdar and Rolla S. Rao. {With a text-figure) .. 123

New Descriptions:

A new species of Soiichus L. (Asteraceae) from south India. By M. Chandra-

bose, V. Chandrasekaran and N. C. Nair. ( With seven text-figures ) .. 149

Description of two new species and one new record of Cryptostigmatid Mites (Acari: Oribatei) from Maharashtra, India* By A. K. Sanyal. (With four text-figures) . . 151

A new species of Ophiorrhiza (Rubiaceae) from Andhra Pradesh. G. V. Subba

Rao and G. R. Kumari. ( With six text-figures ) 156

Description of a new genus and some new species of tcrrenticole Diptera

of the northwest Himalaya. By B. K. Kaul. ( With thirty-four text-figures) 158

Three new species of genus Isoetes L. from Rajasthan, India. By C. B. Gena

and T. N. Bhardwaja. ( With a pi ate) . . 165

A new 'Sim pi ocos Jacq. ( Symplocaceae) from southern India. By A. N. Henry, R.

Gopalan and M. S. Swaminathan. ( With seven text-figures) . . 169

Reviews: :

1. Sunlight and Shadows. (Divyabhanusinh Chavda) .. 172

2. The IUCN Aphibia-Reptilia Data Book — Part I. (P. Kannan) , . 174

3. Symbiosis in the Mango-hopper: A study in Comparative Cytopathology.

(T. S. S. Dikshith and R. K. Varshney) .. 175

4. Supplement to Duthie’s Flora of the Upper Gangetic Plain and of the adjacent

Siwalik and Sub-Himalayan Tracts. (A. R. Daruwalla) .. 176

Miscellaneous Notes :

Mammals: 1. Record of the Fulvous Fruit Bat, Rousettus leschenaulti (Desmarest, 1820) from Sikkim, with notes on its interesting feeding habit and status. By R. K. Ghose and D. K. Ghosal (p. 178); 2. A feeding association between a Heteropteran Bug and Langurs. By Paul N. Newton (p. 180); 3. White dots on the legs of Barking Deer. By J. Mangalraj Johnson (p. 182); 4. A note on cannibalism in desert rodents. By A. P. Jain (p. 182);

5. Bandicoot rat seizing a snake. By N. C. Nandi (p. 183); 6. Record of a foetus of the Unless Black Porpoise from Goa Coast. (With six text-figures). By M. Hafeezullah (p. 183).

Birds: 7. Cattle Egrets (Bubulcus ibis) feeding on cicadas on trees. By S. G. Monga and Pan ish Pandya (p. 186) ; 8. Unusual plumage in a Cattle Egret Bubulcus ibis coro- mandus (Boddaert). By Natin Jamdar (p. 187); 9. Some notes on the Indian Reef Heron. By K. S. Dharmakumarsinhji (p. 188); 10. Exceptionally large eggs of the common House Crow, Corvus s. splendens Viell. By S. G. Monga (p. 189); 11. Laboratory observations on the incubation period of the Indian Black Ibis Pseudibis papillosa (Temminck). ( With a text- figure). By C. Salimkumar and V. C. Soni (p. 189); 12. Seasonality and occurrence of birds in the Eastern Ghats. By Humayun Abdulali (p. 191); 13. A curious experi- ence with a small Minivet ( Pericrocotus cinnamomeus) . By Sadiq A. Futehally (p. 191); 14. Frequency and duration of incubation of the eggs for Aegithina tiphia. By H. Daniel Wesley (p. 193); 15. Hypsipetes madagascariensis sinensis (La Touche): a first record for India. By S. Dillon Ripley and S. A. Hussain (p. 195); 16. The diagnostic plumage characters of the Redheaded Babblers Stachyris ruficeps and S. rufifrons. By C. J. O. Harrison (p. 197); 17. Behavioural response of a male Magpie-Robin ( Copsychus saularis Sclater) to its own song. By E. Narayanan (p. 199); 18. Mouse, a nest-parasite of Baya Weaver bird ( Ploceus philippinus L.). ( With three text-figures) . By D. Regupathy and T. A. Davis

(p. 200).

Fishes: 19. First record of the freshwater Grey Mullet. Rhinomugil corsula (Hamilton) from Maharashtra. By M. S. Pradhan and D. F. Singh (p. 202); 20. Preliminary observa- tions on the migratory behaviour of the Garhwal Himalayan Mahseer. ( With a text-figure) . By Prakash Nautiyal and M. S. Lai (p. 204).

Insects: 21. The spider as bee enemy. ( With a text-figure) . By A. K. Thakur and O. P. Sharma (p. 208); 22. New records of Aphids (Homoptera: Aphididae) from Uttar Pradesh. By Basant K. Agarwala. D. Ghosh and D. N. Raychaudhuri (p. 211); 23. New record of insect pests infesting Kastnri Bhendi. Hibiscus abehnoschus Linnaeus, a medicinal plant. By R. Rajashekhargcuda, M. C. Devaiah and Suhas Yelshetty (p. 212); 24. A new record of Neopheosia fasciata (Moore) on apple. By Ramesh Chander (p. 213).

Other Invertebrates: 25. On a glossiphonid leech. ( With three text-figures) . By H. V. Ghate (p. 214); 26. Studies on the biological control of two common vector snails of

Punjab by predatory insects. By H. S. Bali. Sawai Singh and Sunita Sharma (p. 216);

27. Necrophagous habit in the Giant African Snail, Aehatina fulica fulica Bowdick. (With a plate). By A. K. Das and R. M. Sharma (p. 219).

Botany: 28. A contribution to the vegetation of Chaibasa (North), Singhbhum Dist. (North Bihar). By Dilip Kumar Biswas (p. 221); 29. Occurrence of Desmodium seorpiurus (Swartz) Desvaux in Western India. (With five text-figures) . By V. D. Vartak and M. S. Kumbhojkar (p. 224); 30. Identification and distributional note of a few species of Epilo- bium Linn, in India. By G. S. Giri and R. N. Banerjee (p. 227); 31. Cucumis melo Linn, in Punjab — a taxonomic reappraisal. By M. Sharma (p. 229); 32. On the identity of two species of Oldenlandia L. (Rubiaceae). By D. B. Deb and Ratna Datta (p. 232); 33. Ceropegia pusilla Wight et Arn. (Asclepiadaceae) in Hoshiarpur District (Punjab).

By Anil K. Goel and Surendra Singh (p. 233); 34. Two noteworthy plants from West

Bengal. By S. N. Das and S. C. Roy (p. 234); 35. Abnormal flowering of Agave angusti- folia Haw. ( With a plate). By Anand Kumar and P. G. Diwakar (p. 235); 36. Gregarious flowering of Carvia callosa Bremek and Nilgirianthus reticulatus Bremek at Amboli. By Ulhas Rane (p. 236); 37. Pteris scabripes Wall, ex Hook. — a new find from India. By S. R. Ghosh (p. 237); 38. Studies in Leguminosae XXX - — Further contributions to Dalbergia L. f. and Denis Lour. {With three text-figures) . By K. Thothathri (p. 238).

\

J. Bombay nat. Hist, Soc 81

Plate

Narcondam Hornbills: Female above; Male below

JOURNAL

OF THE

BOMBAY NATURAL HISTORY

SOCIETY

1984 APRIL

Vol. 81

No. 1

SOME ASPECTS OF THE BIOLOGY AND ECOLOGY OF NARCONDAM HORNBILL ( RHYTICEROS N A RCONDA Ml )

S. A. Hussain1 2

( With one coloured & five Black-and -White plates and five text -figures)

Introduction

Two species of hornbills of the genus Rhyti- ceros are represented in the Indian sub-conti- nent- Of these R. undulatus occurs in north- eastern India, Burma down to Malay penin- sula and the Mergui Archipelago. The other, R. narcondami is restricted to Narcondam, an off lying island in the Andaman group. Very little is known about the biology and ecology of the latter. Hume (1873) on an expedition to the Andaman group collected several horn- bills from Narcondam and named it narcon- dami. Prain (1893), St. John (1898), Cory (1902) and Osmaston (1905) visited Narcon- dam subsequently to collect specimens. The last spent five days, (the longest period of time spent in the island by a visitor), in search of stands of the timber tree Pterocarpus dal-

1 Accepted January 1984.

2 Project Scientist, Avifauna Project, Bombay Natural History Society.

bergoides. He also made some notes on the fauna and flora including Hornbills, whose number he estimated to be about 200. No further information on the hornbills was avail- able until two of my colleagues at the BNHS* Robert B. Grubh and R. J. Pimento visited the island briefly in 1969. Abdulali (1971) visited the island in the following year and spent a few hours to collect specimens. In 1972 along with Mr. N. J. George of Prince of Wales Museum, I visited the island at the instance and direction of Mr. Humayun Ab- dulali. We visited South and North Andamans and Narcondam island from 4th March to 25th April 1972 and the field data and speci- mens collected by us were reported in the Journal (Abdulali 1974). The Narcondam island (the name Narcondam is derived from Sanskrit Naraka — Hell; Kundam — Pit, — an obvious reference to the origin of the island which is believed to have been an active vol- cano not long ago) is difficult to approach, except during the months of March, April

JOURNAL . BOMBAY NATURAL HIST. SOCIETY, Vol. 81

and May when the sea around is comparati- vely calm.

The present paper records observations made by me during my stay on Narcondam from 16th March to 14th April 1972, and the sub- sequent observations on the two hornbill chicks brought back which lived in captivity at the Society’s premises.

T axonomical notes :

Hume (1873) while describing narcondami stated that it resembled R. plicatus of Borneo and due to the difference in size as well as the absence of a zoogeographic connecting link between these species gave the former the sta- tus of a species. Baker (1927) treated it as a full species in the absence of intermediates and stated that systematists may consider it to be a small island race of R. plicatus of which R. everetti of the Moluccas was thought to be an intermediate form. Blyth (1845) had in the meantime, described subruficolUs from N Burma, which he differentiated from R. plica- tus ruficollis by the absence of any ridges on the sides of bill and by its smaller size. Peters (1945) accepted this nomenclature and consi- dered subruficolUs a valid race of plicatus. Sanft (1960) who has authoritatively reviewed the family Bucerotidae, did not accept subru- ficollis as a race of plicatus preferring to synonymise it with undulatus. His argument was that undulatus and subruficolUs are from

3 Sanft, (IBIS 95: 702-703) after studying 16 museum specimens of R. undulatus, R. subruficolUs and intermediates argued that the ranges of the two overlap with the intermediates showing characteristics of the both, and therefore subruficolUs is synony- mous with undulatus. However, Elbe! ( Condor 71 (4) : 434-435) on the evidence of the mellophaga present in the above two species concluded that subruficolUs is distinct from undulatus and is closer to plicatus.

the same ancestral stock, differing only in deve- lopmental stages as well as localised varia- tions. One of the main differences is in the structure of the bill i.e. presence of ridges on the side of the basal half of the bill (= undu- latus) and absence of it (= subruficolUs) , which according to him, are linked with sexual matu- rity and tend to develop as the bird becomes older. The difference in body size, accord- ing to him, was ecologically linked to the types of habitats in which they occur. Thus the larger birds of the mountainous region are undulatus and the smaller occurring in low hill zones subruficolUs ■ However, he had overlook- ed two other distinct characteristics that differentiate the two. The colour pattern of the head and neck of males, colour of gular pouch, and presence or absence of a black band on throat. These patterns are ap- parently not linked with ecological distribu- tion. Are they then linked with age? Does the yellow colour of gular pouch in S and black band on the pouch in both female and male develop as they grow older?3 (Table 1).

A 16 year old specimen of R. p. subruficolUs at the San Diego zoo shows all the characteri- stics of the typical ruficollis with blue gular pouch without the black band (K. C. Lint, pers. comm.). Under these circumstances the taxonomic and zoogeographic position of nar- condami is quite intriguing. If one were to accept Sanft’s proposition, narcondami is a smaller form showing immature characters of undulatus isolated in the islands long ago and gradually evolving into the present form (ende- mic?) and in the process losing the adult characteristics of the undulatus. On the other hand, the plicatus link theory, with the recogni- tion of subruficolUs as a distinct subspecies of the former, would perhaps open up a new line of possibilities on the zoogeography of the re- gion. Another species which perhaps raises

4-*

ECOLOGY OF NARCONDAM HORNBILL

3

Fig. 1. Distribution of Hornbilis in SE Asia (After Sanft 1960).

JOURNAL , BOMBAY NATURAL HIST. SOCIETY, Vol. 81

Table 1

R. undulatus R. p. subniftcollis R. narcondami

Bill

Wing

Weight Head & Neck

Gular

pouch

Distri-

bution

Side of mandibles ridged at base

458-505 $ 9 2.500 gm

$ dark brown crown and hindneck-almost black lower down. Throat & upper neck whitish

Bright yellow with black band 8

Dark blue and black band 9

NE India, Burma. Singapore, Sumatra, Java & Borneo

side of mandibles smooth at base 420-445 $ 9

1.900 gm

8 rufous head & hind neck, white on throat

Pale blue $ 9

Without black band

S. Burma. SW Thailand Sumatra, Borneo

side of mandibles smooth at base 303-305 $

285-287 9 600-750 gm

$ rufous head and neck

Pale blue $ 9 without black band

Narcondam I.

similar questions is R. everetti an endemic of Sumba islands, SE Asia (Fig. 1). There are similarities in the evolution of these two species. Both are endemic to small

islands, are smaller versions of neigh-

bouring forms, and have distinct morpho- logical characters (Fig. 2). Ali and Ripley

(1970), followed Peters’ nomenclature and call- ed it R. ( undulatus ) narcondami. Flowever, Ripley (1982) after seeing the live specimens in the BMHS and personal discussions with me agreed that narcondami is closer to plicatus than undulatus. Kemp and Kemp (1975) mention the long-hop flights of the SE Asian hornbills which sometimes cross the sea to offshore

islands. These hornbills have been observed to take off from the mainland and fly in “follow the leader” formation for some distance straight out over the sea and return eventually to the starting point. Is this behaviour then an instinctive urge of a long forgotten “migra- tory” habit? The significance of the white tail

in these hornbills which can be seen from long distances and which may probably act as a visual stimulus for the following hornbills, is worth noting.

Physiography and vegetation :

Narcondam island (13°30' N; 94°38' E) is situated c 500 km NW off Mergui archipelago and c 300 km SW of the Gulf of Martaban off the Burmese mainland, and c 125 km east of North Andaman in the Andaman and Nicobar group of islands in the Bay of Bengal. The island has a total area of about 682 hectares and is a part of a submerged chain of mountains in the Andaman archipelago. Narcondam is one of the two off-lying volcanic islands in the eas- tern sector of the group. It rises abruptly from the sea to a height of c 750 m sloping west- eastwards with a succession of steep spurs emanating from the main summit which is situated on the western portion of the island. The very mountainous nature of island (there

4

ECOLOGY OF NARCONDAM HORN BILL

Fig. 2. Ridges and gular black band is absent in plicatus plicatus, everetii everetti and narcondami. Latter two are 1/3 the size of undulatus with plicatus being

intermediate (see text).

Abbreviations : A — Ridges; B — Black band.

<$

R. EVERETTI

CARL P.

6

R. NARCONPAMI

(5

R. PLICATUS

JOURNAL, BOMBAY NATURAL HIST. SOCIETY, Vol. 81

is virtually no continental shelf around the is- land) provides no landing place except for a small boulder-strewn bay on the southern side, which also provides the only small flat bit of ground for camping. A small spring in the bed of a dry nallah about 25 m above the sea level is the only fresh water source in the is- land known so far.

Climate :

The climate of the Andaman group of is- lands is tropical wet and humid with daily temperatures ranging from 27.8°C maximum and 21.8°C minimum. The rainfall is heavy both during SW and NE monsoons, lasting from May to October. Cyclonic storms occur during this period with rough weather con- ditions prevailing almost throughout the season. The average annual rainfall recorded for 17 years at Mayabunder (12°55' N; 92°55' E) the nearest weather station to Narcondam, is 3055.5 mm with an average of 13.4 rainy days per year. The month of July recorded highest average (538.5 m / 18.7 rainy days) and March lowest (4.8 mm / 0.4 rainy days).

Vegetation:

Parkinson (1923) and Thothathri (1960, 1962), and Balakrishnan (?) give some de- tails of the flora of the Andaman and Nico- bar group of islands. Prain (1893) described some aspects of the flora of Narcondam. The vegetation structure of the Narcondam island is more or less similar to that of the tropical N Andaman group. The vegetation can be divided into three categories (a) littoral (b) deciduous /evergreen and (c) moist evergreen. The very limited ‘shoreline’ of the island con- tains Ipomoea biloba, Scaevola koenigi. Hib- iscus tiliaceus, Panclanus sp., Thespesia popu - Inea, Barringtonia speciosa and Sterculia rubiginosa. Introduced plants like Coconut,

Papaya and Banana grow wild in this zone. The lower hills immediately following the ‘shoreline’ have both deciduous and evergreen trees. Some of the typical plants of this zone are Terminalia catappa, T. bialata, Parishia in- si gnis, and Cary ota mitis interspersed with numerous thorny creepers. The flora in the higher zones of the hill contains evergreens like Dipterocarpus sp-, Sideroxylon sp.. Ficus sp. etc. The vegetation still higher and close to the summit appears to be moist evergreen, with numerous epiphytes. Some of the seeds collected from a hornbifl’s nest were later identified as Anamirta cocculus, Capparis sepiaria, C. tenera var. latifolia, Garuga pin- nata, Amoora rohituka, Terminalia catappa and Ixora brunniscens. Apart from these, seve- ral other fruiting trees including the ones men- tioned above no doubt occur in the island.

Mammals :

No large mammals have been recorded in the island. Large rats ( Rattus sp.) obviously introduced, are common around the landing bay. Giant fruit bats (Ptcropus melanotus satyrus ) are common and other smaller bats may also occur.

Reptiles :

One of tne commonest snakes seen in the island is the flying snake Chrysopelia paradisi which is mostly arboreal. On the seashore oc- casionally sea snakes Laticauda colubrina are encountered. The giant water monitor Var anus salvator is common in different parts of the island. One specimen, which was collected, measured 1 m and weighed 4.5 kg. Skinks, Mabuya tytleri, Lygosoma maculatus and lizards, Cnemaspis kandiana, Cyrtodactylus rubidus and Phelsuma andamanense (endemic to Andamans) are common.

Land Crabs ( Car disoma hirtipes ) are very

6

J . Bombay nat. Hist. Soc. 81 Plate i

Hussain: Rhyticeros narcondami

Above : Narcondam island from western side. The central peak is perpetually under a shroud of cloud.

Below : Male feeding female (and young) at nest ‘B\

(Photos'. Pat Louis)

J. Bombay nat. Hist. Soc. 81 Hussain: Rhyticeros narcondami

Plate II

Above : Debris from nest ‘A’ (27/3/1972).

Below : A week old chick from nest (27/3/1972).

( Photos : S. A. Hussain)

ECOLOGY OF NARCONDAM HORN BILL

common throughout the island, occuring even higher up in the hills. Of the invertebrates, Danaidae; Nymphalidae; Lycaenidae (Butter- flies); Chryoschroa ignita, Mimila prenceps (Beetles); and spiders of the families Heterop- idae, Aregiopidae, and Thomsidae are record- ed.

Results

An attempt was made to locate as many nests of the hornbill as possible and to census

the population. A regular count of all the hornbills seen each day was made (see table 2). Increase in number of the females sighted may be due to their emergence from nest confinement after a successful brooding. It was not possible to identify all the nesting trees though a few nests were spotted on Sideroxylon sp. and Sterculia sp. Most of the nests were dis- covered from the debris and excreta and seeds below the nest-trees while a few others by observing the movements of the male bringing food to the nest.

Table 2

NARCONDAM HORNBILL

Date

No. of $ seen

No. of $ seen

No. of nests

No. of 8 9 at nest

17 / iii/76

25

Nil

3

3(3)

18

19

Nil

Nil

—

19

26

Nil

2

2(3)

20

14

Nil

Nil

—

21

28

1 (with $ )

Nil

—

22

31

4 ( 2 with 8 )

Nil

—

23

8

Nil

Nil

—

24

11

4

1

—

25

16

2

Nil

—

26

14

3

1

H8)

27

19

4 (2 with 8 )

Nil

—

28

16

2

1

H8)

29

59

11

Nil

—

30

72

28

1

—

31

40

10

—

—

l/iv/76

31

9

—

—

2

59

14

—

—

3

13

2

—

—

4

42

12

—

—

5

39

11

—

—

6

21

8

—

—

7*

—

—

—

—

8

48

9

—

—

9

23

4

—

—

10

52

11

—

—

* Rain

Note : The birds were counted randomly each day. The numbers may be biased on certain days as the birds congregating in feeding trees were counted as well as males on feeding forays may have been counted several times over !

7

JOURNAL, BOMBAY NATURAL HIST. SOCIETY, Vol. 81

3. Nest ‘A’ sketch

8

ECOLOGY OF N A RCON DA M HORN BILL

Nesting site :

A rough estimate of the heights of the nests observed varied between 2.4 m to 15.2 m. Two nests situated at 2.5 m and 2.74 m respectively were studied in detail. The nest ‘A' (Fig. 3) was situated on the outer bend of one of the main boughs of a tree facing west. The en- trance, though not concealed, was not easy to spot as the ground below the nest sloped down- wards steeply. The outer rim of the hole breadthwise measured 30 cm. Depth from the entrance to inner wall about 180 cm gradually tapering inwards. Nest i.T (Plate I) was on a bare tree facing east situated about 22.9 m from nest ‘A’. The ground rose into a steep ascent in front of the nest which enabled one to gaze directly into the nest hole from a cer- tain height. The entrance measured about 25 cm with a depth of about 149 cm. The flooor of the nest was horizontal. The contents re- moved from nest ‘A’ weighed 1360 g and con- sisted of eight varieties of seeds apart from feathers and powdered plaster. Some of the seeds were identified by the Botanical Survey of India.

