*
JOURNAL
OF