The female and the young in the nest sat with their tail held up vertically. (They con- tinued to remain in this posture for quite some time even after they were removed from the nest.) The female attended to nest sanitation after every feeding visit of the male. She was observed tossing out what appeared to be the excreta of the young with her beak while she herself turned around and forcibly ejected her own excreta. On 6th April one of the chicks in the nest ‘B’ was seen making feeble attempts to defecate by bringing the anal region to- wards the nest entrance. Thereafter both the chicks regularly defecated in this manner.

Behaviour at nest :

The male starts fetching the food just be-

fore sunrise. No marked territorial behaviour by the breeding pair was observed. Occasio- nally an alien male or female was tolerated in the vicinity of the nest (i.e- on the same tree) though the minimum distance measured bet- ween two nests was about 22.8 m. Frequency of feeding varies with distance covered to the foraging tree. The shortest time recorded was 10 minutes and the longest 30 minutes. On arrival the male always perched on a particular branch of a tree depending on the direction of his arrival. If undisturbed, he would fly directly to the nest-hole, perching on a con- venient branch or clinging to the nest itself and proceed feeding the female. The food is coughed up, brought to the tip of the beak which is inserted into the slit opening and is offered to the female. The number of the in- sertions depends on the size of the food brought in. Large berries are offered piecemeal while smaller ones, whole. No attempt was made to retrieve the food that fell down in the course of feeding. A minimum of 10 insertions were counted when berries offered were large and a maximum of 93 when they were smaller. Some times the insertions are ‘false' when the female is not ready to receive the next berry. (Is she in turn feeding the young?) All this time the young would keep calling continuous- ly. Once the feeding was over the male would clean his beak on the branch a few times and after preening himself for a while fly away on the next foraging trip.

In the beginning of my observations the male refused to approach the nest in my pre- sence. He kept flitting from branch to branch and finally flew away. Fie seemed to rely on sight and showed no reactions to normal sounds but was wary of human voice. This particular male did not allow me to observe from any position below the line of its nest but allowed me to remain in full view at a dis-

9

JOURNAL , BOMBAY NATURAL LUST. SOCIETY, Vol. 81

*

tance of about 13-7 m, above the line of its nest. (This was possible as the ground rose upwards from the nesting tree).

It was not possible to ascertain the roles of the male and the female in nest building. The female in the nest ‘A’ was seen tamping the plaster of her nest by applying material with the sides of her bill on 18th March. The female sheds her flight feathers in the nest. The female taken out of the nest ‘A’ had 3rd, 4th and 5th primaries on the left wing and 3rd, 4th, 5th and 6th on the right in moult. The rest had fallen. Of the tail the 3rd pair was in moult. She weighed about 680 g, while the bill measured 108 mm, tarsus 43 mm, tail 198 mm, (moulting). She was found to be incap- able of flight.

General behaviour

Call :

In flight, adults of both sexes emit a con- tinuous ‘Ka . . ka . . ka’ to the accompani- ment of wheezing laboured wingbeat. When alarmed, the male at nest-site calls a halting ‘ko . . kokokoko . . ko . . kok . . ko kok kok kok’ etc. The female inside the nest is generally silent, but sometimes utters a single ‘krwak’ if the male is late in offering the next morsel during the course of a feeding. If alarm- ed herself she emits a repeated ‘Kraawk kok kok’ resembling the alarm call of a frightened domestic fowl. The young inside the nest call feebly ‘chew . . . chew . . . chew’ continuously like a squeaking sewing machine in operation, especially when the male is feeding.

Courtship :

On 27th March four males and three females were seen perching on different branches of a Ficus tree. All were calling simultaneously- One pair ( d* $ ) was more active than the

others. The female, which perched on the lower portion of a horizontal branch assumed begging posture towards the male perched a little higher next to her on the same branch. The male though silent now, occasionally a gave ‘krawk’ call and ‘touched’ the female’s bill and hopped away. Twice the male brought out a berry and offered it to the female. This went on for some- time as both kept hopping from branch to branch and finally flew away together. Several pairs ( d $ ) were seen together in the different parts of the island. This suggests that court- ship was still in progress.

Relations with other animals :

No predators have been recorded so far but rats and water monitors are the only large animals/ reptiles in the islands. Flying snakes ( Chrysopelia paraclisi ) are very common and on one occasion one was observed passing on branches very close to a hornbiH’s nest con- taining a female and young. Once several hornbills were seen mobbing a whitebellied Sea Eagle ( Haliaeetus leucogaster) from tree to tree. Abdulali (op. cit.) also mentions similar occurrence earlier. A koel ( Eudynamys scolopacea ) was also seen being chased by a hornbill. Human presence in the island is a re- cent phenomenon and though the impact of their presence throughout the year could not be assessed it may be assumed that the nesting pattern of the hornbill, may be affected as they would avoid nesting on lower available sites due to disturbance /predation by man.

Development of the young :

The egg (only one obtained) was earthy brown in colour. This may be due to staining. It measured 33 x 45 mm and weighed 28 g- The same nest contained a chick about a week old. It weighted 75 g and measured 130 mm from tip of the beak to vent (Plate II). The

10

J . Bombay nat. Hist. Soc. 81 Plate III

Hussain: Rhyticeros narcondami

On 26/4/1972 ‘A’ on right, ‘B’ on left.

( Photo : Courtesy Indian Express, Madras)

Above: ‘A7 <3* on 13/11/1972 c 9 month old. Below : ‘B’ $ on 13/11/1972 c 9 month old.

( Photos : S. A. Hussain)

J. Bombay nat. Hist. Soc. 81 Plate IV

Hussain: Rhyticeros narcondami

ECOLOGY OF N ARC ON DAM HORN BILL

body completely naked except about 10 rudimentary rectal barbs in an arc immediately above and between the anus and the oil gland. Similar barbs, numbering about 23 along the basal half of the wing along the alar tract. The upper mandible from gape to tip was c 25 mm, and the lower c 27 mm. The depth of the bill c 13.5 mm and the tarsus measured c 16 mm, the eyes were completely closed. Both the egg and the chick were preserved.

The exact age of the two other chicks obtain- ed could not be ascertained though it is as- sumed that the interval of hatching between the two was about 10 days, but this factor needs further investigation. The present obser- vations on the growth were made from the date (13 April) the birds were removed from the next.

Of the two, one was considerably larger and ultimately turned out to be male. It was not possible to ascertain the sexes of them at this stage as both seemed to have similar plumage i.e. rufous on head and neck. The chicks were named ‘A’ and ‘B’ for the sake of convenience of description through various stages of deve- lopment. (It became apparent in the final stages of growth that smaller ‘B’ though it started of with the rufous plumage of a male, acquired black plumage of the 9 after the post-juvenile general moult and thereafter be- came a full-fledged female). Descriptions of development recorded for the period April 1972 — March 1973 etc are given below. Body measurement and weights are given separately (see Figs. 4 and 5). The actual dates of measurements vary, though taken roughly dur- ing the middle of each month- There are some obvious gaps in data as I was away on other assignments during that period.

Both the parents and chicks, kept in a make- shift cage were brought to Port Blair and then on to Madras by ship and then to Bombay by

passenger train. The parent male died on board ship one week after capture. He had re- fused to eat. The female accepted food occa- sionally but did not feed the young which were kept in the cage along with her. The young however, fed voraciously. The female escaped from the cage when an enthusiastic reporter from a daily newspaper in Madras tried to photograph it in my absence. Though the news- paper sent out an appeal through its columns for information, she was never found and was believed to have died somewhere in the city of Madras. The chicks were brought to Bombay and were temporarily kept at Hornbill House. A cage measuring c 12' x 20' x 8' was subsequently built in the compound adjacent to Hornbill House and the pair remained there till their death 6 years later.

Development :

13 th April 1972 : ‘A’ — Rufous feathers on crown. Auriculars in sheath. Lores, area below the eye, nasal groove, hind-neck chin, throat and upper breast naked. The colour of skin in these areas smalt blue and rest of the area pinkish yellow. Vent, lower abdomen patchily feathered. All feathers in sheath. Upper tail coverts and lower back with a few barbs. Wing coverts well developed- A few feathers on the tarsus — oil gland swollen, and a line of feathers encircling it. Stomach greatly dis- tended. Wing and tail in moult. Bill waxy yellow. Gular pouch pale blue. Eyelashes well developed. Irides pale blue. Soles of feet pale blue. Wing 153 mm. Bill 72 mm, tarsus 47 mm, tail 96 mm.

‘B’ — A few barbs appearing on the crown, nasal groove; whole back, breast, abdomen naked. Wing coverts fully grown. A few tufts of feathers around oil gland. Bill 58, tarsus 41 tail 58, weight 380 gm.

Behaviour : Call monotonous and conti-

11

JOURNAL , BOMBAY NATURAL HIST. SOCIETY, VoL 81

nuous chew, chew, chew . . . Both ignored their parents and vice versa, though kept in the same enclosure. Defecation was carried out by stretching the neck out, raising the wings, projecting the anal region and stepping back- wards towards the edge of the enclosure. When on an open ground, the ‘stepping back’ is con- tinued till the faeces is discharged. Picking up and throwing about whatever object found nearby- Pecking at the toes of the observer (resemblance to seeds?). Both voracious caters, were fed on creamcracker biscuits, bananas, and other fruit.

May 1972 : ‘A’ — A line of rufous feathers in pin immediately below the gular pouch. Auriculars fully developed. Feathers on vent and abdomen fully developed. Upper back and lower hind neck patchily feathered. Base of the upper mandible swollen, showing a faint trace of wreath (furrow). Bill waxy yellow ex- cept at the base where it is reddish.

‘B’ — Crown, ear coverts, upper and lower tail coverts fully developed. The feathers adjoining these areas in pin.

September 1972: ‘A’ — All body feathers fully grown. Crown and nape dark rufous. Throat sulphur-yellow, grading into rufous to- wards the upper breast where it meets the black of the abdomen. The feathers of abdomen and tarsus softer than those on the back, wing coverts and scapulars. Gular pouch light blue. Bare skin around the eye smalt blue. Bill waxy yellow, the basal tinge of red increased in tone. A gap of about 2 mm bet- ween the mandibles about 8mm from the tip.

kB’ — - All body feathers fully grown. Fea- thers adjacent to the gular skin appear blackish. Lower neck where the black of abdo- men merging with the rufous of the neck seems to extend upwards. A few feathers on the crown have a barred appearance. The

rufous feathers on the head and neck dark greyish on the basal half.

November 1972: ‘A’ — Swollen casque of the wreath broad at forehead tapering towards the tip, about 72 mm in length. Depth of the bill including the wreath 50 mm.

‘B’ — Feathers on the crown (Centre streak) turning darker. A line immediately below and along the gular pouch black. Another streak across the ear coverts extending down to hind neck blackish. Swollen casque 50 mm. Depth of the bill including the casque 42 mm, gap between the mandibles about 2 mm.

December 1972 : ‘A’ — Depth of the bill 72 mm. Tip of the swollen casque blunted due to wear.

‘B‘ — Black feathers in sheath in a line above the eye (almost a central streak). Simi- lar streaks below the eye and extending up- wards from the black feathers in the abdomen and breast- Another line of black feathers ex- tending upwards from the upper back towards the crown. Rest of the neck area dark brown, depth of bill 43 mm, wreath 50 mm.

March 1973 : ‘A’ — Depth of bill 54 mm, 2 central tail feathers and 7th primary moult- ing. No body moult.

‘B’ — Extensive dropping of body and flight feathers. Heavy body moult. All the feathers moulting in the head and neck area black.

From March 1973, onwards the plumage of ‘R' showed a marked overall tendency to be- come black. By the end of May the moulting was over. The moulting of the wing and tail feathers was irregular. Both the birds shed their flight feathers irregularly, sometimes even freshly moulted ones. Powdered calcium sandoz was added to the regular diet during this period. By this time ‘B’ attained the full adult female plumage.

In August 1973 the transverse band on the wreath turned opaque and developed a soft

ECOLOGY OF N ARCON DAM HORN BILL

Fig. 4.

wrinkled depression where small blood capil-

laries were seen.

Soft parts : Colour of the irides remained pale grey in both and ? throughout while the eye lids of & turned red in colour. Bare skin around the eye and the gular pouch blue.

Measurements of wing, bill, tarsus, and tail were noted at intervals (Fig. 4). Weights were also noted for the same period (Fig. 5). The measurements of wing and tail were discon- tinued after March 1973, as their tips were

either breaking or wearing out as the birds constantly flew around in the cage.

Food and behaviour in captivity :

Both were fed on suttoo (powdered roasted Bengal gram) mixed with glucose powder and a few drops of ABDEC, supplemented with fruits like bananas, apple, guava, jamun ( Syzigium jambolana ) marshmelon, mango and sapota, hardboiled eggs and chopped meat was given in the initial stages.

13

JOURNAL, BOMBAY NATURAL HIST. SOCIETY, Vol. 81

Fig. 5. Weights of chicks (blank area — male, shaded area — female)

In Tockus hombills nestling weight reaches a peak at the completion of body growth indicated by tarsus and ulna measurements. Thereafter weight declines erratically untill about fledging time and once: again rises to a constant level (Kemp 1976). Figs. 4 & 5 indicate here that the body growth had reached a peak around the beginning of

April and Fig. 5 indicates possible fledging

Sporadic jerky movements of the head, a trait also observed in adults in the wild state. Occasionally tossing up the head and rubbing the crown on upper back. An occasional fruit or a morsel of food would be brought up and swallowed again. Hard seeds are regurgitated, preening is done throughout the day at irregular

period around mid-May.

intervals. Head is scratched by extending the wing, and bring up the leg over it, (indirect scratching) .

‘A’ ( cf ) was aggressive towards ‘B’ (?) and would not allow it to come near, often attacking and chasing it around the cage. There was no significant change in plumage

14

J. Bombay nat. Hist. Soc. 81 Hussain: Rhyticeros narcondami

Plate V

Above : Male about 4 years old.

Below : Male & female before the ‘fight’.

( Photos : S. A. Hussain)

ECOLOGY OF NARCONDAM HORN BILL

or body size / weight in the following years, but in February 1977 the iris of male appeared to change colour from grey to reddish. At the same time the female began to show interest in a nest box (prepared from a sawn-off tree trunk) placed inside the cage. The male how- ever did not show any sign of breeding con- dition. The female became more aggressive and finally both had to be separated by a partition inside the cage. It was possible that the female was imprinted on humans and re- garded the male as a threat to pair bond (A. C. Kemp, per. comm.). The female continued to be aggressive towards the male even after the normal breeding season (March-May) was over and finally on 8th March 1978 she manag- ed to sneak across the partition and attacked the male gripping his throat in her beak till she was separated with considerable effort. The male died the following day. The female was later sent to the Zoological Park at New Delhi where she died within two months of arrival-

Discussion

The very limited scope of the study carried out in Narcondam island restricts any detailed discussion. One of the factors that restricted a more detailed and systematic enquiry was that the expedition was mainly aimed at collecting biological specimens and as such most of the time was spent in that direction. However, it was possible to document, both in the wild as well as in captivity, some hitherto unknown aspects of the biology and ecology of the Narcondam Hornbill.

Ecological status :

Random estimates made by the earlier visi- tors to the island, i.e. c 200 birds is much less than what I believe to be the actual population. Daily counts were made by me during my

‘collecting’ forays (not more than 2 km in various directions in the island) and the birds seen in flight as well as on trees were recorded. The highest counted in one day was 72 males and 28 females (see table 2). These numbers include a point count made on a feeding tree where the hornbills were mobbing a white belli- ed sea eagle. My one month long stay in the island gave me the impression that there are more than 200 birds in the island. All the earlier visitors landed there in the months of March- April which happens to be the breeding period when most of the females would have been confined to nest-holes. However the figures mentioned in the above table cannot be in- terpreted statistically to project the probable population size since the factors contributing •to the regulation of population in the island are not known. What, then, is the optimum population and what is the factor that regu- lates it?

The apparent (?) absence of large predators in the island (including until recently, man) abundance of food resources with perhaps some degree of competition for resource from other frugivores in the island such as Green Imperial Pigeon ( Ducula aenea). Pied Imperial Pigeon (D. bicolor ) and the Giant Fruit Bat, provides an ideal ecological niche for a suc- cessful survival of a species like the hornbill. This is borne out by the fact that the Narcon- dam hornbills raise two chicks while most other larger species of Rhyticeros are able to raise only one chick (Kemp 1979). Com- petition for nesting sites during the breeding season may restrict the actual breeding success as the hornbills do not excavate nest holes. The severe cyclonic storms that lash these is- lands destroy a great number of older nesting trees, even causing mortality of brooding females /chicks confined in nest-holes- The com- petition for nesting sites may actually be acute

15

JOURNAL, BOMBAY NATURAL HIST. SOCIETY, Vol. 81

in the island as evidenced by the fact that two of the nests studied were as low as 2.5 to 2.7 m from the ground. Another possible regulat- ing factor may be perhaps the very insular na- ture of the species where in-breeding is discour- aged among siblings. The female chick's as- suming male plumage upto fledging period may actually be a mechanism evolved for this pur- pose. Was the antagonistic behaviour of the male towards the female initially and then the female towards the male once she attained breeding condition in captivity a part of the behaviour pattern evolved in the wild state to discourage mating among siblings? All this, is of course hypothetical and needs to be studied.

Conservation outlook

Island ecosystems are, in a way, living . laboratories for the study of evolution. The very simplified nature of the isolated islands provide us with an insight into the complexity of nature at work. Some of the factors that contribute to such systems are competition, predation, physical environment and their effect on the insular nature of small popula- tions inhabiting these (Diamond 1982). If any one of these factors is jeopardised there is a danger of such populations becoming vulner- able to extinction. It has been demonstrated repeatedly elsewhere that the single most im- portant cause for species extinction on oceanic islands has been predation by alien elements including man.

The vulnerability of the endemic birds is emphasised by the fact that most of the birds listed in the IUCN’s Red Data Book occur in islands. It is estimated that in the past 400 years two hundred of the estimated 220-odd species and subspecies of birds which have be- come extinct have been island forms. Most of these became extinct either because of the loss of habitat or when the population size was

too small to compete with alien competitors and predators, or even some catastrophy such as disease.

Where does the Narcondam hornbill stand under the present ecological conditions? It is definitely not facing the danger of extinction as yet but it is vulnerable and some conserva- tion strategy has to be evolved to protect the species from possible future exposure to the factors mentioned above. Even if one were to make a liberal estimate of the total population of the hornbills to be about 400 it will be per- haps too small a number to cope with ecologi- cal disasters like an outbreak of an epidemic or an extensive destruction of habitat. What, then could be the strategy to ensure the safety of this unique species?

First of all a complete ecological study of the hornbill is necessary to understand its status. Conservation measures based on such a study will be one of the answers to the ques- tion. Captive breeding, which has been success- fully carried out with several endangered bird and animal species elsewhere, is a tempting prospect- However, it may not be necessary in the case of Narcondam hornbill. On the con- trary, it would be much better to find out other islands within the Andaman group having simi- lar ecological structure and introduce the birds there. There are about 300 islands in the Andaman group of which about 60% are uninhabited. In the north Andaman group, which are close to Narcondam, there are seve- ral off-lying islands like Landfall, East, Inter- view and Barren having close affinities with the ecological conditions in Narcondam. A third alternative is to provide suitable artificial nesting sites in the island itself.

In conclusion it may be suggested that once the need to conserve the species against pos- sible extinction is sufficiently acknowledged, the following strategy may be instituted :

16

ECOLOGY OF NARCONDAM H0RNB1LL

1. Studying the complete ecology of the Narcondam Hornbill.

2. Emphasis on study of ecological require- ment, food niche, and the nesting suc- cess in the island.

3. Study of the habitat, vegetation struc- ture and faunistic composition in the island.

4. A comparative assessment of habitats of nearby islands.

5. Experimental capture and transfer of a few pairs of hornbills to alternate sites under careful supervision.

6. Monitoring the progress of introduced populations in their new habitats.

7. Declaring Narcondam and the island /s selected for transfer of the species as completely protected.

Acknowledgements

Under the direction of Mr. Humayun Abdulali, who first initiated the series of col- lecting expeditions to the Andaman and Nico- bar islands, Robert B. Grubh and R. J. Pimento of BNHS spent a few days on Nar- condam. They collected several specimens of the hornbill and attempted a rough count of its population. In 1971 Mr. Humayun Abdulali himself paid a fleeting visit to Nar- condam and took a few more specimens. Since then, following a spurious territorial claim by Burma, a police picket of 16 men has been posted on the island posing a poten- tial threat to the bird. Therefore, I am par- ticularly grateful to Mr. Abdulali for the opportunity he provided me for this trip to

Narcondam under the Charles McCann Ver- tebrate Zoology Fund for a field study of this unique species while it is still relatively safe and plentiful. I am also grateful to Mr. Har- mender Singh, the then Chief Commissioner, and Mr. S. Vajpayee, Chief Secretary of Andaman Administration; Mr. V. N. Singh IPS the then Superintendent of Police, Mr. Fred Burns, Manager, WIMCO (since deceas- ed); Mr. Bhaktawar Singh, Dy SP; the Master and crew of Police boat M. V. Jawahar (who cheerfully dropped us at Narcondam) the Radio Officer and the Police party, including the cheerful Nicobarese policemen for their as- sistance in various ways, Mr. N. J. George, of Prince of Wales Museum was of great help in collecting specimens and Mr. Pat Louis, who arrived with his cameras, provided a photo- graphic cover to the trip.

I am grateful to the then Honorary Secre- tary of the Society Mr. Zafar Futehally, and the Dynacraft Machine Co. for providing the cage for the hornbills at BNHS. Mr. J. C. Daniel and Dr. Salim Ali gave all encour- agement in my studies. Dr. A. C. Kemp of Transvaal Museum, S. Africa read through the manuscripts and gave useful suggestions and advice on hornbill studies and shared his knowledge about SE Asian hornbills. My col- leagues at the BNHS, specially, Messrs R. J. Pimento and Umapratap Singh helped in car- ing for the hornbills. The expenses for feeding the hornbills were met from a grant from Salim Ali/Loke Wan Tho Ornithological Re- search Fund. Mr. Carl D’Souza and Miss Usha Ganguli helped with graphs and sketches-

References

Abdulali, H. (1971) : Narcondam island and (1974) : The fauna of Narcondam

notes on some birds from the Andaman islands. island. Part I. Birds, ibid. 71: 496-505.

J. Bombay nat. Hist Soc. 68(2): 385-411. Ali, Salim & Ripley, S, D. (1970): Handbook

JOURNAL, BOMBAY NATURAL HIST. SOCIETY, Vol. 81

of the Birds of India & Pakistan. Vol. 4, O.U.P., Bombay.

Baker, E. C. S. (1927): Fauna of British India, Birds. Vol. 4, London. Taylor & Francis, London.

Blyth, E. (1845): Notices and descriptions of various new and little known species of India. Journal of Asiatic Soc. of Bengal 14 : 173-212.

Cory, C. P. (1902) : Some further notes on Nar- condam Hornbills (R. narcondami) . J. Bombay nat. Hist. Soc. 14: 372.

Diamond, J. M. (1982): Population processes operating on Islands. Island Management Sympo- sium 12 Aug. 1982, ICBP XVII World Conference, Cambridge.

Hume, A. O. (1873): Novelties, Stray Feathers 1: 411.

Kemp, A. C. (1976) : A study of Ecology Beha- viour and systematics of Tockus Hornbills (Aves: Bucerotidae) . Transvaal Museum, Pretoria.

(1979): A review of Hornbills:

Biology and Rediation. The living bird, 1978 Annual.

& Kemp, M. I. (1975): A report

on a study of Hornbills in Sarawak, with comments on their conservation. WWF Report.

Osmaston, B. B. (1905): A visit to Narcondam. J. Bombay nat. Hist. Soc. 16: 620-622.

Parkinson, C. E. (1923) : A forest flora of Anda- man Islands. Simla.

Peters, J. L, (1945): A checklist of the birds of the World.

Prain, D. (1893) : On the flora of Narcandam and Barren Islands. J. Asiatic Soc. Bengal 62: 39-86.

Ripley, S. D. (1982): A Synopsis of the Birds of India and Pakistan. 2nd Ed. Bombay Natural History Society.

Sanft, K. (1960) : Bucerotidae. Das Tierrich 76.

St. John, J. H. (1898) : Some notes on Narcon- dam Hornbill. J. Bombay nat. Hist. Soc. 12: 212-214.

Thothathri, K. (1960): Studies on the flora of Andaman Islands. Bull. Bot. Surv. of India 2:357- 373.

(1962): Contribution to the

flora of the Andaman and Nicobar Islands, ibid. 4:281-296.

18

SEASONAL VARIATION IN THE POPULATION OF AC RIDA EX ALT AT A WALK. AT ALIGARH1

Shamshad Ali2 (With seven text-figures)

The population level of Acrida exaltata over three years (1974-76) at Aligarh has been discussed. Information is furnished on seasonal variation, intra and inter year fluctuation and life cycle in natural conditions. Climatic conditions exert marked in- fluence on the rise and fall of population. The timing of the various life history events (i.e. oviposition, hatching and maturation) varies widely from year to year depending upon the particular sequence of climatic conditions prevailing throughout the entire grasshopper cycle.

Introduction

Acrida exaltata Walk, is a serious pest of cotton and tobacco. Besides cotton and to- bacco, it also attacks rice, sugarcane, potato and grasses. It has long been recognized that the wide fluctuations periodically occurring in acridid populations throughout the world are closely linked to weather conditions (Parker 1935, Dempster 1963). The major weather factors involved are apparently temperature and precipitation. In some characteristically very dry regions, rainfall may be the principal limit- ing factor in grasshoppers distribution through its influence on food (Scharff 1954), breeding behaviour (Uvarov 1956). Putnam (1954) said that grasshoppers outbreak usually coincide with extended period of hot, dry weather. Des- camps (1975) studied factors influencing the distribution and abundance of acridid popu- lation in general.

Studies were made to note the seasonal

1 Accepted July 1980.

2 Section of Entomology, Department of Zoology, A.M.U. Aligarh. Present address : Assistant Professor, Institute of Biology, University of Constantine, Algeria.

variation in the population of Acrida exaltata Walk., due to various environmental factors at Aligarh.

Material and Method

The field observations were undertaken for three years from January, 1974 to December, 1976, during different months of the year. The samples of hoppers and adults were obtained by sweeping. A standard net was used for collection. The insects were collected in the morning on every tenth day for an hour from the acridid field station (Scindia Fort, Ali- garh). Meteorological records were obtained from the weather station. Department of Physics, Aligarh Muslim University, Aligarh. Data on peak density recorded each month in the area for the period (1974-76) were used for analysis. These monthly peaks were then analysed for the three months, each con- stituting four seasons. Winter (December to February), Spring (March to May), Summer (June to August) and Autumn (September to November). Only the mean values of various seasons were considered to reveal intra and inter year fluctuations. The reason for using

19

JOURNAL, BOMBAY NATURAL HIST. SOCIETY, Vol. 81

seasonal instead of monthly population was to obtain ‘nil’ population values, invariably en- countered for months at a time, specially during Winter and Spring seasons.

Inter year fluctuation was measured by the deviations of the seasonal mean from that of annual mean for three years. Comparative be- haviour of two types of fluctuations was also studied.

Studies were made on the life history of Acrida exaltata Walk, under natural condi- tions.

Observations

Topography : The geographical position of

Aligarh is 27° 53' 38"N. Latitude and 78° 04' 30"E. Longitude. The district of Aligarh lies in the upper Doab of the Ganga and Jamuna rivers.

Climate : Aligarh experiences tropical monsoon type of climate. The year is generally divi- ded into the following three seasons —

1. The cold weather: Winter (Late Octo- ber to February),

2. The hot weather season: Summer

(March- June),

3. The season of general rains: (Mid- June-September).

During winter the temperature is generally

low. The mean maximum temperature is 80°F, however, the mean minimum temperature re- mains around 50°F. The prevailing direction of wind during the season is from West and North-West to South and South-East. The winds are generally light with an average speed of 2 miles/hour. These winds are supposed to be of continental origin and are mostly dry. The month of December and January are the coldest and often register light rains due to western disturbance otherwise the weather is generally fine and pleasant due to bright and sunny days with clear sky. The month of May and June are the hottest with mercury shoot- ing sometimes upto 115°F, however, the mean maximum temperature is 115°F and the mean minimum temperature 65°F. Strong dust rais- ing hot and dry westerly winds during day time is common feature of the summer. The peculiar phenomenon of the summer is the frequent occurrence of dust and thunder storms with an average velocity of 30-40 miles /hour gales. The humidity sometimes falls to 2 or 3% whereas the general level is 20%.

With the onset of monsoon generally by late June, the direction of winds is reversed due to low pressure area developed in the north western India. With the arrival of the humid oceanic currents from the Arabian Sea as well as from the Bay of Bengal, the tempera-

Table 1

Average monthly rainfall at Aligarh (1974-76)

Year

January

February

March

1 .

April

May

y

c

3

(in mm)

July

August

Seplembe

October

Novembc

Decembei

1974

0.0

0.0

0.0

0.6

20.9

31.8

230.9

193.4

5.5

19.6

0.0

17.2

1975

19.4

10.0

0.7

0.0

23.7

74.1

247.9

146.4

312.6

68.1

0.0

0.0

1976

0.0

13.0

5.0

11.8

22.6

35.7

354.4

426.4

73.9

0.0

0.0

0.0

SEASONAL VARIATION IN THE POPULATION OF ACR1DA EXALTATA

120

ioc£

-o

E

80 2

>

60

&

O'

40 c

if

20

Days

FIG-1 CLIMATIC DATA 1974

ture falls and the air becomes cool. The mean monthly temperature falls to 80°F in July. The relative humidity increases to 70-74% R.H. The sky is generally overcast in the rainy season. This season receives nearly 90% precipitation of the whole year and the mean seasonal ranfall is 25" (Table 1 and Figs. 1, 2 and 3).

Vegetation : Water penetration plays an impor- tant part in determining the distribution of vegetation. Scindia Fort is rich in green veeetation. Abundance of food is available for feeding by grasshoppers, and is surrounded by cultivated crop fields.

It was found that hoppers and adults were most abundant during and after the monsoon period (July-October) due to the optimum ecological conditions, particularly temperature, relative humidity and food for their develop- ment and biological activities. As is evident from Figs. 4, 5 and 6 the population was lowest in winter (December-March) and summer (May-June). This is due to slow reproductive activities during this period. Copulation was observed to be higher in July to October, Ovi- position was also higher.

Egg pods laid during April to June, hatched after the monsoon showers in July since suffi-

2\

JOURNAL, BOMBAY NATURAL HIST. SOCIETY, Vol. 81

cient moisture was available for the develop- ment of eggs. But during the heavy rains, hatching of eggs decreased, but again increased in August to October. The population of hop- pers and adults was at its maximum from July to October, but decreased in subsequent months due to the advent of winter (Figs. 4, 5 and 6). This decrease continued till February.

Seasonal Life History at Aligarh under Natural Conditions :

Copulation : From 15th March to 31st Octo- ber. Maximum in July, August and Sept- ember.

Oviposition : From last week of March to middle of November. Maximum in August and September.

Hatching : From first week of March to Dec- ember last maximum hatching was observ-

ed during the months of August and September.

Hopper Stage : Found throughout the year. Large number of nymphs were found during the months of August, October and March.

Adults were found throughout the year. Large number of adults were found in Octo- ber, November and April.

Climatic conditions were suitable for the re- production only for a short period at the end of the dry season. During the rainy season, it results in rapid growth of grass cover, which favours the grasshopper; the temperatuie and humidity remain favourable for about two months. However, the heavy rains in July and August increased soil humidity and the Oothecae were destroyed and from November temperature frequently falls below the thre-

Days

FIG. 2 CLIMATIC OATA 1975

22

SEASONAL VARIATION IN THE POPULATION OF ACRIDA EXALTATA

100 »>

• «S>

TJ

• «*

80 i

<2

60 I

QC

40 ^ c 01 u

20 l

F M A M

J J A Days

FIG- 3 CLIMATIC DATA 1976

S 0 N 0

shold for development. During the dry season, low temperatures induce degeneration of the oocytes, disrupt spermatogenesis and change the sex ratio in adult populations. It was found that the populations in the area was maintain- ed only by migration from other localities.

Inter and Intra Year Fluctuations :

Apparent intra year fluctuation of the mean peak density populations in respect of the Winter, Spring, Summer and Autumn seasons are shown for three years from 1974-76 in Fig. 7. The highest peak usually occurs during the Autumn season with the characteristic sharp fall in the next winter. However, for the years 1975-76 Summer peaks were maximum.

Discussion

In the present three year study, the rela- tionship between egg production and tempera-

ture was particularly apparent during the month of September which showed extremely variable climatological conditions from year to year. September is a transitional period bet- ween Summer and Winter, when rapid change in temperature again occur. In order, grass- hoppers may take advantage of the optimum egglaying periods during mid Summer and so attain their maximum reproductive potential, it is essential that they mature early in the season. Early maturation is dependent upon a continued sequence of high temperatures through all stages of development extending even back to the previous fall following depo- sition of eggs. Continued high temperatures permit rapid development of nymphs to the adult stage followed by advanced matu- rity, early mating and oviposition. On the other hand, a complete reversal of the weather patterns just outlined, with consistently low

23

JOURNAL, BOMBAY NATURAL HIST. SOCIETY, Vol. 81

240

i "" ■ ■

Jan Feb Mar Apr May Jun July Aug Sep Oct Nov Dec

FIG-4 MONTHLY POPULATION -1974

■24

Number

SEASONAL VARIATION IN THE POPULATION OF ACR1DA EXALTATA

220

200

180

160

140

120

100

80

60

40

20

□ Nymphs ■ Adults

Jan Feb Mar Apr MayJun July Aug Sep Oct Nov Oec

FIG- 5 MONTHLY POPULATION -1975

25

Number

JOURNAL, BOMBAY NATURAL HIST. SOCIETY, Vo! . 81

240

Jan Feb Mar Apr May Jun July Aug Sep Oct Nov Dec FIG. 6 MONTHLY POPULATION -1976

26

Number

SEASONAL VARIATION IN THE POPULATION OF ACRIDA EXALT AT A

Winter Spring Summer FIG. 7 SEASONAL POPULATION

74 7 5 76 Autumn

(1974-76)

JOURNAL, BOMBAY NATURAL HIST. SOCIETY, Vol. 81

temperatures through the various stages of growth and development will result in drastic reduction of the number of eggpods laid. In eggs laid after mid September the percentage of hatching was drastically reduced, these eggs while still apparently viable showed only minor embryonic development.

Similar observations were made by Suslik (1975), who observed that seasonal dispersal of grasshopper was found to depend on the height of the grass stand, and population den- sity varied at different times of the year. Pick- ford (1970) gave the reason for population increase as due to favourable environmental conditions, which resulted in increased fecun- dity and survival of more eggs Randell and Mukherji (1974) observed that increase in population was due to high temperatures at the time of egglaying.

Temperature during peak population density was highly significant (Edwards 1960). Increase in population due to high temperature at the time of egglaying was also supported by Ran-

R E F E

Albrecht, F. O. (1956): Limitations des effectifs chez un acridien. Influece de la secheresse du sol sur les oeufs de Nomadacris septemfaciata (Serv.). Locus ta, 4 : 1-21.

Cassimir, M. (1962): History of outbreaks of the Australian plague locust, Chortoicetes termini- fera (Walk.) between 1933 and 1959, and analysis of the influence of rainfall on their outbreaks. Aust. J. Agric. Res., 72(4): 674-700.

Dempster, J. P. (1963) : The population Dyna- mics of grasshoppers and locusts. Biol. Rev., 38: 490-529.

Descamps, M. (1975): Etude du penplement acri- dien de 1 Etat de veracruz (Mexique). Folia Ento- mologica Mexicana. 31/32 : 3-98.

Edwards, R. L. (1960): Relationship between grasshopper abundance and weather conditions in Saskatchewan, 1930-58. Canad. Ent., 92(8) : 619-624.

Parker, J. R. (1935): Factors largely responsible for years of grasshoppers abundance. Proc. World’s Graon Exhibition Conf., Ottawa, 2: 472-473.

dell and Mukherji (1974). Heavy rain in an outbreak area gave rise to increase in popula- tion (Cassimir 1962). Descamps (1975) studied factors influencing the distribution and abun- dance of acridid population in general.

This study on seasonal variation indicates that in Acrida exaltata Walk. Populations in- directly affected by climatic factors through their effects on the sexual activity of the adults. It is clear that mating is restricted or even curtailed during periods of cool, cloudy wea- ther.

Acknowledgements

I am highly indebted to Prof. S. M. Alam, Head, Department of Zoology, A. M. U. Aligarh for providing financial assistance and encouragement. Thanks are also due to Mr. Abdul Qayyum, Department of Physics, A. M. U. Aligarh for his help in collecting climatic data and to University Grants Com- mission, New Delhi for providing financial assistance.

R e n c e s

Pickford, R. (1970): The effect of climatic fac- tors on egg survival and fecundity in grasshoppers. Proc. International Conf. Acridids: 257-260.

Putnam, L. G. (1954): Development in grass- hopper research and control in Canada. Rep. 6th Common. Ent. Conf., London: 133-137.

Randell, R. L. & Mukherji, M. K. (1974): A technique for estimating hatching of natural egg population of Melanophus sanguinipes (Orthoptera: Acrididae). Canad. Ent., 106(8): 801-812.

Scharff, D. K. (f954) : The role of food plants and weather in the ecology of Melanoplus mexica- nus mexicamis Sauss. J. econ. Ent., 47 : 485-489.

Suslik, V. (1975) : Disperzia, abundanica a bio- rnasa konikovz celade acrididae (Orthoptera) na pasienkupri Basukej staavnica. Biologia Czechoslo- vakia, 39(11) : 847-851.

Uvarov, B. P. (1956): The locusts and grass- hoppers problem in relation to the development of arid lands. The Future of Arid Lands, 43: 383-389.

28

LARVAL CULTURE OF THE HERMIT CRAB CUBAN ARIUS AEQUABILIS VAR. MERGUIENSIS DE MAN (DECAPOD A, ANOMURA, DIOGENID AE ) REARED IN THE LABORATORY1

Venkatray N. Nayak2 (With seven text-figures)

The paper describes the complete life history of an intertidal hermit crab, Clibanarius aequabilis var. merguiensis, as observed in the laboratory. The larvae pass through four zoeal stages and a glaucothoe stage. All the developmental stages are fully illustrated and described. Characters of generic importance are listed.

Introduction

The present paper deals with the complete larval history of Clibanarius aequabilis var. merguiensis de Man as observed in the labo- ratory.

Materials and Methods

An ovigerous female of C. aequabilis var. merguiensis was collected on September 16, 1974 from a tide pool of Kinkade rocky shore along the west coast of India, and was kept in a glass trough containing filtered sea water until the larvae hatched out on 28th September 1974. The larvae were separated into groups of 5 per bowl with approximately 150 ml of sea water. Newly hatched Artemia nauplii were added as food. Every day the larvae were transferred to fresh sea water. Exuviae and the dead larvae were removed regularly and were preserved in a special preservative (Thakur 1960). During the course of the ex- periment the temperature ranged from 25° to 28°C and salinity about 25 ppt.

1 Accepted September 1981.

2 Department of Marine Biology, Kamatak Uni- versity, Kodibag, Karwar-581 303. Present address : Government Arts and Science College, Karwar- 581 301, India.

The larvae were dissected in 5% glycerine under a binocular microscope. All the draw- ings were prepared with the aid of a camera lucida. The total length of the larva was measur- ed from the tip of rostrum to median posterior margin of telson. The carapace length was measured from tip of rostrum to postero- dorsal margin of carapace. The term ‘stage’ is used herein in the sense of instar or inter- moult.

The eggs are oval, violet to pinkish (when young) turning transparent and pale when about to hatch; egg size ranged from 0.37- 0.44 x 0.30-0.38 mm.

Results

The larvae of Clibanarius aequalibis var. merguiensis de Man reached the glaucothoe after passing through four successive zoeal stages in about 22 days after hatching. A sum- mary of the various instars and duration of instars is shown in Table 1.

Table 1

Larval stage

Duration in days

I zoea

6 — 8

II zoea

4 — 7

III zoea

5 — 8

IV zoea

6 — 9

Glaucothoe

None moulted to crab

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JOURNAL, BOMBAY NATURAL HIST. SOCIETY, Vol. 81

Fig. 1. First zoea, Clibanarius aequabilis var. merguiensis de Man. a, entire larva; a 1? rostrum; b, antennule; c, antenna; d, mandibles; e, first maxilla; f* second maxilla; g. first maxilliped; h, second maxilliped; i, third maxilliped; r, telson.

THE HERMIT CRAB CLIBANARIUS AEQUABILIS VAR. MERGUIENSIS

Chromatophores :

The chromatophores of the larvae are of diffuse type, giving orange yellow to red appea- rance to the larvae. The chromatophores are stellate with mainly orange, crimson red and light yellow components.

Description of larval stages :

First Zoea (Fig. 1)

Carapace length: 1.0 mm; Abdomen length: 1.2 mm

Rostrum beak like, acutely pointed at the tip and broad at the base, projects beyond the antennule and antenna (fig. 1, a); eyes sessile; carapace smooth, rounded on postero-lateral angle, 4 exopod setae each on first and 2nd maxillipeds; third being rudimentary; abdomen nearly as long as carapace, inclusive of rostrum; telson process formula 7+7, 1st slightly laterally situated and blunt, finger-like.

Antennule (fig. 1, b) with 2 terminal aesthe- tascs and 2 unequal setae; inner ramus repre- sented by a long, plumose seta. Antenna (fig. l,c): Endopod nearly 2 /3rd the length of scale, with 3 terminal setae; scale, long and narrow with 11 plumose setae; endopod and scale distinctly articulated to peduncle; pedun- cle with a minutely serrated ventral spine on the distal margin. Mandibles (fig. l,d) asym- metrical and stout; ventral plate with 4-5 large but unequal teeth in the middle, whereas the dorsal provided with several unequal small teeth all along the edge. First maxilla (fig. l,e): Coxal endite with 6 setae of which 3-4 simple and others bristle-like; basal armed with 2 serrated spines and one short spine-like seta; unsegmented palp with 2 terminal and 1 subterminal setae. Second maxilla (fig. l,f): Coxal and basal endites bilobed; proximal of coxal with 5 terminal and single sub- terminal setae; distal with 3 terminal and 1 subterminal; proximal of basal with 4

and distal with 3 terminal and a short sub- terminal setae; endopod with 2 groups of 2 setae each; scaphognathite bears 5 marginal plumose setae. First maxilliped (fig. l,g): Endopod nearly as long as exopod, 5-segment- ed, setation being, 1, 2, 1, 2 and 4+1 (outer) distalwards; exopod 2-segmented with 4 nata- tory setae; basis with 8 setae as in figure. Second maxilliped (fig. l,h): Endopod 4- segmented, 3rd being the longest, setation, 2, 2, 2 and 4+1 (outer) distalwards; exopod as in first maxilliped; basis with 4 setae as illus- trated. Third maxilliped (fig. 1, i) rudimentary and uniramous. Abdomen (fig. 1, a) 5-seg- mented, segments smooth and broader than long. Telson (fig. 1, r) broader than long, pro- cess formula 7 + 7; 1st finger-like process arti- culated and situated laterally; 2nd reduced ‘anomuran’ hair; 3rd to 7th, plumose setae; 4th being the longest, 7th alone spinose on the outer margin; posterior margin of telson and median notch fringed with microscopic hairs. Second Zoea (Fig. 2)

Carapace length 1.2 mm; Abdomen length 1.6 mm

Larvae increase in size; eyes stalked; num- ber of setae on exopod of first two maxillipeds increased to 6; exopod of third maxilliped well developed with 5 setae, but endopod bud-like; telson process formula 8 + 8.

Antennule (fig. 2, b): Peduncle with 3 long plumose setae at its distal margin; outer ramus now distinctly articulated with peduncle, bear- ing 2 aesthetascs (of which one is quite pro- minent) and 3 unequal setae. Antenna (fig. 2, c) : No considerable change over previous stage. Mandible (fig. 2, d) : No change except for a slight increase in size and prominence of teeth. First maxilla (fig. 2, e) : Coxal endite and endopod unchanged; basal now with 4 serrated spines and 2 short setae. Second maxilla (fig. 2, f ) : Coxal and basal endites

31

JOURNAL, BOMBAY NATURAL HIST. SOCIETY, Vol. 81

a, entire larva; b. antennule; c, antenna; d, mandibles; e, first maxilla; f. second maxilla; g, first maxiliiped; h, second maxilliped; i, third maxilliped; r, telson.

32

THE HERMIT CRAB CLIBANARIUS AEQUABILIS VAR. MERGUIENSIS

with 7 + 4 and 4-5 + 3 setae respectively; scaphognathite now with 6-7 marginal setae; no change in endopod. First maxilliped (fig. 2, g) : Exopod with 6 natatory setae; addition of a plumose seta each on the outer margin of first 3 segments of endopod. Second maxi- lliped (fig. 2, h): Exopod as in 1st maxilliped; addition of 1 seta each on 2nd and 3rd seg- ments of endopod. Third maxilliped (fig. 2, i) : Exopod well developed, partially 2-segmented with 5 natatory setae; endopod as indistinct bud. Telson (fig. 2, r) : Slightly broader than long; process formula 8 + 8; median pair being spinulose distally.

Third Zoea (Fig. 3)

Carapace length: 1.4 mm; Abdomen length: 1.7 mm

Zoeae increase considerably in size and can be distinguished by the following characters: antennal endopod with a single seta; 3rd maxi- lliped now with 6 setae; 4 pairs of pereiopod buds developed; abdomen 6-segmented; telson process formula 8 + 1 + 8, 4th process reduced to a spine; uropod biramous.

Antennule (fig. 3, b) : Peduncle with 4 long, plumose and 4 fine hair-like setae distally; inner ramus separated from peduncle; outer now with 3 aesthetascs and 4 unequal setae. Antenna (fig. 3, c) : Endopod elongated, reach- ing upto the tip of the scale and with a single seta; scale with 12 setae. Mandible and first maxilla (fig. 3, d & e) as in previous stage. Second maxilla (fig, 3, f) : Coxal and basal endites with 8 + 4 and 4 + 3 setae respectively; scaphognathite with 10 plumose setae; no change in endopod. Maxillipeds (fig. 3, g, h & i): No change in 1st and 2nd; exopod of 3rd with 6 natatory setae; endopod slightly elongated. Pereiopods (fig. 3, k) : Four pairs of rudimentary buds clearly seen. Abdomen (fig. 3, a): Sixth segment separated from tel- son; Telson (fig. 3, r) : Somewhat rectangular

in shape; process formula 8 + 1 + 8; 4th pro- cess now reduced to an unarticulated spine; all the processes plumose except the 1st and 4th, somewhat spinose at the tip. Uropods (fig.

3, r) biramous, with a functional exopod and a rudimentary endopod; exopod with 8 plu- mose setae on its posterior margin; endopod present as bud.

Fourth Zoea (Figs. 4 & 5)

Carapace length: 1.7 mm; Abdomen length:

1 . 9 mm

This stage exhibits following advanced features over the previous stage: inner ramus of antennule now gets distinctly separated from peduncle; mandibular palp developed as rudi- mentary bud; 5 pairs of pereiopod buds; 4 pairs of pleopod buds; endopod of uropods functional.

Antennule (fig. 4, b) : Peduncle now with 4 long plumose and 3 hair-like setae distally; inner ramus clearly separated and tip devoid of setae; outer with 3 aesthetascs and 4 un- equal setae terminally. Antenna (fig. 4, c) : Endopod 2-segmented, about l\ times longer than scale and with a single terminal seta. Scale with 13 marginal setae. Mandible (fig. 4, d) : Palp developed as bud.FzV.st maxilla (fig.

4, e) : No change except for the addition of a plumose seta on the coxal endite. Second maxilla (fig. 4, f ) : Coxal and basal endites with 8-9 + 3 and 4-5 +5 setae respectively; endopod being unchanged; scaphognathite fringed with about 14 setae. Maxillipeds (fig.

5, g, h & i) : 1st and 2nd unchanged; endopod of 3rd segmented with a terminal seta. Pereiopods (fig. 5, k) represented as 5 pairs of elongated buds showing partial segmentation, 1st being chelate and 5th subchelate. Abdo- men (figs. 4, a & 5, 1) : Four pairs of pleopods present as buds from 2nd to 5th segments. Telson (fig. 5, r) : Process formula 8+1 + 8; 4th process still present, but reduced. Uropods

33

3

JOURNAL, BOMBAY NATURAL HIST. SOCIETY, Vol. 81

Fig. 3. Third zoea, Clibanarius aequabiiis var. merguiensis de Man. a, entire larva; b, antennule; c, antenna; d, mandible; e, first maxilla; f, second maxilla; g, first maxilliped; h: second maxilliped; i, third maxilliped; k, pereiopods;

r, telson.

THE HERMIT CRAB CLIBANARIUS AEQUABILIS VAR. MERGUIENSIS

Fig. 4. Fourth zoea, Clibanarius aequabilis var. merguiensis de Man. a, entire larva; a1? rostrum; b, antennule; c, antenna; d, mandibles; e, first maxilla;

f, second maxilla.

35

JOURNAL, BOMBAY NATURAL HIST. SOCIETY, Vol. 81

Fig. 5. Fourth zoea, Clibanarius aequabilis var. merguiensis de Man. g, first maxilliped; h, second maxilliped; i, third maxilliped; k. pereiopods; 1, pleopod;

i, telson and r1? uropods.

THE HERMIT CRAB CLIBANARIU5 AEQUABILIS VAR. MERGUIENSIS

(fig. 5,1*0: Both exopod and endopod func- tional; exopod rather oval with 9-10 plumose setae; endopod with 4-6 setae.

Glaucothoe (Figs. 6 & 7)

Carapace length: 1.2 mm; Abdomen length:

1 .9 mm

Rostrum short and broad at the base; abdo- men about 1J times the length of carapace; chelipcds more or less equal; eye-stalks nearly twice as long as broad and almost reaching the distal end of antennular and antennal peduncle; ophthalmic scales small; fourth and 5th pereiopods smaller, subchelate and chelate respectively; 4 pairs of biramous pleopods; uropods slightly unequal.

Antcnnule (fig. 6, b) : Peduncle 3 -segment- ed, first segment somewhat globular, second and third segments somewhat cylindrical; outer ramus 5-segmented with 0, 5-6, 4 and 2 aesthe- tascs respectively on 1st to 4th segments, distal wards and 3-4 setae on 5th, in addition to few simple setae on 2nd and 3rd segments. Antenna (fig. 6, c) : Peduncle 5-segmented; scale with 3 outer marginal and 2 distal setae, reaching nearly to the distal margin of 4th segment, with a pointed process on the outer distal angle; 9-segmented flagellum with 0, 3, 3, 3, 4, 3, 4, 5 and 7 setae from 1st to 9th segments distalwards. Mandible (fig. 6, d) : As in adult; palp 3 -segmented with about 12 bristle-like setae distally on terminal segment. First maxilla (fig. 6,e): Both coxa and basis membranous bordered with setae and plumose hairs; coxa with about 19 setae and basis 15-17 setae; inner margin of basis with 2 simple setae; endopod short and unsegmented with a short knob-like projection terminally on the outer side bearing a single seta; inner margin with 1 or 2 setae. Second maxilla (fig. 6, f) : Similar to adult except that palp is simple; scaphogna- thite well developed and fringed with about 50 setae. First maxilliped (fig. 6, g) : As in

adult, except that exopod lacks the terminal flagellated portion. Second maxilliped (fig. 6, h) : Of the endopod segments merus is the longest as in adult; exopod shows distinct flagellated and nonflagellated portions, flagellat- ed portion 3 -segmented bearing 5 to 6 plumose and a few simple distal setae. Third maxilliped (fig. 6, i) : Distal 2 segments of endopod bear many setae, remaining 3 segments with 2 + 2, 2+1 and 4 + 1 setae distalwards; the flagellated portion of exopod do not show clear segmen- tation and with 6 plumose setae terminally. Pereiopods (fig. 7, k2-5): First pair of cheli- peds (fig. 7, kj), as in adult, distinctly chelate, in almost horizontal plane; merus is the lon- gest segment; carpus somewhat triangular, 1 /3rd the length of merus; propodus longer than broad and devoid of tubercles; fingers hoofed; dactylus more or less equal to propo- dus in length with slightly curved but somewhat hoofed tip; few scattered setae on all segments but no spines and tubercles. Second pereio- pods (fig. 7, k2), more or less similar; seg- ments long and cylindrical; propodus is the longest segment; carpus 1/2 the length of pro- podus; dactylus nearly Jth the length of pro- podus, unlike in adult wherein dactylus is 1.7 times propodus, with 2-3 spinules on its poste- rior margin. Third pereiopods (figs. 7. k3) similar to the second leg except for 2-3 spinu- les on its posterior margin. Fourth pereiopods (fig. 7, k4) quite small, smaller xhan fifth leg; merus and carpus more or less of equal length; propodus as broad as long and with about 4-6 pectinate granules and tufts of setae distally; dactylus claw-like with no spines but bearing tufts of setae and a long seta. Fifth pereiopods (fig. 7, k5) minutely chelate; merus being the longest segment; carpus nearly 2/ 3rd the length of merus and bears few setae on either mar- gins; anterior part of propodus and proximal part of dactylus show corneous granules; the

37

JOURNAL, BOMBAY NATURAL HIST. SOCIETY, Vol. 81

Fig. 6. Glaucothoe of Clibanarius aequabilis var. merguiensis dc Man. a, lateral view of entire larva; a1? dorsal view of entire larva; b, antennule; c, antenna, d, mandible; e, first maxilla; f, second maxilla; g, first maxilliped; h, second maxi

lliped; i, third maxilliped.

38

T

THE HERMIT CRAB CLIBANARIUS AEQUABILIS VAR. MERGUIENSIS

Fig. 7. Glaucothoe of Clibancuius aequabilis var. merguiensis de Man. kj, cheliped; k2, second pereiopod; k„. third pereiopod; k4, fourth pereiopod; k5 fifth pereiopod; 1, pleopod; r. telson and uropod; rv telson

JOURNAL , BOMBAY NATURAL HIST. SOCIETY, Vol. 81

latter forming a chelate structure with dacty- lus; very long setae both on propodus and dactylus. Abdomen (fig. 6, a) nearly \\ times as long as carapace; 6-segmented; 2nd to 5th segments with a pair of pleopods each. Pleo - pods (fig. 7, 1) biramous with a long pedun- cle; a setose exopod and 2 hooks in the inner margin of endopod. Telson (fig. 7, r) some- what obtuse, slightly broadening anteriorly, with about 9 plumose setae on its plain poste- rior margin and 2 pairs of small simple setae laterally and dorsally and a pair of submar- ginal setae posteriorly. Uropods (fig. 7, r) well developed; both the rami being somewhat rectangularly triangular and armed with cor- neous granules on the posterior end; with 14 plumose setae on the posterior and inner lateral margins of exopod and about 11 on posterior margin of endopod.

Discussion

Based on the hitherto described larvae, the features of generic importance of Clibanarius may be summarised as under for first zoeal stages : carapace and abdominal somites

smooth; rostrum long, broad at the base, blunt or acutely pointed at the tip; reaching beyond the antennule and antenna, beak-like; antennal scale without a terminal spine, endopod with three long plumose setae; telson deeply notched; first process laterally situated, blunt, finger-like except for C. erythropus wherein it is a small spine.

Of the three species wherein published in- formation on the laboratory reared larvae in the genus Clibanarius is available, the larvae of C. vittatus (Lang & Young 1977) pass through 5 zoeal instars, rarely 4, whereas, C. infraspinatus (Shenoy & Sankolli 1977) and C. padavensis (Shenoy & Sankolli 1975) pass through 4 zoeal instars before a glaucothoe.

Thus, stage to stage comparison is possible with only the latter two species.

The first stage larva of Clibanarius aequa- bilis var. merguiensis differs from those of C. padavensis and C. infraspinatus in the following: the posterior border of telson

along with the notch fringed with fine hairs; exopods of first and second maxillipeds two- segmented; outer margin of endopod of second maxilla without hairs. Also, the larvae of the present species differ from other larvae in seta- tion of appendages.

The third zoea of Clibanarius aequabilis var. merguiensis differs from those of C. pada- vensis and C. infraspinatus in the telson, having an unarticulated spine as the fourth process in the present species as against a minute tubercle in C. infraspinatus and C. padavensis.

Shenoy & Sankollii (1977) consider the re- duced fourth process of C. infraspinatus as a generic character. However, in the larvae of C. vittatus (Lang & Young) and C. aequabilis var. merguiensis (present species) the fourth process changes into an unarticulated spine and retains its prominence throughout the zoeal stages. Hence the reduced fourth telson pro- cess in C. infraspinatus should not be consider- ed as a generic feature.

The glaucothoe of Clibanarius aequabilis var. merguiensis exhibits generic features sum- marised by Shenoy & Sankolli for C. infra- spinatus (1977). The differences observed are mainly in the armature of appendages, hence a detailed comparison is not made.

Ack nowledgements

I am thankful to K. N. Sankolli and Shakun- tala Shenoy, Taraporevala, Aquarium, Bombay, for checking the drawings, and to the Kama- tak University for providing laboratory faci- lities. Also, thanks are due to Dr. V. B. Nadkarni, Head of the Department of Zoo-

40

THE HERMIT CRAB CLIB AN ARIUS AEQUABILIS VAR. MERGUIENSIS

logy, Karnatak University, for his valuable by an award of Junior Fellowship from the suggestions. The research work was supported C.S.I.R., which is gratefully acknowledged.

References

Lang, W. H. & Young, A. M. (1977) : The larval development of Clibanarius vittatus (Bose) (Crusta- cea: Decapoda: Diogenidae) reared in the labora- tory. Biol. Bull. 152: 84-104.

Shency, Shakuntala & Sankolli, K. N. (1975): Metamorphosis of an estuarine hermit crab, Cliba- narius padav crisis de Man, in the laboratory (Deca- poda, Anomura). Bull. Dept. Mar. Sci. Uni. Cochin

VII (3): 671-683.

(1977):

Laboratory culture of the hermit crab Clibanarius infraspinatus Hilgendorf (Crustacea, Decapoda, Anomura). Proc. Symp. Warm Water Zooplankton: 660-670.

Thakur, M. K. (1960): A new technique for preserving prawn larvae. Curr. Sci. 29: 128.

41

TOURIST ACTIVITY AND BEHAVIOUR OF THE LEOPARD PANT HERA PARDUS FUSCA (MEYER, 1794) IN THE RUHUNA NATIONAL PARK, SRI LANKA1

M. R. Chambers2, Charles Santiapillai3

AND

N. ISHWARAN4 (With two text -figures)

The activity of the leopard ( Panthera pardus fusca) was carefully monitored in three areas of Block I of the Ruhuna National Park, Sri Lanka, from July 1979 to June 1980. In 67 hours of observations there were 16 sightings. Despite few records, the following conclusions were made: — (a) leopards had two activity peaks, early morn- ing and late evening; (b) leopards were seen more frequently in the drought months; (c) leopards were not seen with the same frequency in the different areas; (d) sight- ings were short, mostly 10 seconds or less, and (e) most sightings caused obvious disturbance to the leopards. Tourist activity in the Park mav account for these

characteristics of leopard behaviour.

Introduction

In Sri Lanka the leopard, once widespread, is now mainly limited to the country’s major National Parks, and there are probably no more than a few hundreds remaining in the country (Santiapillai et al. 1982).

The National Parks are visited by large num- bers of tourists and the wildlife authorities are concerned that these visitors may be having harmful effects such as altering the behaviour and distribution patterns of ungulates and carnivores.

In the Ruhuna National Park the leopard is confined mainly to forested areas, and direct

1 Accepted, June 1983.

2 Department of Biology, University of Papua New Guinea.

3 Department of Zoology, University of Perade- niya, Sri Lanka.

4 Department of Zoology, University of Perade- niya, Sri Lanka.

observations on the animal are not easy. Sight- ings are frequently brief and restricted to early morning and late afternoon periods of activity. The present study reports an attempt to make brief sightings of the leopard of quantitative value so that reasonable inferences may be made concerning its behaviour and abundance. In addition, good quantitative data on leopard sightings may be useful for measuring future changes in leopard behaviour and/or abun- dance.

The study area

The Ruhuna National Park is situated in the arid south-east corner of Sri Lanka (for- merly Ceylon). The annual rainfall is less than 1000 mm unequally spread throughout the year. There is a prolonged drought from June to September, a marked rainy season from Octo- ber to December and intermittent rains from January to May. The Park is 1,160 km2 and is

42

THE LEOPARD IN THE RU HUN A NATIONAL PARK, SRI LANKA

divided into several Blocks. The present study was carried out in Block 1, the main tourist area of the Park which currently attracts about 90,000 visitors each year. Most tourists enter in the early morning or late afternoon. The

vegetation of Block 1 consists principally of climax riverine forest, thorn scrub (degraded climax forest) and grasslands (edaphic climaxes maintained by seasonal floodings and by ungulate and small mammal grazing pres-

Fig. 1. Map of Block I, Ruhuna National Park, Sri Lanka showing the locations ( • ) of the leopard sightings in the main study Areas A, B & C.

43

JOURNAL, BOMBAY NATURAL HIST. SOCIETY, Vo!. 81

sure). The predominant vegetation is a dense thorny scrub through which visibility rarely exceeds 20 m and is frequently much less. The grasslands of Block 1 are found throughout the area although more commonly in the cen- ' tral and coastal regions. They are small in size, the largest being about 0.5 km2.

The Park supports a large biomass of herbi- vores, the most important being spotted deer ( Axis axis Erxleben), water buffalo ( Bubalus bubalis L.), elephant ( Elephas maximus L.), wild pig ( Sus scrofa L.), sambar ( Gervus uni- color Kerr) and the black-naped hare ( Lepus nigricollis F. Cuvier). The leopard is the only large carnivore in the Park and in Sri Lanka. For a fuller description of the Ruhuna National Park see Comanor (1971) and Mueller-Dom- bois (1968 and 1972).

Methods

All observations were made between July 1979 and June 1980. During the course of our studies on the herbivores of the Park (Bala- subramaniam et al. 1980, Santiapillai and Chambers 1980, 1981 and 1982, Santiapillai et al. 1981 and 1982) we travelled extensively in Block 1. For the purposes of studying the leopard population. Block 1 was divided into three Study Areas A, B and C (Fig. 1). Area A comprised the main road running the length of Block 1 from the entrance at Palatupanu to its termination at the Yala bungalow and all roads east of this. Area B comprised the central roads and Area C the northern roads. A fourth major road running along the western side of Block 1 was not surveyed frequently enough to be included in this analysis although leopards have been seen there. Each day was divided into 2-hour time intervals from 0600 to 1800 with an additional interval later than 1800 h. The amount of time spent travelling in

each Study Area was carefully logged and whenever a leopard was sighted the following information noted:- date, time, duration of sighting, minimum distance from the leopard and its behaviour during the sighting. Every effort was made to reduce disturbance during the sighting and each sighting was continued until the leopard disappeared from view.

Results

The total observation time within the Park was 4020 minutes (67 hours) and there were 16 leopards sighted (l/4.2h). For ease of comparison, sighting frequencies are reduced to a standard unit of number of sightings per hour of observation.

(a) Pattern of daily sightings

The total amounts of time spent on the look- out for leopards in each time period, together with the number and frequency of sightings, are given in Table 1. The data show two peaks of activity each day, a smaller one between 0600 and 0800 h (0.43 sightings /h) and a second larger one after 1800h with a frequency of sightings (0.79/h) double that of the morn-

Table 1

Number and frequency of leopard sightings in

EACH TIME INTERVAL, RUHUNA NATIONAL PARK, JULY

1979 to June 1980

Time interval

Minutes of obser- vations

Number of Frequency sightings of sight-

ings (no./h)

06. CO-08. 00

415

3

0.43

08.00-10.00

820

1

0.07

10.00-12.00

475

1

0.12

12.00-14.00

385

0

• —

14. CO-16. ©0

340

0

—

16. 00-18. GO

830

1

0.07

From 18.00

755

10

0.79

44

THE LEOPARD IN THE Rli HUN A NATIONAL PARK, SRI LANKA

ing. The leopards in Block 1 were therefore mainly active after 1800h, and most of these sightings were made after sunset (about 1830 h). There may also be considerable activity before 0600 h. The activity peaks, at least during the daylight hours, were very short indeed. Between 0800 and 1 800 h there were only three leopard sightings in 2850 minutes (0.06/h) and none between 1120 and 1755 h.

(b) Seasonal variations in sightings

The frequency of leopard sightings during the dry season, wet season and the period out- side the peak dry season are given in Table 2.

Table 2

Seasonal variation in the frequency of leopard SIGHTINGS, RUHUNA NATIONAL PARK, JULY 1979 TO

June 1980

Season

Minutes of obser- vations

Number of Frequency sightings of sight- ings (no.|h)

Peak dry season

920

7

0.46

(July /Aug) Peak wet season

815

0.22

(Nov/Dec) Period outside peak dry season

3100

9

0.18

(Nov/June)

These frequencies varied with season. Leopards were seen most frequently during the drought months of July and August (0.46/h). During the wet season (November and December) sightings were 0.22/h and for the whole of the period outside the drought. 0.18 /h. It was therefore apparent that leopard behaviour varied between dry season and the rest of the year.

(c) Frequency of sightings in the Study Areas The total observation times together with

the number and frequency of sightings in each of the three Study Areas are given in Table 3.

Table 3

Variation in the frequency of leopard sightings in the study areas of Ruhuna National Park, July 1979 to June 1980

Area

Minutes of observations

Number of sightings

Frequency of sightings (no./h)

A

2415

9

0.22

B

890

5

0.33

C

715

2

0.17

Leopards were seen most frequently in Area B (0.33/h) and least in Area C (0.17/h). It was not possible (because of the few sightings and seasonal and daily variation in sighting frequencies) to assess whether or not the diffe- rent rates from the Areas reflected chance observations or real differences in leopard be- haviour and/or abundance. A possible indica- tion of the differences between Areas A and B was the frequency of early morning sightings (0600 to 0800 h). In Area A no sightings were made during 190 minutes, whereas in Area B, 3 leopards were seen in 225 minutes. Similarly nearly all night sightings (after dusk) were in Area A. These observations tend to suggest that leopards avoided the busy roads of Area A during the daylight hours but came to them after sunset once the tourists had left the Park.

(d) Duration of sightings

The duration of each of the 16 sightings is shown diagrammatically in Fig. 2. The total duration of all 16 sightings was 896 seconds and the average 56 seconds. If the one long sighting of 600 seconds is excluded, the average duration of the remaining 15 was 20 seconds.

45

Number of Sightings

JOURNAL, BOMBAY NATURAL HIST. SOCIETY, Vol 81

12

Duration (s) of Sightings

Fig. 2. Histogram showing the duration and frequency distribution of leopard sightings in the Ruhuna National Park, Sri Lanka, from July 1979 to June 1980.

Ten of the sightings were of 10 seconds or less. These generally very short sightings were in marked contrast to the long ones noted by Eisenberg and Lockhart (1972) and Mucken- hirn and Eisenberg (1973) that averaged from 8 to 10 minutes in the Wilpattu National Park, Sri Lanka.

Of the 16 sightings there were 12 (75%) in which the leopard was obviously alarmed by us. In the remaining 4 the leopard was appa- rently unconcerned by the sometimes close encounters with the vehicle. On two occasions leopards strolled with 5 m of the jeep without apparently being aware of its presence.

46

THE LEOPARD IN THE RUHUNA NATIONAL PARK, SRI LANKA

Discussion

The leopard sightings documented here form a part of our studies on the fauna of the Ruhuna National Park. In anticipation of in- frequent sightings we attempted to collect the data in such a way that it would be possible to draw conclusions about leopard behaviour, possible disturbance by tourists and to provide baseline information for comparison with future studies. During the year we recorded only 16 sightings in 67 hours of travels along the roads and tracks of the Park. Although this was not an ideal data base from which to make interpretations, several tentative conclu- sions can be drawn.

Firstly, leopards had morning (06.00 to 08.00) and evening (after 18.00) activity peaks. Between these periods there was very little observed activity. Such a daily behavi- our pattern is typical of large carnivores. FIow- ever the daylight activity peaks were shorter than those observed by Eisenberg and Lock- hart (1972) in Wilpattu National Park, Sri Lanka. This had considerably fewer visitors than Ruhuna. It was possible therefore that daytime leopard activity in Ruhuna was cur- tailed by the early morning and late evening rush of visitors to the Park. More detailed studies would be required to show if leopards were active in the daytime but in areas away from the roads.

The results also show that leopards were seen more frequently during the July /August drought than at any other times of the year. One possible reason for this is that Ruhuna was closed to visitors at this time. Consequently very few vehicles were in the Park and the leopards were less frequently disturbed. Since the normal reaction of the leopard on seeing or hearing a vehicle was to disappear quickly into the adjacent scrub, it follows logi-

cally that the fewer vehicles in the Park the greater the chance of seeing leopards on or by the roadside. The drought season was also the leopard breeding season (Santiapillai et al 1982) and therefore the animals may have been more active and wider ranging at this time which could account for more frequent sightings.

Other possible explanations for increased frequency of sightings at this time — loss of leaves by much of the scrub vegetation and changes in prey distribution — were unlikely to account for them. This was because all sightings were in open spaces — on roads, verges or grasslands and therefore unaffected by increased visibility into the scrub. Similarly during the drought one of the leopard’s main prey species (spotted deer) moved into the scrub away from the roads (Balasubramaniam ei al. 1980). If the leopards followed them closely, reduced sightings would be expected at this time.

The average duration of leopard sightings in Ruhuna was 56 seconds and the majority were 10 seconds or less. These short observa- tion periods were undoubtedly due to the facts that virtually all sightings took place on or by roads and that the leopards moved quickly into the scrub when disturbed. Large numbers of vehicles travelling within the Park therefore virtually ensured that very few leopards would be seen.

The affinity of leopards for roads has long been documented (Storey 1907). The results of this study suggest that at Ruhuna this affi- nity was such that daily and seasonal behavi- our patterns were to some extent at least gear- ed to human activity on the roads.

The data collected during this study gave no information on the density of leopards in Ruhuna. The frequency of leopard sightings however, especially when related to time of the

47

JOURNAL, BOMBAY NATURAL HIST. SOCIETY, Vol 81

day and locality, provide baseline information that can be used to monitor any future changes in numbers. Any significant increase or de- crease in the leopard numbers would be ex- pected to show up as corresponding changes in the frequency of sightings during a similar future survey, providing the behaviour of the leopards was unchanged.

Acknowledgements

We wish to thank Professors K. D. Arud- pragasam and H. Crusz of the Universities of

Refer

Balasubramaniam, S., Santiapillai, C. & Cham- bers, M. R. (1980): Seasonal shifts in the pattern of habitat utilization by the spotted deer ( Axis axis Erxleben 1777) in the Ruhuna National Park, Sri Lanka. Spixiana 3: 157-166.

Comanor, P. L, (1971) : An analysis of the woody scrub vegetation of Ruhuna National Park, Ceylon. Tropical Ecology (India) 72: 209-222.

Eisenberg, J. F. & Lockhart, M. (1972) : An eco- logical reconnaissance of the Wilpattu National Park, Ceylon. Smithsonian Contrib. zool. 101. Smithsonian Institution Press, Washington, D.C.

Muckenhirn, N. & Eisenberg, J. F. (1973): Home ranges and predation of the Ceylon leopard (Panthera pardus fusca). In: Eatob, R. (ed.). The World Cats, volume 1. World Wildlife Safari, pp. 142-175.

Mueller-Dombois, D. (1968): Ecogeographic

analysis of a climate map of Ceylon with particular reference to vegetation. Ceylon Forester 8: 39-58.

(1972) : Crown distortion

and elephant distribution in the woody vegetation of the Ruhuna National Park, Ceylon. Ecology 53: 208-227.

Santiapillai, C. & Chambers, M. R. (1980): Aspects of the population dynamics of the wild pig

Colombo and Peradeniya respectively for their help and encouragement in our Ruhuna studies. We also wish to thank Mr. Childers Jayawar- dhana. Assistant Director of the Department of Wildlife Conservation for his help and hospi- tality during our visits. We are grateful to the National Science Council of Sri Lanka for financial support. Our thanks to Mr. Tissa Alagoda of the Department of Zoology, Uni- versity of Peradeniya for technical assistance. Finally we wish to thank Dr. Brian Mitchell of the Institute of Terrestrial Ecology, Banchory, Scotland, for helpful comments and criticisms.

ENCES

(Sus scrofa L.) in the Ruhuna National Park. Spixiana 3: 239-250.

Santiapillai, C. & Chambers, M. R. (1981): Observations on the Sambar ( Cervus unicolor Kerr 1792) in the Ruhuna National Parte, Sri Lanka, Ceylon Journal of Science 14: 193-205.

(1982):

The social organisation and calving patterns of the water buffalo (Bubal us bubalis L.) in the Ruhuna National Park, Sri Lanka. In: Workshops on water buffalo research in Sri Lanka, SAREC report R. 3. Published by the Swedish Agency for Research Co- operation with Developing Countries, Stockholm, pp. 59-67.

Santiapillai, C, Balasubramaniam, S. & Cham- bers, M. R. (1981): A preliminary study of bark damage by cervids in the Ruhuna National Park, Sri Lanka. Spixiana 4: 247-254.

Santiapillai, C., Chambers, M. R. & Ishwaran, N. (1982) : The leopard ( Panthera pardus fusca Meyer 1794) in the Ruhuna National Park, Sri Lanka and observations relevant to its conservation. Biological Conservation 23: 5-14.

Storey, H. (1907): Hunting and shooting in Cey- lon. (Reprinted 1969). Tissara Prakasakayo. Dehi- wela, Sri Lanka.

48

SOME OBSERVATIONS OF SCARCE BIRDS IN NEPAL1

N. J. Redman2, F. Lambert3 and R. Grimmett4

Sightings of three species of birds previously unrecorded in Nepal are documented. Brief details of a further eleven species of ornithological significance to Nepal are also presented.

Introduction

From December 1978 to February 1979 the authors, R. Filby, C. Murphy and L. Norton visited Nepal to observe birds. C. Murphy and N. J. Redman made a return visit from April to June 1979. During these two periods 585 species of birds were identified, three of which: Baer’s Pochard Aythya baeri , Sanderling Calidris alba and Common Gull Larus canus, were recorded for the first time in Nepal. The main purpose of this paper is to document these sightings. Brief details are also given of other species seen for which there are very few pre- vious Nepalese records or for which informa- tion on breeding or unusual behaviour was gathered.

A wide variety of habitats and altitude zones in central and eastern Nepal was visited. Areas of significant ornithological interest included the Jomosom trek (January), Helambu/Gosain- kund trek (May), Kathmandu Valley, central tarai, Ilam district and Kosi Barrage. The latter proved to be of particular interest since many scarce species were found here, including all three new species for Nepal. We visited Kosi

1 Accepted March 1983.

2 I Westfields, Saffron Walden, Essex.

" 15 Bramble Rise, Westdene, Brighton, Sussex.

4 12 Hobbs Way. Rustington, Littlehampton, W. Sussex.

Barrage on four separate occasions, 10-13 February, 20-21 February, 17-20 April and briefly on 23-24 April. During these four short periods totalling eight full days, the majority of our time was spent in areas on the north side of the barrage.

New Species for Nepal baer’s pochard Aythya baeri (Radde)

Baer’s Pochard was first identified on 12 February 1979 when two males and one female were seen on open water just north of the Kosi Barrage amongst a flock of several hundred Aythya spp. On 20 February we saw this species again and counted at least 17 males and 3 females. Prolonged observation of both sexes was possible and comparisons were made with accompanying Tufted Duck A. fuligula , Common Pochard A. ferina and Ferruginous Duck (White-eyed Pochard) A. nyroca.

In shape Baer's Pochard was comparable to Ferruginous Duck and Common Pochard, and intermediate in size.

Males were readily identifiable by the com- bination of iridescent green head and rich chestnut breast, unlike any other Aythya species. Other features included white belly, dusky brown flanks, white under-tail coverts and gleaming white eye. Females were similar ir coloration to the males but duller, the

49

4

JOURNAL . BOMBAY NATURAL HIST. SOCIETY, Vol. 81

dusky head showing only a slight greenish gloss. At a distance, female Baer’s Pochard appeared confusingly similar to other female Ay thy a species. Common Pochard and Tufted Duck, however, both lacked white undertail coverts and Ferruginous Duck could generally be separated by its uniform chestnut coloura- tion, less white on the belly and flanks, and smaller size.

Baer’s Pochard breeds in eastern Siberia wintering primarily in eastern China. In the Indian sub-continent, Ripley (1982) states that it is an uncommon and erratic winter visitor to Manipur, Assam and associated states, Bangladesh and Bengal; also recorded from Bihar and Maharashtra. Ali and Ripley (1978) suggest that it may be less rare than records indicate. There are no previous records from Nepal and this species was not seen at Kosi in the subsequent three winters (C. and T. Tnskipp in lift.).

sanderling Calidris alba (Pallas)

A single Sanderling was discovered at Kosi Barrage on 11 February 1979 feeding on a mudbank with nine Dunlin Calidris alpina and fifty Little Stints C. minuta. It was not seen on subsequent days.

In size it was considerably larger than Little Stint and about the same as Dunlin but with a shorter, straight bill. Bill and legs were black. The very pale plumage, pale grey above and pure white below, served to distinguish the bird at some distance. A small black shoulder patch and black primary tips were also noted, and in flight it showed a more prominent white wing bar than Dunlin or Little Stint. A distinctive feature was its rapid feeding action. All the observers are familiar with this species in Europe.

Sanderling breeds in the Arctic, wintering on coasts almost throughout the world.

Although a regular winter visitor to the coasts of India, it has apparently not been previously recorded inland in the subcontinent (Ali and Ripley 1980). Our sighting constitutes the first record from Nepal.

common gull Lotus canus Linnaeus

On 12 February 1979 a Common Gull was found amongst a group of Black-headed Gulls L. ridibundus resting on a sandbank at Kosi Barrage. It remained in the area all day and allowed direct comparison with the Black- headed Gulls and also with Brown-headed Gulls L. brunnicephalus and a single adult Slender-billed Gull L. genei. It was still pre- sent on 21 February. Despite its rarity in Nepal, Common Gull is a familiar bird in Europe and was readily identified by all the observers as a first-year bird.

In appearance it was an elegant, medium- sized gull with a rounded head and a small yellowish bill with a dark tip. The head, neck and underparts were white with some faint brownish markings on the sides of the breast and head. A pale whitish oval patch on the closed wing contrasted with the grey mantle. The protruding primaries were dark brown and the legs were pale yellowish. In flight it showed a broad whitish band across the inner wing, contrasting with the brown flight feathers. The tail was white with a broad dark terminal band.

The Common Gull was easily separated from all other gulls present by its size, shape and colour of bare parts. Black-headed Gull was noticeably smaller and slimmer with red- dish bill and legs. Brown-headed and Slender- billed Gulls, although of similar size, were different in shape and also had reddish bills and legs. The only species of gull likely to cause confusion within the Indian subcontinent is the Herring Gull L. argentatus, but this species is considerably larger and deeper-

50

SOME OBSERVATIONS OF SCARCE BIRDS IN NEPAL

chested with a more angled head and much stouter bill. Its wings are broader and flight heavier.

Common Gull is a widely distributed and abundant holarctic species breeding across the northern palearctic and moving south in winter. Surprisingly it has rarely been recorded from the Indian subcontinent. It is not included in Ali and Ripley (1981) or Ripley (1982) and the only records from the Indian subcontinent of which we are aware are two individuals from the Punjab area of Pakistan: L. J. Djiksen, F. Koning and A. Vittery saw an adult at Rasul Barrage on 27 January 1974 and A. Vittery identified a first winter bird at Rawal Lake, near Islamabad on 17 February 1974 (A. Vittery, pers. comm.). Our sighting represents the first record from Nepal. Subse- quently, an adult in winter plumage has been observed at Phewa Tal near Pokhara on 21 January 1981 (del-Nevo and Ewins 1981).

Other records of outstanding interest fulvous whistling duck (Large Whistling Teal) Dendrocygna bicolor

A single individual of this species was iden- tified amongst a flock of 1500 Lesser Whistling Duck (Teal) D. javanica at Kosi Barrage on 12 February 1979. This record constitutes the only sighting of Fulvous Whistling Duck in Nepal this century. Although overlooked by recent authors, a Hodgson specimen from Nepal is listed by Sharpe (1894). The speci- men is still present in the British Museum and was located by C. Inskipp in 1981. In the Indian subcontinent this species is scattered widely, but sporadically in many areas and everywhere rather scarce. It is apparently more common in Bengal and Bangladesh (Ali and Ripley 1978).

ruddy shelduck Tadorna ferruginea

A pair with 8 newly-hatched young was

present on a lake at Gosainkund on 27 May 1979 at an altitude of approximately 4300 m. This represents the first positive breeding re- cord for Nepal (R. L. Fleming Jr., pers. comm. ) .

white-tailed (sea) eagle Haliaeetus albicilla Seven birds were seen from December to February including three at Kosi Barrage. An adult at Begnas Tal near Pokhara on 2 Janu- ary 1979 was seen to catch a Purple Gallinule (Purple Moorhen) Porphyrio porphyrio in flight. The gallinule was subsequently dropped and defied repeated attempts at recapture by diving. Attacks on flying birds are apparently rare (Cramp et al. 1980).

black-tailed godwit Limosa limosa A flock of 29 was seen at Kosi Barrage on 18 April 1979. In the 19th century Hodgson obtained at least one specimen in April and five between mid- August and late October (no year given), from the Kathmandu Valley. He wrote: “small flocks feed by day on chours or moist cultivated plots”. J. Scully obtained a specimen on 7 September 1876 or 1877 in the Kathmandu Valley (Scully 1879), stating that “the godwit is a winter visitant to the Nepal Valley, but does not appear to be common there”. In this century, a single bird was obser- ved in the Kathmandu Valley in August 1978 (Fleming et al. 1979) and a specimen was taken at Chobar on 30 August 1981 by H. S. Nepali. There have been at least five other re- cords at Kosi Barrage in Spring since 1979 (C. and T. Inskipp in lift. 1983).

little owl Athene noctua A single Little Owl was found at Kagbeni in the upper Kali Gandaki Valley on 14-15 January 1979 at an altitude of approximately 2800 m. This species was first recorded in Nepal

51

JOURNAL, BOMBAY NATURAL HIST. SOCIETY, Vol. 81

in July 1978 when two specimens were collect- ed in Dolpo (H. S. Nepali pers. comm.). It has been seen subsequently at Kagbeni and nearby at Muktinath by several observers (C. and T. Inskipp in litt. 1983).

Indian cliff swallow Hirundo jluvicola Up to ten birds were present at Begnas Tal near Pokhara on 3 January 1979. This consti- tutes the second record from Nepal, the first being a single bird at Kosi on 10 April 1975 (Fleming et al. 1979).. There have been at least five subsequent records (C. and T. Ins- skipp in litt. 1983).

rufous-bellied robin Tarsi ger ( Erithacus ) hyperythrus

A pair was feeding young on 24-25 May 1979 at about 3500 m on the west side of the Gandak-Kosi watershed. The nest was not seen but was sited on or close to the ground at the side of a shady wooded ravine. A second female was found nearby. Fleming et al. (1979) state that the species is known from the Kosi- Gandaki watershed ridge eastward but there have subsequently been several records west to the Kali Gandaki (C. and T. Inskipp in litt. 1983). The nest is undescribed (R. L. Flem- ing, Jr., pers. comm.).

EYE-BROWED THRUSH (DARK THRUSH)

T urdus obscurus

Two birds were identified at Kokarna in the Kathmandu Valley on 20 December 1978 amongst a flock of thrushes comprising five species. Excluding a Hodgson specimen of unknown origin this was the first record of this species for the Kathmandu Valley. All other records are from East Nepal (C. and T. Inskipp., pers. comm.).

dusky thrush T urdus naumani

An adult was found at Lete in the Kali

Gandaki Valley on 22-24 January 1979 in the company of a flock of sixty Dark-throated Thrushes T. ruficoilis. A second bird, duller in plumage and presumed to be in first winter plumage, was discovered on 4 February 1979 at Gokarna in the Kathmandu Valley, also amongst a flock of Dark-throated Thrushes. In the 19th Century at least two specimens were obtained by Flodgson in the Kathmandu Valley in January (Gray and Gray 1846, Seebohm 1881). The only subsequent reference to Dusky Thrush in Nepal is a report of large parties on Nagar Jong at about 1500 m in Spring 1948 and a few in the Kathmandu Valley the following winter (Proud 1949). There have been several individuals reported since our observations, in both the Kali Gandaki and Kathmandu Valleys (C. and T. Inskipp in litt. 1983). In the Indian subcontinent, Dusky Thrush is stated to be an irregular winter visitor, rare in the west and more frequent in the east (Ripley 1982).

yellow-browed tit Sylviparus modestus

An occupied nest hole of a pair of Yellow- browed Tits was found on 2 May 1979 on the slopes of Phulchowki in the Kathmandu Valley, at an altitude of 2000 m. The pair was feeding young. This was only the third nest ever discovered of this species. Full details have already been published (Lohrl 1981).

SCARLET-BACKED FLOWERPECKER Dicaeum

cruentatum

A male and two females were seen at Dharan in East Nepal on 21 April 1979. This is only the second record of this species in Nepal (Fleming et al. 1979).

Acknowledgements

We are indebted to Carol and Tim Inskipp for providing valuable information on the

52

SOME OBSERVATIONS OF SCARCE BIRDS IN NEPAL

status and distribution of birds in Nepal and for commenting on an earlier draft of this paper. Thanks are also due to Bob Fleming

Refer

Ali, S. & Ripley, S. D. (1978): Handbook of the Birds of India and Pakistan. Vol. 1 (2nd edition).

— (1980) : Handbook of the

Birds of India and Pakistan. Vol. 2 (2nd edition). Bombay.

(1982): Handbook of the

Birds of India and Pakistan. Vol. 3 (2nd edition). Bombay.

Cramp, S. et a/. (1980) : Birds of the Western Palearctic. Vol. 2. Oxford.

Del-Nevo, A. & Ewins, P. (1981): Bird Watch- ing in Nepal, 7th December 1980 — 19th February 1981. Unpublished.

Fleming, R. L. et al. (1979) : Birds of Nepal, (2nd edition). Kathmandu.

Gray, J. E. & Gray, G. R. (1846) : Catalogue of the specimens and drawings of Mammalia and

Sr. and Bob and Linda Fleming for much assistance and encouragement during our stay in Nepal.

ENCES

birds of Nepal and Tibet, presented by B. H. Hodg- son, Esq to the British Museum, London.

Hodgson. B. H. (undated). Original paintings held in the Zoological Society of London Library. Unpublished.

Lohrl, H. (1981) : “Zur Kenntnis der Laubmeise, Sylviparus modestus”. J. Orn. 122 : 89-92.

Proud, D. (1949): Some Notes on the Birds of the Nepal Valley. J. Bombay nat. Hist. Soc. 48: 695-719.

Ripley, S. D. (1982): A Synopsis of the Birds of India and Pakistan. 2nd edition. Bombay.

Scully, J. (1879) : Contribution to the Ornitho- logy of Nepal. Stray Feathers 8: 204-368.

Seebohm, H. (1881) : Catalogue of the Birds in the Collection of the British Museum. Vol. 5.

Sharpe, R. B. (1894): Catalogue of the Birds in the Collection of the British Museum. Vol. 23.

53

SPAWNING OF SOME IMPORTANT COLDWATER FISH OF THE GARHWAL HIMALAYA1

S. P. Badola2 and H. R. Singh3 {With a plate )

The present study gives an account of the spawning of some important coldwater fish of the Garhwal Himalaya. Most of them have one breeding season and breed in summer, monsoon and postmonsoon months. However, Schizothorax species show a long spawning season (July to January) and their breeding is at a peak from Sept- ember to November. Puntius chilinoides and P. hexastichus spawn twice a year during two different but relatively short spawning periods, May-July and December-January. Increased pH and flooding (turbidity) is necessary for the spawning of Noemaeheilus, Glyptothorax, Pseudecheneis and Tor species of the Garhwal streams. High concen- tration of dissolved oxygen and relatively low pH are necessary for the spawning of Schizothorax, Labeo and Barilius. Besides flooding and varying values of pH and oxygen content of the waters, a varying suitable temperature is also necessary for the spawning of these species. The natural breeding grounds of these fishes are also disturbed by the transportation of timber in the Garhwal rivers.

There is little or no information available on the breeding habits of coldwater fishes of the Garhwal Himalayas. Hence this study was undertaken.

Material and Methods

The period of spawning as inferred by the presence of mature ova and testes in the body cavity was confirmed by actually finding the spawn, fry and fingerlings. In some cases the ova attached to stones, lying in a particular breeding place were collected and counted (Plate 1). The breeding site of a particular fish was decided by the occurrence of its eggs and fry in that place. The temperature, pH, depth, gradient, current, and dissolved oxygen

1 Accepted December 1980.

2 Department of Zoology, Government Postgra- duate College, Rishikesh, Dehra Dun.

3 Department of Zoology, Garhwal University, Srinagar Garhwal. (U.P.).

of the water in the breeding ground were re- corded. The specimens were collected and exa- mined from different snow-fed and non-snow- fed rivers and streams.

This study was made in respect of the fol- lowing fishes:

Schizothorax sinuatus (Heckel), S. plagio- stomus (Heckel), S. richardsonii (Gray), Tor tor (Ham.), T. putitora (Ham.), Labeo dyo- cheilus (McClell.), L. dero (Ham.), Barilius bendelisis (Ham.), B. vagra (Ham.), B. barna (Ham.), Puntius chilinoides (McClell.), P. hexastichus (McClell.), Noemaeheilus monta- nus (McClell.) M. multifasciatus (Day), N. rupicola (McClell). Glyptothorax pectinopterus (McClell.), and Pseudecheneis sulcatus, (McClell.).

1. Schizothorax sinuatus, S. plagiostomus and S« richardsonii. These species are most common in snow-fed rivers and streams of this region. It was found that they start spawn- ing gradually from July and end in January.

54

J. Bombay nat. Hist. Soc. 81

Singh: Spawning of coldwater fish

Plate

Above : Showing fish ova attached to stones.

Below : Showing destruction of ova in the breeding ground by timber logs

X

SF AWNING OF SOME IMPORTANT COLDWATER FISH

The peak spawning period was observed in September-October and November. In these months the water velocity, temperature, and dissolved oxygen remain quite favourable for their breeding. They breed in shallow running semistagnant water along the banks of the rivers among gravel and stone. The eggs re- main attached to stones in batches. The water temperature of the breeding ground ranged between 7.8 and 15.6°C, the atmospheric temperature was 12.9° to 31.2°C, pH of water was between 7.0 and 8.2 dissolved oxygen 9.0 to 18.1 ppm. The depth of breeding ground varied from 25 to 32 cm. The gradient was less and velocity was observed to be 0.496 metre /second.

2. Tor tor and T. putitora: These species spawn from April to July when the water of Alaknanda becomes turbid due to the melting of snow at the peaks. They move from the deeper waters or lower regions upward for breeding. This type of local movement was noticed right from April, when the water starts becoming turbid. It is believed that mature specimens travel upstream from Rishikesh or Hardwar, first to the upper reaches of the Ganga and then to Alaknanda and Bhagirathi for breeding. The eggs are laid on and under stones at a depth of 35 to 50 cm where the gradient is less. The water temperature of the breeding place was from 15° to 17.5°C, atmospheric temperature 26.1° and 31.2°C, pH of the water 7.0 to 8.2 and dissolved oxy- gen 9.0 to 9.9 ppm. The fertilized eggs were slightly brown in colour and found attached to the stones and rocks and other objects such as logs, etc. However, we could not get adult Tor species in Alaknanda after July- August when only large number of fry and fingerlings were found in the backwaters and sidestreams of the Alaknanda and Bhagirathi rivers. It appears that after spawning the adults move

down the Ganga probably due to the effect of the low water temperature. Thus we could observe the breeding only from April to July.

3. Lafoeo dyocheilus and L. dero : These two species abound in Alaknanda, Bhagirathi and Finder, etc. from March to June when they come from Ganga for the purpose of breeding. During these months, they are found with mature gonads. The eggs are laid on and among the stones towards the bank of the river at a depth of 30 to 35 cm in slow running water. The water temperature in the breeding grounds was 12.6° to 17.5°C, atmospheric temperature 26.8° (in April) to 31.2°C (in June), maximum pH was 8.2 and dissolved oxygen from 9.3 to 9.9 ppm. The velocity of water was 0.616 to 1.234 metre /sec. Thus, they prefer clear shallow water for breeding. The fertilized eggs were somewhat greenish in colour.

4. Bardins bendelisis9 B. vagra and B. barna : These species always prefer small rivers and streams, where the water is clear and shallow with high percentage of dissolved oxygen. They do not survive in polluted water. In this investigation it was found that these species breed from April to June. The breed- ing grounds were observed in Nayar, Khoh and in the side stagnant waters of Alaknanda. These small fishes are found schooling in abundance and lay their eggs in shallow pockets of water under stones and weeds (algae), as well as in the sand mixed gravel bed at a depth of 15 to 28 cm. The water tem- perature of the breeding grounds ranged from 20.5° to 22.5°C with the pH from 7.0 to 7.3. The dissolved oxygen was 9.5 to 10.2 ppm. The water was semistagnant, with low velocity (0.197 to 0.204 metre/second) and less gra- dient.

5. Puntius chilinoides and P. hexastichus :

The two species are very common in Nayar,

55

JOURNAL, BOMBAY NATURAL HIST. SOCIETY, Vol. 81

Mandakini and Pindar rivers and their breed- ing period was noticed to be from May to July and December to January. However, the actual breeding grounds of these species could not be located. But from the fry it was esti- mated that they breed in shallow water under stones and rocks, with the water temperature at 8.9°C, pH of water 7.0 and dissolved oxygen 16.8 ppm. The gradient and velocity of water was also low.

6. Noemacheilus momtanus, N. rupicola and N. multifasciatus : These species spawn from July to August in small streams and rivulets. Being small in size, their breeding ground could not be located. But it is almost definite that they breed in small streams and rivulets and not in large snow-fed rivers like Alaknanda, Bhagirathi, etc.

7. Glyptothorax pectinopterus : This species breeds from April to August and is quite com- mon in streams throughout the year. In Alak- nanda, Bhagirathi and Pinder, etc. it occurs in abundance in the rainy season. Probably having been swept in from small streams and rivulets by the swift current. Its breeding niches could not be observed due to the increased water level and velocity (2.493 metres | second).

8. Pseudecheneis sulcatus : P. sulcatus is found in snow-fed streams all the year round, but in Alaknanda, Pindar, Bhagirathi and Jam una it is available after April, when the water becomes turbid. Its spawning period was observed to be from April to August. The mature testes are branched. In female the abdomen is bulged out by large number of eggs. However, its breeding ground could not be located due to the high speed of water in the rainy season.

Destruction of eggs

During the course of this study it was found that the breeding grounds in Alaknanda and

Nayar are disturbed by the floating of timber logs (Plate 1). In winter and summer the log- ging is most common in Alaknanda and Nayar. In this period most of the fishes are in spawn- ing stage, for instance Schizothorax species breed from September to January and the Puntius species from December to January. The Tor and Labeo species breed from April to June-July. Wc have noticed that the logs that float in the main current of the river sometimes reach the banks of the river and strike the stones and rocks, and thus crush thousands of eggs and disturb the natural breeding grounds. It was estimated that about 26% eggs are damaged by this type of trans- portation of the timber. (Table 1).

Table 1

Destruction of fish eggs by timber in Alaknanda River

Spot Nature No. of sub- stratum

Total

eggs

examined

No. of

living

eggs

No. of damaged eggs

Percen- tage of damaged eggs.

1.

Stony

347

235

112

32.27

2.

Stony

286

160

126

44.05

3.

Stony &

rocky

360

292

68

18.88

4.

Stony

198

178

20

10, 10

5.

Stony

307

307

nil

nil

6.

Stony &

Rocky

401

401

nil

nil

Discussion

According to this study most of the fishes of the Garhwal Himalaya breed in the summer and monsoon months. However, only Puntius chilinoides and P. hexastichus breed twice a year, i.e., from May to July and December to January. Of all the species included in this study, Schizothorax spp. have the longest

56

SPAWNING OF SOME IMPORTANT CO LOW A TER FISH

breeding period ranging from July to January. Their breeding is at peak during September to November, the eolder days of the year, when the water has ample amount of dissolved oxygen (about 18.1 ppm). According to Jhin- gran (1975), S. richardsonii of Himachal Pradesh spawns from March to June. Bhat- nagar (1964) in his studies on Bhakra reser- voir fishes has reported that Schizothorax plagioslomus breeds in July-August and Dec- ember- January. However, our observations are that the Schizothorax species of the Garhwal waters do not spawn intermittently. Probably one reason for the availability of the Schizo- thorax in the snow-fed rivers and the streams throughout the year is that the fish has a long and continuous breeding period. The other reasons are the favourable water spread and a large amount of dissolved oxygen.

Bhatnagar (1964) has reported that the T. putitora of the Bhakra reservoir first spawns in July and this activity continues intermit- tently throughout the year. According to Karamchandani et al. (1967), T. tor of Nar- bada river has a prolonged breeding season which commences in July- August and conti- nues upto December with peak breeding from July to September. Some other studies on the breeding of the Tor species of the other re- gions arc of Khan (1939), Qasim & Qayyum (1962), and Sehgal et al. (1971), and accord- ing to them Tor species spawn two to three or more times a year. But our studies show that the Tor tor and T. putitora of the Garh- wal hills spawn once a year, i.e. from April to July. These species come to the Alaknanda and Bhagirathi from the Ganga for breeding, ft appears that the presence of suitable isolated

and well protected breeding grounds, the high amount of dissolved oxygen and the abundance of insect larvae in Bhasirathi and Alaknanda waters, which form the food of these species are some of the factors that attract Tor for this breeding migration.

Bhatnagar (1964) described the spawning of Labeo dero in July and according to him flooding was necessary for its breeding. How- ever, our observations indicate that L. dero and L. dyocheilus of Garhwal rivers prefer shallow water for breeding. Similarly Barilius bendelisis, Barilius barna and Barilius vagra were also found to breed in shallow pockets of clear waters. According to Khanna (1958) and David et al. (1967), the presence of flood water in the breeding ground and a current of moderate intensity were essential for breed- ing and increased pH did not seem to be necessary for fish breeding. Mookerjee (1945) pointed out that change of pH may be one of the principal factors which may induce carp to spawn. Das & Das Gupta (1945) stated that although an increased pH and high oxygen content of water play an important role in the spawning of carps, they have no independent position. However, our observations indicate that high pH and flood water were necessary for the breeding of Glyptothorax pectinopte- rus, Pseudecheneis sulcatus and Noemacheilus species of this region. But high oxygen con- tent. relatively low pH values and mild velo- city of the water were essential for the breed- ing of Schizothorax, Labeo, Puntius and Bari- lius species of the Garhwal streams. On the basis of this study it may be pointed out that a suitable temperature is also necessary for the breeding of the different species.

57

JOURNAL , BOMBAY NATURAL HIST. SOCIETY, Vol. 81

References

Bhatnagar, G. K. (1964) : Observations on the spawning frequency of certain Bhakra reservoir fishes. Indian J. Fish. 1 : 285-502.

Das, K. N. & Das Gupta, B. N. (1945) : Breeding of the Principal carps in Bengal. Proc. Nat. Inst. Sci. India 11: 324-327.

David, A., Govind, B. V., Rao, N. G. S. & Raj- gopal, K. V. (1967) : Fish seed resources of some rivers in South India. Indian J. Fish. 74(1-2) : 54-84.

Jhingran, V. G. (1975): Fish and Fisheries of India. Flindustan Publishing Corporation (India), Delhi.

Karamchandani, S. J., Desai, V. R., Pisolkar, M. D. & Bhatnagar, G. K. (1967): Biological in- vestigations on the fish and fisheries of Narbada river (1958-66). Bull. Cent. Ini. Fish. Res. Inst. Bar-

rack pore (10) : 40p.

Khanna, D. V. (1958): Observations on the spawning of the Major carps at a Fish Farm in Punjab. Indian J. Fish. 5(2) : 282-290.

Mookerjee, H. K. (1945) : Factors influencing the spawning of principal carps in India. Symposium on “The factors influencing the spawning of Indian carps*’. Proc. Nat. Inst. Sci. India. 11: 312-315.

Qasim, S. Z. & Qayyum, A. (1962): Spawning frequencies and breeding seasons of some fresh water fishes with special reference to those occurring in the Plains of Northern India. Indian J. Fish. S(l) : 24-43.

Sehgal, K. L., Shah, K. L. & Shukla, J. P. (1971) : Studies on some aspects of cold water fisheries in Himachal Pradesh and Kashmir (1966-67) (M.S.).

58

REPRODUCTION BIOLOGY OF THE SOFT-FURRED FIELD RAT, R ATT US MELT AD A PALLIDIOR (RYLEY, 1914) IN THE RAJASTHAN DESERT1

B. D. Rana2 and Ishwar Prakash3 (With three text-figures )

Introduction

The soft-furred field rat, Rattus meltada is distributed throughout India. It is found abun- dantly in the crop fields of the south-eastern region of the desert (Rana and Prakash 1980). It usually inhabits irrigated crop fields but is also found in grasslands on heavier soils. This species is one of the economically important among crop inhabitants and inflicts losses to standing crops and grain stores of cereals, chiefly wheat, jowar ( Sorghum vulgare) and bajra (Pennisetum typhoides). Thus, keeping in view, its economic importance, an attempt has been made to study its ecological rela- tionships in the desert environment (Rana 1981) and in this paper its reproduction bio- logy is described.

The Study Area

Bisalpur (25°7'N, 73°10'E), is situated in western Rajasthan, on the south-eastern fringe of the Thar desert, very near to Aravalli ranges. The mean maximum and minimum temperatures are observed to the order of 32.8°C and 20.0°C respectively. The average amount of precipitation received is about 500 mm but 90 per cent falls during Julv-

1 Accepted October 1981.

2 Rodent Biologist.

8 Professor of Eminence, Central Arid Zone Re- search Institute. Jodhpur- 342 003 (Rajasthan), India.

September period. The vegetation is rich due to the water retaining capacity of sandy loam. Rodents were trapped in a protected grassland and from surrounding crop fields. Two crops are raised in the region: rainfed, July to Octo- ber and irrigated, November to March. There- fore, green food is available to rodents for the greater part of the year, except from April to June.

Material and Methods

The Soft-furred field rat, or the metad, Rattus meltada pallidior (Ryley, 1914) were collected every month at Bilaspur from Janu- ary 1978 to December 1979. Soon after their collection, they were weighed, sexed and dis- sected. Ovaries were checked for the number of corpora lutea and uterine horns for im- planted embryos in freshly killed material. The teats in females were also examined for any sign of lactation.

Epididymal smears were examined for the presence of sperms and uterine horns for an indication of embryonic mortality. The testes and ovaries were weighed to the nearest 0.001 g on a semi-micro Mettler balance.

Results

MALE FECUNDITY Epididymal smear

Sexual maturity of male metads was attain- ed at 45 g body weight during both the years

59

BODY WEIGHTS ( q )

JOURNAL , BOMBAY NATURAL HIST. SOCIETY, Vol. HI

* MALE WITH SPERMS v MALE WITHOUT SPERMS

o FEMALE WITH GORPQRALUTEA x FEMALE WITHOUT CORPORALUTE A

»©@l

90<

to*

T- ■ T T ■ . ■ ^B3ua"B»gf

J FMAM J JASOWDJFMAMJASONO • ------ — 197© — — » i~ — — • 1979 —

Fig. 1. Gross body weights of R. m. pallidior during 1978 and 1979.

of study (Fig. 1). However, a few metads weighing 70 to 73 g during 1978 and 1979 res- pectively did not exhibit sperms in their epidi- dymis. It may possibly be due to regression of the testes. The regression of the testes dur- ing a part of the year has also been observed

in many rodents (Asdell 1946). Prasad (1961) and Jain (1970) also found regressed testes in T. i. cuvieri and T. i. indica respectively. The present study points out that in male R. m. pallidior the reproductive activity does not cease completely during winter as pregnant

60

MEAN PAIRED TESTES WEIGHTS MEAN MONTHLY RAINFALL (mm)

REPRODUCTION BIOLOGY OF THE SOFT-FURRED FIELD RAT

Q

Z

450* 400- 350 300 250 200 1 50-* 100 50J

• PERCENT FEMALE PREGNANT

* PERCENT MALE FECUND MEAN MONTHLY RAINFALL

1978*

1979

Fig. 2. The breeding intensity of R. m. pallidior in relation to the mean monthly rainfall.

PERCENT MALE FECUND — k MEAN PAIRED TESTES WEIGHT

00i

80

•60

)0-

30*

50-

*0*

o

z

Z)

o

yj

u.

LU

o

CE

U

a.

Fig. 3. The male fecundity rate and mean paired testes weights of R. m. pallidior.

61

;

JOURNAL, BOMBAY NATURAL HIST. SOCIETY, Vol. 81

The weights of right and left testes were, there- fore, pooled. The paired testes weight, differed significantly (F = 13.38, P < 0.05) between months. They were found to be maximum during June to October and lowest during Dec- ember. During 1979, two peaks in paired testes weight, one during March and the other in the months of August to October was observed. However, the lowest testes weights were re- corded during January (Fig. 3). The mean monthly paired testes weights run parallel to the per cent fecund males during both the years of the study.

FEMALE FERTILITY Ovary weights

The mean paired ovary weights of adult metads exhibits two main peaks, one during March to May and the second during June to October period during 1978 (Table 2). Lowest weights were observed in December, soon after attaining their peak level in Septem- ber, which may be due to ovarian refractoriness during winter. Similar trend in the variations of ovary weights was found in 1979 (Table 3).

Table 1

Monthly testes weights (mean + se) of R. m. pallidior

Months

Right

1978 Weight of testes (g)

Left

1979

Right

Left

Jan.

0.3225+0.10

0.3394+0.10

0.2402 +0.04

0.2453+0.05

Feb.

0.3160+0.18

0.2925 +0. 14

0.4182+0.11

0.3695 + 0.35

Mar.

0.5210+0.00

0.6630+0.00

0.5520+0.04

0.5452+0.04

April

0.3306+0.05

0.3327+0.04

0.3623 + 0.08

0.3441+0.08

May

0.4320+0.23

0.4145+0.19

0.4036+0.22

0.4156+0.20

June

0.6700 + 0.19

0.6620+ 0.19

0.5135+0.06

0.4816+0.03

July

0.6543+0.06

0.6269 +0.08

Collection was

not made

Aug.

0.8685+0.35

0.9050+0.30

0.5741+0.04

0.5754+0.01

Sept.

0.6590+0.00

0.5930+0.00

0.4574+0.25

0.6787+0.30

Oct.

0.3275+0.06

0.3050+0.04

0.8725+0.11

0.8212+0.08

Nov.

0.4710+0.08

0.4150+0.01

0.3914+0.05

0.3962+0.05

Dec.

0.2211+0.08

0.3129+0.09

0.4079 + 0.65

0.3826+0.59

metads were found throughout the two years (Fig. 2).

The male metads were found to be fecund throughout the two years. But the maximum rate of fecundity (100 per cent) was observed from March to October, 1978. Thereafter, it reduced to 50 per cent in the month of Dec- ember. Surprisingly, a minor peak in January was also observed. During 1979, two peaks of fecundity rate, one during spring and the second in monsoon seasons, were recorded.

A comparison of fecundity rate between the two years revealed that it remained at higher rate during 1978 (50 to 100 per cent) as com- pared to that of 1979 (35 to 100 per cent). The proportion of fecund males during both the years was also found to be closely similar to that of pregnancy rate in the population (Fig.

3).

Testes weights

Mean monthly weights of right and left testes did not differ significantly from each other in any of the months during the two years except during March 1978, when the right testes was found to be lighter than the left (Table 1).

62

REPRODUCTION BIOLOGY OF THE SOFT-FURRED FIELD RAT

Table 2

Prevalence of pregnancy and ovary weights of R . m. pallidior during 1978

Months

Total

number of females collected

Number of adult female metads

Prevalence

of

pregnancy

Paired ovary weights (Mean±SE)

Not

pregnant

Pregnant

Pregnant & lactating

January

4

2

2

0.50

0.0135± .005

February

3

1

2

-

0.66

0.0150± .007

March

2

-

1

1

1.00

0.0220± —

April

2

-

2

-

1.00

0.0213± .006

May

2

1

1

-

0.50

0.0215±0.007

June

5

2

3

-

0.60

0.0310± .001

July

5

-

5

-

1.00

0.0452± .005

August

2

-

2

-

1.00

0.0441± .004

September

2

-

2

-

1.00

0.0463± .001

October

2

-

1

1

1.00

0 . 0267 ± .002

November

7

5

1

1

0.28

0.0172± .002

December

7

4

3

—

0.42

0.0101± .004

Prevalence of

Table 3

PREGNANCY AND OVARY WEIGHTS OF R. m.

pallidior during

1979

Months

Total

number of females collected

Number of adult female metads

Prevalence

of

pregnancy

Paired ovary weights (Mean±SE)

Not

pregnant

Pregnant

Pregnant & lactating

January

8

7

1

_

0.12

0.0173± —

February

8

5

3

-

0.37

0.0273±0.002

March

10

2

7

1

0.80

0.0377±0.002

April

4

2

1

1

0.50

0.0229±0.002

May

4

1

3

-

0.75

0.0276±0.001

June

4

2

2

-

0.50

0.0176±0.001

August

4

-

4

-

1.00

0.0596± 0.006

September

3

-

3

-

1.00

0.0663±0.005

October

6

2

3

1

0.66

0.0263±0.001

November

13

8

4

1

0.38

0.0221 ±0.001

December

24

19

5

-

0.20

0.0200±0.003

Production of ova

Sixty-eight pregnant females exhibited 282 corporalutea in right ovary and 238 in left ovary, which are significantly different (x2(i) =

3.60, P < 0.05) from each other, ova pro- duction being more in the right ovaries. During 1978, the right ovary (110) possessed signifi- cantly (x2G)=7.40, p < 0.01) larger num-

63

JOURNAL, BOMBAY NATURAL HIST. SOCIETY, Vol. 81

ber of corporalutea than the left (73, Table 4). However, during 1979, the difference was not significant (Rt. 172: Lt. 165, x2(i)=0.14; Table 5).

The production of ova per pregnant female varied from 5.50 to 10.00 (av. 6.53) during 1978 and 5.00 to 12.00 (av. 8.25) during 1979. The average number of ova produced

Table 4

Pre-implantation losses in R. m. palliclior during 1978

Number of embryos Number of corpora lutea Pre-implanation losses

Mths. — — — — —

Right

Left

Total

Right

Left

Total

Mean±SE

Right

Left

Total

%

Jan.

5

3

8

6

5

1 1

5.50±0.50

1

2

3

27.2

Feb.

5

4

9

6

6

12

6.00+0.00

1

2

3

25.0

Mar.

7

6

13

8

7

15

7.50+0.50

1

1

2

13.3

Apr.

4

'j

J

7

6

5

11

5.50+0.50

2

2

4

36.3

May

4

3

7

6

4

10

10.00±0.00

o

1

3

30.0

June

9

O

J

12

14

5

19

6.33+0.91

5

2

7

36.8

July

19

5

24

22

7

29

5.80+0.89

3

2

5

17.2

Aug.

n

1

5

12

9

5

14

7 . 00+0.79

2

-

2

14.2

Sept.

7

6

13

8

7

15

7. 50±0. 68

1

1

2

13.3

Oct.

5

6

1 1

6

9

15

7 . 50±0. 50

1

3

4

26.6

Nov.

5

4

9

7

6

13

6.50+0.65

2

2

4

30.7

Dec.

10

6

16

12

7

19

6. 33±0. 82

2

1

3

15.7

Total

87

54

141

410

A

73

183

6. 53 ±0. 98

23

19

42

22.9

Table 5

Pre-implantation losses in R. m. palliclior during 1979

Number of embryos Number of corpora lutea Pre-implanation losses

Mths. — — —

Right

Left

Total

Right

Left

Total

Mean±SE

Right

Left

Total

%

Jan.

3

2

5

4

1

5

5.00+0.00

—

—

_

Feb.

10

9

19

14

13

27

9.00±0.00

5

5

10

37.0

Mar.

27

25

52

32

30

62

7 . 75±0. 59

5

5

10

16. 1

Apr.

7

6

13

10

7

17

8.50+0.28

n

J

1

4

23. 1

May

9

12

21

13

12

25

6.05+Q.00

6

9

36.0

June

9

8

17

13

11

24

12,00+0.00

4

A

J

7

29.0

Aug.

12

11

23

19

18

37

9.25+0.47

7

1

14

37.8

Sept.

10

9

19

15

13

28

9.33+0.45

5

4

9

32.2

Oct.

11

10

21

15

16

31

7.75±0.75

4

6

10

28.6

Nov.

12

13

25

17

18

35

7.00+0.70

5

5

10

28.6

Dec.

15

16

31

20

26

46

9.20+0. 89

5

10

15

32.6

Total 125

121

246

172

165

337

8. 25±2. 35

49

49

98

29.08

64

REPRODUCTION BIOLOGY OF THE SOFT-FURRED FIELD RAT

Table 6

Transfer of blastocyst

Female

nos.

Corpora lutea Right Left

Embryos present Right Left

Transfer of blastocyst

1

2

3

1

4

From right

14

4

3

3

4

— do —

15

4

5

3

6

— do —

24

4

1

3

2

— do —

50

2

6

3

5

From left to right

per pregnant female did not vary significantly between months and between seasons except during winter when it was significantly (p < 0.01) lower than that in the summer season during both the years (Tables 4 & 5). The high production rate of ova during summer months may be due to influence of day length which is maximum in this period in the desert region. The mean number of produced ova (8.2) during 1979 did not differ significantly

(x2G) = 0.2416) than that of 1978 (6.5). Transfer of blastocyst

The transfer of blastocyst was investigated by comparing the number of corpora lutea and the actual number of embryos in both the ute- rine horns. Out of 68 pregnant females exa- mined during both the years, in five cases the transfer of blastocyst was observed. In female nos. 1, 14, 15 and 24 the blastocyst was trans- ferred from right to left, whereas, in female no. 50, it was transferred from left to right (Table 6). Transfer of blastocyst was also re- ported in Cutch Rock Rat, Rat t us cutchicus (Prakash et al. 1973).

Implantation rate in right and left uterine horns

During 1978, the number of implanted em- bryos was significantly higher (x2(i)=7.72, P < 0.05) in the right uterine horn. However, no significant difference (x2(!)=0.26) was found between right (125) and left (121) im- planted embryos during 1979. After pooling the embryo numbers for both the years of

Table 7

Distribution of litters of various sizes in the monthly collection of R. m. pallidior during 1978

Months

Frequency of occurrence of litters No. of implanted embryos

3 4 5 6

7

Total number of embryos

Mean±SE

Jan.

1

—

1

—

_

8

4.

00±0.66

Feb.

-

1

1

-

-

9

4.

, 50±0. 64

March

-

-

-

1

1

13

6.

,50±0.61

April

1

1

-

-

-

7

3.

,50±0.01

May

-

-

-

-

1

7

7.

00 ± —

June

1

1

1

-

-

12

4,

00±0.42

July

-

3

1

-

1

24

4.

80±0.69

Aug.

-

-

1

-

1

12

6.

,00±0.51

Sept.

-

-

-

1

1

13

6.

,50±0.58

Oct.

-

-

1

1

-

11

5,

, 50±0. 56

Nov.

-

1

1

-

-

9

4,

,50±0.61

Dec

—

1

1

-

1

16

5,

,33±0.86

Total

3

8

8

3

6

141

5,

03±0. 89

65

5

JOURNAL, BOMBAY NATURAL HIST . SOCIETY, Vol. 81

Table 8

Distribution of litters of various sizes in the monthly collection of R. m. pallidior during 1979

Months

Frequency of occurrence of litters

Number of implanted embryos 4 5 6 7

8

9

Total

number of

embryos

Mean±SE

Jan.

—

1

—

—

—

—

5

5.00±0.00

Feb.

-

1

1

-

1

-

19

6.33±0.93

Mar.

1

1

2

2

1

1

52

6.50±0.56

Apr.

1

-

-

-

-

1

13

6.50±0.50

May

1

1

2

-

-

-

21

5 . 25±0. 51

June

-

-

-

-

1

1

17

8.50±0.85

Aug.

1

-

2

1

-

-

23

5.75±0.62

Sept.

-

1

-

2

-

-

19

6.33±0.62

Oct.

-

3

1

-

-

-

21

5. 25 ±0. 39

Nov.

2

2

-

1

-

-

25

5.00±0.63

Dec.

1

1

—

2

1

-

31

6.20±0.73

Total

7

11

8

8

4

3

246

6. 15±1 . 87

study, it is observed that the embryonic rate in right uterine horn is significantly (x2(x) = 4.24, P < 0.05) higher than that of left uterine horn (Tables 4 & 5).

Prevalence of pregnancy

Pregnant females were found during the two years of the study but the intensity of repro- duction activity in the metad population was more during 1978 in which the pregnancy rate of 100 per cent was observed during 6 months, whereas, during 1979 it was only for two months. The period of maximum breeding activity (March-September, 1978) coinciding with the occurrence of larger litter size also. During 1979, however, 100 per cent pregnancies occur- red during August-September, the period of maximum precipitation, but the largest litters occurred during February to June only. It is also observed that after a low pregnancy rate during winter, a peak occurs during spring, followed by a low during summer and then again with a peak during monsoon (Tables 2

& 3) — a typical reproduction cycle observed in other desert rodents also.

Litter size

Litter size varied from 3 to 7 (av. 5.03) and 4 to 9 (av. 6.15) during 1978 and 1979 respectively (Tables 7 & 8). The two yearly average was found to be 5.69 while in 1978 the larger litter size was scattered over the period March to September, they were restrict- ed to the February- June period in 1979. Smallest litters were observed during the win- ter (November to January).

Super-foetation

One female R. m. pallidior bore four em- bryos in the left uterine horn, their mean crown rump length was 25 mm (new born young ones measure 34 mm in crown rump length). But the right uterine horn carried two embryos (crown rump length = 5 mm) which were quite healthy. Considerable difference in the size of embryos in the two uterine horns, suggests

66

REPRODUCTION BIOLOGY OF THE SOFT-FURRED FIELD RAT

that the second conception might have occur- red around 9 to 12 days of first pregnancy (gestation period is 20 days, Bindra and Sagar, 1968). This phenomenon in metad is probably being reported for the first time though it is common in mice (Barnett and Munro 1970), in Indian gerbil, T. i. indica (Jain 1970) and in the shrew, S. m. sindensis (Rana and Prakash 1979).

Lactation

Out of all the pregnant females collected during both the years, during the period. March-April and October-November, seven female metads (10.3 per cent. Tables 2 & 3) were found to be pregnant as well as lactating. The difference between the first year (3 preg- nant & lactating) and latter year (4 pregnant & lactating) did not vary significantly. The pre- sence of pregnant and lactating females in the collection suggests the possibility of occurrence of postpartum oestrus in R. m. pallidior and therefore, a regular littering throughout the breeding season is logical. In various species of small mammals, the presence of postpartum oestrus has been reported earlier (Prakash et ai. 1973, Asdell 1946, Barnett and Little 1968).

Embryonic Mortality Pre-implantation loss

Pre-implantation losses were judged by com- paring the number of corporalutea and the implanted embryos whereas, post-implantation losses were judged by identifying resorbed and mummified embryos in the uterus. In the sample of metads, the former type of mortality accounted for 22.9 and 29.0 per cent of the total number of ova produced during 1978 and 1979 respectively. The preimplantation mortality ranged from 13.3 per cent to 36.8

per cent during 1978 and 16.1 to 37.8 per cent in 1979 (Tables 4 & 5). The intensity of loss of ova during both the years was more in right (72) than in the left (68) uterine horn, the difference was, however, not statistically significant. Thus, the average annual pre- implantation mortality amounted to 26.92 per cent.

Post-implantation loss

One female (body weight, 54 g) bore four embryos, two in each uterine horn (their crown rump length was measured to 12-15 mm). Two in left horn were in healthy state but the two in right uterine horn were in mum- mified stage. Out of 387 total implanted em- bryos examined in this study, only 0.51 per cent account for post-implantation loss. In another case, a female during December possessed one embryo in right and three in left uterine horn. These four embryos were found to be resorbed, which amounts to 1.03 per cent loss. The total post-implantation loss comes to 1.55 per cent.

Annual Productivity

The number of young produced by an adult metad during the breeding season can be deter- mined by dividing the number of days of breed- ing season by number of days of gestation period (Prakash and Taneja 1969). The period of gestation in R. m. pallidior is 20 days (Bindra and Sagar 1968), and applying the data from this study we find that (365/20 = 18.20) litters can be produced by a female in a breeding season. Correcting this figure by multiplying it by the average prevalence of pregnancy (18.20 x 0.51 = 9.28), the number of litters per adult female comes to 9.28 Multiplying this with 5.69, the average num- ber of embryos per pregnant female during

67

JOURNAL, BOMBAY NATURAL HIST. SOCIETY, Vol. 81

the breeding season, it appears that (9.28 x 5.60 = 52.80) young ones can be produced by a female annually. If the pre-implantation (26.92 per cent) and post-implantation (1.55 per cent) losses are accounted for, the figure is corrected to 52.52 young per female per year.

Discussion

Rattus meltada is essentially a sub-mesic species, distributed all over the Indian plains but its subspecies, R. m. pallidior occurs in the arid and semi-arid zones of northern India (Prakash 1975). Its frequency of occurrence is poor in arid zones but it is found in fair numbers on the foothills and in desert areas which receive over 500 mm rainfall annually. Its breeding season appears to be similar to those rodents inhabiting tropical regions of the country and not that of the Saharo-Indian rodents which breed during winter as stated by Bodenheimer (1957). It breeds throughout the year with two peaks, one during spring and the other during monsoon and two lows, one during summer and the other in winter, the latter shows very low pregnancy rates as well as smallest litter size. Our study area at Bisalpur, on the foothills of the Aravalli ranges, does not experience severe climatic and vege- tational fluctuations as the low rainfall areas do. The metads being nocturnal and fossorial are not exposed to severities of climatic con- ditions. Secondly in this area, due to the pre- sence of irrigation cropping, green food is consumed by them for the greater part of the year. Experiencing these environmental condi- tions R. m. pallidior maintain rather a high prevalence of pregnancy rate with bimodal peaks, one in spring and the other in monsoon. It is surprising, however, that R. meltada in- habiting similar environmental conditions in

Uttar Pradesh, Punjab and Pakistan breed from March to October only (Bindra and Sagar 1968, Guraya and Gupta 1975, Srivastava 1966, Smiet et al. 1980). In southern India, however, R. m. meltada breeds all the year round (Blanford 1888-91, Lloyd 1909, Chan- drahas and Krishnaswami 1974). In Uttar Pradesh and Punjab the non-reproductive state of female metads and low breeding rate at our study area during winter may be due to the partial quiescent state of ovary. After this re- fractory period, their inherent physiologic activity is accelerated causing a spring peak in pregnancy rate in R. m. pallidior. The breeding activity slightly decreases during May and June, the hottest months of the year. Second main peak then occurs during the monsoon season. These two peaks well coincide with the availability of green food to R. m. pallidior which in turn influences their repro- ductive activity. R. m. pallidior , the northern subspecies of metad was even able to maintain a fairly high (50 to 80 per cent) prevalence of pregnancy during summer of 1978 and 1979 at the study site, when the air temperature was high and the relative humidity was low. The maintenance of a relatively high rate of repro- duction during summer is presumably a reflec- tion of the rodent’s efficient eco -physiological adaptive mechanisms to cope with the arid environment.

The mean litter size of R. m. pallidior was 5.69 which is slightly lower than 6.0 report- ed by Bindra and Sagar (1968) at Ludhiana, but more or less equal to that of Kanpur metads (5.7, Srivastava 1966). However, the lowest litter size was reported by Lloyd (1909) in India as a whole.

A comparison of mean litter size and lati- tudes of the localities revealed that it was the lowest, 4.8 in Karnataka (latitude 12°), and the highest, 6.0 in Punjab (latitude 30°) and

68

REPRODUCTION BIOLOGY OF THE SOFT-FURRED FIELD RAT

a mid position was occupied (5.7) by Bisalpur field metads (latitude 25°). Thus, a relation- ship between litter size and latitude exists, which was larger in Northern than in Southern India, irrespective of wide variations in their geographical range.

Another very interesting point emerged out of this study is the occurrence of superfoeta- lion which is not recorded in any other species of R. meltada in India.

Annual productivity of R. m. pallidior is comparatively higher, 52.52 young ones /year than other species of desert rodents such as R. cutchicus (19.96, Prakash et al 1973). T. i. indica (17.75, Jain 1970); and more or less equal to R. norvegicus (53.4 young/adult female/year; Southwick 1966).

Acknowledgements

We are grateful to the Director, Dr. H. S. Mann for providing necessary facilities and constant encouragement throughout the course of this study, to Dr. P. K. Ghosh, Head of Division of Animal Studies for various useful suggestions. Thanks are also due to staff mem- bers of Cartography Section of the Institute for drawing. The assistance of Sarvashri R. P. Mathur, B. K. Soni and Mala Ram during field work are also acknowledged.

Summary

The Soft-furred field rat, Rattus meltada pallidior (Ryley) were snap trapped every month during 1978 and 1979 at Bisalpur

R E FE

Asdell, S. A. (1946) : Patterns of mammalian reproduction. Comstock Publishers. Ithaca, N.Y., xi+670.

Blanford, W. T. (1888-91): The fauna of British Tndia. including Ceylon and Burma. Mammalia.

(25°7'N, 73°10'E) situated on the fringe of the Thar desert in India. The male R. m. palli- dior were found to be fecund in every month from January 1978 to December 1979. The fluctuations in testes weights coincided with the number of pregnant females. The left testes of adult rodents collected over two years were observed to be heavier than the right. The average testes weights were also found to vary significantly (P < 0.05) from month to month.

The production of ova ranged from 5.5 to 10.0 (av. 6.5) and 5 to 12 (av. 8.2) during 1978 and 1979 respectively. The maximum number of ova per pregnant female metad was produced during the period of maximum preci- pitation. The implanted embryos per pregnant female varied from 3 to 7 and 4 to 9 during 1978 and 1979, average being 5.03 and 6.15 respectively.

Embryonic mortality before implantation was found to be 22.9 and 29.0 per cent dur- ing 1978 and 1979 respectively and due to mummification and resorption 0.51 per cent and 1 . 03 per cent respectively.

Considering these losses, litter size and pre- valence of pregnancy during both the years, the annual production was estimated to be 52.52 young ones per annum.

R. m. pallidior litters all through the year. The prevalence of pregnancy was found to be broadly correlated with the day length. Maxi- mum production appears to have been influ- enced by the level of nutrition which was found to be maximum during monsoon due to availa- bility of green forage in the desert.

E N C E S

Taylor and Francis, London.

Barnett, S. A. & Little, M. J. (1968) : Concep- tion and parturition of mice at two temperatures. /. Reprod. Fertil. 15: 295-304.

Barnett, S. A. & Munro, K. M. H. (1970) :

69

JOURNAL, BOMBAY NATURAL HIST. SOCIETY, Vol. 81

Superfoetation of mice. Nature, 227 : 1343-1344.

Bindra. O. S. & Sagar, Prem (1968): Breeding habits of the field rat, Rattus meltada. J. Bombay nat. Hist. Soc., 65: 477-481.

Bodenheimer, F. S. (1957) : The ecology of mam- mals in Arid Zone. In: Human and Animal Ecology, Review of Research, Arid Zone Research, UNESCO, Paris, 8: 100-137.

Chandrahas, R. K. & Krishnaswami, A. K. (1974) : Ecology of the Soft-furred field rat, Rattus meltada meltada (Grey) in Kolar, Mysore State. J. Bombay nat. Hist. Soc. 70: 447-457.

Guraya, S. S. & Gupta, S. (1975) : Observations on the reproductive biology of the Soft-furred field rat ( Millardia meltada ). Proc. All India Rodent Seminar, Ahmedabad, 70-75.

Jain, A. P., (1970): Body weights, sex ratio, age structure and some aspects of reproduction in the Indian gerbil, Tatera indica indica Hardwicke, in the Rajasthan desert, India. Mammalia 34: 416-432.

Lloyd, R. E. (1909): The races of Indian rats. Rec. hid. Mus. 3: 1-100.

Prakash, Ishwar, (1975): The ecology and zoo- geography of mammals. Chapter XIX in “Environ- mental Analysis of the Thar Desert” (Edited by Dr. R. K. Gupta & Dr. I. Prakash) . English Book Depot, Dehradun: 448-467.

Prakash, Ishwar, & Taneja, G. C. (1969): Re- production biology of the Indian desert hare, Lepus

nigricollis dayanus Blanford. Mammalia, 33: 102-117.

Prakash, Ishwar, Rana, B. D. & Jain, A. P. (1973) : Reproduction in the Cutch Rock Rat, Rattus cutchicus cutchicus in the Indian desert. Mammalia, 37: 452-467.

Rana, B. D. (1981): Ecological studies on Soft- furred field rat, Rattus meltada pallidior (Ryley, 1914) with special reference to its feeding and breeding habits, population structure and interspecific relation- ship in mixed population of small mammals (Un- published Ph.D. Thesis submitted to Agra Univ., Agra) .

Rana, B. D. & Pra-kash Ishwar, (1979): Repro- ductive biology and population structure of the house shrew, Suncus murinus sindensis in western Rajas- than. Sauget. Kunde, 44: 333-343.

Rana, B. D. & Prakash Ishwar, (1980): The metad — a serious rodent pest of Indian Agriculture. Indian Farming, xxxix(10: 21 & 23.

SouTHWiCK, C. H. (1966): Reproduction, morta- lity and growth of murid rodent populations. Proc. Indian Rodent Symp., Calcutta USAID & John Hop- kins CMRT, New Delhi, 152-176.

Smiet, A. ., Fulk, G. W. & Lathiya, S. B. (1980) : Rodent ecology in sugarcane in lower Sind, Pakistan. Acta Theriol., 25: 81-87.

Srivastava. A. S. (1966): Rodent control for in- creased food production. Rotary Club (West) Kanpur, 1-52.

70

FLOWERING PLANTS AROUND THE HOLY SHRINE OF KEDARNATH, UTTAR PRADESH1

J. K. Semwal2 (With a text-figure)

This paper gives an account of the angiospermic flora of Kedarnath, which is an alpine zone in Garhwal Himalaya. A list of 262 species of flowering plants represented by 149 genera and 52 families is given from 3200 to 3800 metre altitude a.s.l. The dominant families of the area are: Ranunculaceae, Brassicaceae, Caryophyllaceae, Rosaceae, Apiaceae, Asteraceae, Ericaceae, Primulaceae, Scrophulariaceae, Lamiacea, Polygonaceae, Orchidaceae, Liliaceae and Poaceae.

Introduction

Garhwal Himalaya is famous for its vegeta- tion and some of the noblest peaks of the world mountain system namely, Nanda Devi, Kamet, Trisul, Dunagiri, Chaukhamba, Nil- kantha, Badrinath, Kedarnath and others. Kedarnath is the first among the Holy ‘Panch (five) Kedars’ of Uttarakhand, the other four being Madmaheshwar, Tungnath, Rudranath, and Kalpeshwar respectively. Mandakini river has its origin from Kedarnath glacier, and con- fluences with Alaknanda at Rudraprayag and Bhagirathi at Deoprayag whenceforth it is named ‘Ganga’. The explored area lies in bet- ween 30° 40-45' N and 79° 0-5' E. The main rock components are of crystalline and meta- morphic nature like other parts of the Hima- layas (Fig. 1).

Compared with other sectors of the Hima- layas, this region is humid. The characteristic feature of Kedarnath is the presence of marshy habitats, clothed with the typical marshy vege- tation consisting of Primula munroi, J uncus

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1 Accepted May 1982.

2 High Altitude Plant Physiology Research Centre, Garhwal University, Srinagar- 246 174.

spp. Triglochin palustre, sedges and species of Epilobium and others. Callitrache verna is a delicate aquatic herb floating on the water surface of ditches in these marshy localities.

In this region timberline zone is not conspi- cuous, as is observed in most of the alpine re- gions of Garhwal Himalaya, however, the dis- tributional pattern of the common taxa is the same. The trees other than Betula utilis are present up to the elevation of 3200 m or slightly more according to the aspect of the slopes. The tree population is very scarce at the upper limits and gradually increases in density to- wards lower elevations. The major elements of temperate forests are Aesculus indica, Lyonia ovalifolia, Fraxinus excelsior, Buxus walli- chiana and species of Acer, Pyrus, Carpinus, Quercus and Rhododendron. The tree species reaching the highest limit are Betula utilis, Syringa emodi, Viburnum foetens, Prunus cor- nuta, Abies pindrow, Taxus baccata and species of Acer, Pyrus and Rhododendron associated with the shrubs like the species of Berberis, Spiraea, Rosa, Cotoneaster and climbers like the species of Clematis, Smilax and occasion- ally Codonopsis. Elaegnus umbellata and Pyrus lanata can be seen here and there on road sides up to 3000 m along with Piptanthus

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Fig. 1.

/ LOWERING PLANTS AROUND THE HOLY SHRINE OF KEDARNATH

nepalensis , Cotoneaster affinis and some others. Unlike other alpine zones of the western Himalayas pure stands of the scrubs of Rhodo- dendron campanulatum are lacking in this part.

The pattern of seasonal appearance of vari- ous plant species in this alpine zone is similar to that of Tungnath (Semwal et al, 1981). The vegetation of the glacial bed is quite distinct compared to that of the meadows and marshy lands. The species found in these habitats are tufted and very much reduced in size forming the continuous interwoven matted cushions on the grounds. Androsace villosa and Arenaria per- levis are the typical examples of such vegeta- tion, Primula nivalis , P. minutissima and Helerpestes sermentosa are found in rock crevices in these harsh habitats. The plant species on boulders and dry rocky faces are the species of Sedum, Lychnis, Potentilla, Arcyosperma, Rheum, Senecio, Ligularia, Seli- num. Goody era and many others. The vege- tation of the ravines is constituted by the species of Pedicularis, Epilobium, Impatiens, Polygonum and Rumex etc. Chamaenerion latifolium and Polygonum affine are the most attractive species of the ravines. Danthonia cumins ii is the dominant alpine grass on steep slopes like in other alpine zones.

This alpine zone was previously explored by M. A. Rau during the last week of Sept- ember 1958 (Ran 1961). He recorded about 53 species of flowering plants from Kedarnath. T have been working on the alpine flora of Tungnath (Semwal. 1981; Semwal et al, 1981; Semwal and Gaur, 1981). During the course of these studies it was considered useful to ex- plore the neighbouring alpine zones for com- parison. Collections were made during early Tune 1978 and late July 1981. The specimens were compared with the authentic specimens of the ‘Herbarium of Botanical Survey of India, Northern Circle Dehradun’ (BSD) and are de-

posited in the herbarium of Garhwal Univer- sity, Srinagar, U.P. Bentham and Hooker f.’s system is followed with slight modifications as proposed by Hutchinson (1973) in the arrangement of families.

The little known taxa recorded in the pre- sent work are, Circaeaster agrestis, Gentiana albicalyx, Cypripedium elegans and Herminium pugioniforme. Circaeaster agrestis was collect- ed from an elevation of 3200 m and the other three were collected between 3500-3600 m alti- tudes.

Enumeration

DICOTYLEDONS

Ranunculaceae

Aconitum heterophyllum Wall, ex Royle Herb with dull green purple veined flowers. July, 1981. Semwal 7210.

Anemone narcissifolsa Linn, var polyanthes Finnet et Gagnep.

White flowered hairy herb on rocks. July, 1981. Semwal 7242.

A. oMusiloba D. Don

Herb in meadows with white and blue flowers. June, 1978. Semwal 7157.

A. rivularis Buch.-Ham.

Large herb with white flowers. July, 1981. Semwal 7167.

A. tetrasepala Royle

Herb with cream -white flowers. June, 1978. Semwal 7179.

Caltha palustris Linn.

Marshy herb with brilliant-yellow flowers. June, 1978. Semwal 7156.

Clematis barbellata Edgew.

Climbers, sometimes spreading on grounds in alpine slopes, flowers purplish. June, 1978. Semwal 7168.

Halerpestes sarmentosa (Adms) Kom. et Klob Small glabrous herb with yellow flowers. July, 1981. Semwal 7221.

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Oxygraphis polypetala (Royle) Hook. f. et Thoms.

Scapose glabrous herb with yellow flowers and persistent calyx. June, 1978. Semwal 7138. Ranunculus diffusus DC.

Glabrascent herb with yellow flowers. June, 1978. Semwal 7162.

R. hirtellus Royle

Herb abundant in marshy meadows, flowers yellow. June 1978. Semwal 7159.

Thalictrum alpinism Linn.

Small glabrous herb with minute, drooping purple flowers. July, 1981. Semwal 7211.

T. minus Linn.

Erect herb with purplish flowers. July, 1981. Semwal, 7234.

Berberidaceae

Berberis edgeworthiana Schneid.

Tall shrub with yellow flowers, spines few. June 1978. Semwal 7169.

B. jaeschkeana Schneid.

Spinous shrub with yellow flowers. July, 1978. Semwal 7184.

Podophyllum hexandrum Royle Herb with palmate leaf and drooping fruit, very rare. July. 1981. Semwal 7212.

ClRCAEASTERACEAE

Circaeaster agrestis Maxim.

A small glabrous herb growing in crevices, perianth very much reduced, rare. July, 1981. Semwal 7213.

Papaveraceae

Meconopsis aculeata Royle

Aculeate herb with sky-blue flowers, in ravines. July, 1981. Semwal 7217.

M. robusta Hook. f. et Thoms.

Robust herb with light yellow flowers, not common. July, 1981. Semwal 7171.

Fumariaceae

Corydaiis cornuta Royle Spreading yellow flowered herb. July 1981. Semwal 7165.

C. cashmeriana Royle

Small delicate herb with bulbous base and blue flowers. June, 1978. Semwal 7136.

C. govaniana Wall.

Tufted herb with yellow flowers. June 1978. Semwal 7147.

C. vaginans Royle

Glabrous herb with yellow flowers. July, 1981, Semwal 7215.

Brassicaceae

Arcyosperma primulifoSium (Toms) O. E. Schulz

Lithophytic white flowered herb. June, 1978. Semwal 7141.

Barbarea vulgaris R. Br.

Erect herb with yellow flowers. June, 1978. Semwal 7158.

Draba gracillima Hook. f. et Thoms.

Weak herb with minute yellow flowers. June, 1978. Semwal 7142.

Lepidium ruderale Linn.

Glabrous herbs with fruits. July, 1981. Sem- wal 7208.

Megacarpaea polyandra Benth.

Large herb with dull white flowers. July 1981. Semwal 7371.

Sisymbrium wallichii Hook. f. et Thoms.

Herb with white flowers with long pods. July 1981. Semwal 7219.

Thlaspi andersosiif (Hook. f. et Thoms.) O. E. Schulz

Herb with white flowers. June, 1978. Semwal 7148.

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FLOWERING PLANTS AROUND THE HOLY SHRINE OF KEDARNATH

VlOLACEAE Viola biflora Linn.

Small herb with yellow flowers. July, 1981. Semwal 7214.

V. serpens Wall.

Herb with violet flowers. June, 1978. Sem- wal 7170.

Caryophyllaceae

Arenaria perlevis (Williams) Handl.-Mazz.

Lithophytic white flowered herb. July, 1981. Semwal 7222.

Cerastium vulgatum Linn.

Herb with white flowers. June, 1978. Sem- wal 7127.

Gypsophila cerastioides D. Don Herb with pink streaked flowers. July 1978. Semwal 7105.

Lychnis indica Benth.

Herb with fimbriate petals. June, 1978. Sem- wal 7172.

L. nutans Benth.

Pubescent herb with purplish flowers. July, 1981. Semwal 7254.

L. pilosa Edgew.

Herb on rocks with white flowers. July, 1981. Semwal 7246.

Sagina saginoides (Linn.) Karsten

Dark green glabrous white flowered prostrate herb. June, 1978. Semwal 7114.

Silene vulgaris (Moench) Garcke Large glabrascent herb with greenish- white flowers. July, 1981. Semwal 7220.

Stellaria uliginosa Edgew. et Hook. f.

Apetalous spreading herb. July, 1981. Sem- wal 7185.

S. cherleriae (Fisch.) Williams Cushioned herb with white flowers. July, 1981. Semwal 7223.

S. himalensis Majumdar Glabrous herb with bright-white flowers. June, 1981. Semwal 7191.

S. patens D. Don

Diffused herb with white flowers. July, 1981. Semwal 7301.

Hypericaceae

Hypericum hookeriana Wt. et Arn.

Small shrub with large yellow flowers. July, 1981. Semwal 7312.

H. nepaulense Choisy

Herb with yellow flowers. July, 1981. Sem- wal 7224.

CtERANIACEAE

Geranium coliinum Stephan ex Willd.

Glabrous herb with dark pink flowers. July, 1981. Semwal 7227.

G. pa lustre Linn.

Large glabrascent herb with pink flowers. July, 1981. Semwal 7235.

G. wallichianum Sw.

Straggling herb with pink-purple flowers. July, 1981. Semwal 7236.

Balsam i naceae

Impatiens amplexicaulis Edgew.

Erect pink flowered herb. July, 1981. Sem- wal 7303.

I. giandulifera Edgew.

Tall herb in ravines with dark pink flowers. July, 1981. Semwal 7309.

I. thomsonii Hook. f.

Marsh herb with umbellate pale flowers. July, 1981. Semwal 7248.

Kutaceae

Skimmia laureola Sieb. et Zucc.

Prostrate shrub, flowers greenish. June, 1978. Semwal 7186.

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Aceraceae

Acer acuminatum Wall, ex D. Don Tree with white flowers and pink winged fruits. June, 1978. Semwal 7199.

A. caesium Wall, ex Brandis

Large tree, flowers white. June, 1978. Sem- wal 7188.

Fabaceae

Astragalus chlorostachys Lindl.

Erect herb with yellow flowers. July, 1981. Semwal 7225.

Lotus corniculatus Linn.

Decumbent herb with yellow or orange flowers. July, 1981. Semwal 7241.

Parochetus communis Buch.-Ham ex D. Don Prostrate herb with blue flowers. July, 1981. Semwal 7187.

Piptanthus nepalensis D. Don Pubescent shrub with yellow flowers. June, 1978. Semwal 7189.

Trifolium repens Linn.

Herb with white flowers in globose heads. June, 1978. Semwal 7174.

Trigonella pubescens Edgew. ex Baker Diffused herb with yellow flowers. June, 1978. Semwal 7125.

Rosaceae

Cotoneaster acuminatus Lindl.

Erect shrubs, flowers white. June, 1978. Semwal 7175.

C. rotundifolius Wall, ex Lindl.

Procumbent shrub with pinkish-white flowers. July, 1981. Semwal 7226.

Fragaria nubicola Lindl. ex Lacaita Herb with white flowers. June, 1978. Semwal 7128.

F. daltoniana Gay

Stoloniferous herb with dark green leaves and white flowers. June, 1978. Semwal 7119. Geum urbanum Linn.

Erect herb with yellow flowers. July, 1981, Semwal 7302.

G. datum (Royle) Hook. f.

Leaves forming rosette, flowers large, yellow and drooping. July, 1981. Semwal 7255. Potentilla cuneifolia Betrol.

Base shrubby, leaves glabrous, trifoliate, flowers yellow. July, 1981. Semwal 7228.

P. arbuscula D. Don

Shrubs on rocks with yellow flowers. July 1981. Semwal 7243.

P. argyrophylla Wall, ex Lehm.

Herb with silvery trifoliate leaves and yellow flowers. July, 1981. Semwal 7304.

P. atrosanguinea Lodd.

Herb with crimson-red flowers, abundant, June, 1978. Semwal 7159.

P. doubjouneana Camb.

Glabra scent herb on rocks with trifoliate leaves and yellow flowers. July, 1981. Semwal 7229.

P. fulgens Wall, ex Hook.

Silky herb with yellow flowers. June, 1978. Semwal 7176.

P. mkrophylla D. Don var. commutata Hook. f.

Herb, leaves green-glaucous above and pubescent beneath, flowers yellow. June 1978. Semwal 7149.

P. polyphylla Wall, ex Lehm.

Pubescent diffused herb with yellow corym- bose flowers. July, 1981. Semwal 7262.

Rosa macrophylla Lindl.

Erect shrubs with large pink flowers. June, 1978. Semwal 7177.

R. sericea Lindl.

Shrub with white flowers. July, 1978. Sem- wal 7230.

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FLOWERING PLANTS AROUND THE HOLY SHRINE OF KEDARNATH

Rubus nutans Wall.

Creeping shrubby herb with white flowers. June, 1978. Semwal 7178.

R. pedunculosus D. Don

Rambling shrubs with light pink flowers. June, 1978. Semwal 7183.

Sibbaldia cuneata Hornem. ex O. Ktze.

Creeping, base shrubby, flowers minute yellow. June, 1978. Semwal 7123.

S. micropetala (D. Don) Hand.-Mazz.

Diffused herbs with minute yellow flowers.

July, 1981. Semwal 7315.

S. purpurea Royle

Similar to S. cuneata but with pentafoliate leaves and pink flowers. June, 1978. Semwal 7102.

Spiraea bella Sims.

Shrub with light pink flowers. July, 1981. Semwal 7263.

S. canescens D. Don

Shrub with white flowers. July, 1981. Sem- wal 7319.

S. vestita Wall, ex G. Don Herbs, leaves pinnate, flowers greenish- white. July, 1981. Semwal 7249.

Saxifragaceae

Chrysospleniuni tenelium Hook. f. et Thoms.

Delicate creeping glabrous herbs with small green-yellow flowers. June, 1978. Semwal 7132. Parnassia nubicola Wall, ex Royle Glabrous herb with single, sessile, cauline leaf, flowers solitary, terminal. July, 1981. Sem- wal 7207.

Saxifraga brachypoda Wall. var. fimbriata (Wall.) Engl, et Irmsch.

Tufted herbs with yellow flowers. July, 1981. Semwal 7247.

S. bmnoniana Wall, ex Sternb.

Stoloniferous with solitary yellow flowers. July, 1981.

S. diversifolia Wall, ex DC.

Erect glabrous herbs with corymbose yellow flowers. July, 1981 Semwal 7305.

S. pallida Wall, ex DC.

Herb with white flowers. July, 1981. Sem- wal 7250.

Gross ulariaceae

Ribes glaciaSe Wall.

Shrub with unisexual purplish flowers. June, 1978. Semwal 7117.

Crass ulaceae

Seduin bupleuroides Wall, ex Hook. f. et Thoms.

Fleshy herbs with purplish-red flowers. July, 1981. Semwal 7203.

S. heterodontum Hook. f. et Thoms.

Fleshy herbs with greenish or yellowish flowers. June, 1981. Semwal 7118.

S. imbricatuni (Edgew.) Walp.

Herb on rocks with dull yellow flowers. July, 1981. Semwal 7232.

S. qtiadriffduni Pall.

Densely tufted herbs with small red flowers. July, 1981. Semwal 7206.

S. trullipetaluin Hook. f. et Thoms.

Herbs on boulders with yellow flowers. July, 1981. Semwal 7233.

Callitrichaceae

Callitriche verna Linn.

Delicate aquatic herbs floating on water surface. July, 1981. Semwal 7306.

Onagraceae

Chamaenerion latifoSium (Linn.) Sw.

Herbs with beautiful purple-pink flowers in glacial beds. July. 1981. Semwal 7238.

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Epilobium iaxum Royle Erect glabrous herbs in marshy localities. July, 1981. Semwal 7307.

E. palustra Linn.

Herbs with pinkish-white flowers. July, 1981. Semwal 7317.

E. royleamini Hausskn.

Herbs with pink flowers. June, 1981. Sem- wal 7180.

Apiaceae

Acronema tenera Edgew.

Small herb in crevices with minute purplish flowers. July, 1981. Semwal 7239.

Bupleurum longkaule Wall, ex DC.

Glabrous procumbent herb with brownish- black flowers. July, 1981. Semwal 7308. Heracleum brunonis (DC.) C. B. Clarke Erect herb on rocky slopes with white flowers. July, 1981. Semwal 7318.

Selinum candoSSei DC.

Robust aromatic herbs with white umbels. July, 1981. Semwal 7279.

S. vaginatum C. B. Clarke Herb in marshy places with dark-green glossy leaves and white flowers July, 1981. Semwal 7310.

Trachydium roylei Lindl.

Herb in the meadows with white flowers and black tipped fruits. July, 1981. Semwal 7369.

Yicatia coniifolia DC.

Glabrous herb with fruits. June, 1978. Sem- wal 7152.

Y. millefolia (Klotzsch) C. B. Clarke Herb with small flowered umbels. June, 1978. Semwal 7164.

Caprifoliaceae

Lonicera asperifolia (Decne) Hook. f. et Thoms.

Shrub with yellow flowers. June, 1978. Sem- wal 7130.

L. myrtillus Hook. f. et Thoms, var depressa Rehder

Decumbent shrub with white flowers. June 1978. Semwal 7121.

L. obovata Royle ex Hook. f. et Thoms.

Shrub with white corolla gibbous at the base. June, 1978. Semwal 7116.

Yiburnum foetens Decne Small tree with white flowers and red drupes. June, 1978. Semwal 7181.

Rubiaceae

Galium acutum Edgew.

Spreading herb with minute greenish-white flowers. July, 1981. Semwal 7240.

G. asperuloides Edgew.

Decumbent herbs with dull white flowers. July, 1981. Semwal 7244.

G. mollugo Linn.

Scabrous herbs with whitish flowers. July, 1981. Semwal 7311.

G. rotundifolium Linn.

Decumbent herbs, leaves in whorls of four, flowers white. June, 1978. Semwal 7182.

Valeria naceae

Yaleriana hardwickii Wall.

Herb with white flowers. July, 1981. Semwal 7197.

Dipsacaceae

Morina Songifolia Wall, ex DC.

Spinous herbs with pink flowers. July, 1981. Semwal 7245.

Asteraceae

Anaphalis cuneifolia Hook. f.

Herbs with cuneate leaves and white flowers. June, 1978. Semwal 7190.

A. nepalensis (Spreng.) Handl-Mazz.

Woolly herbs with lanceolate leaves and white heads. July, 1981. Semwal 7341.

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FLOWERING PLANTS AROUND THE HOLY SHRINE OF KEDARNATH

A. royleana DC.

Woolly herbs with dirty white heads. June, 1978. Semwal 7196.

Artemisia roxburghiana Besser var. grata Hook. f.

Erect herbs, leaves green above and white beneath. July, 1981. Semwal 7348.

Aster peduncularis Wall.

Herbs with violet flowers. July,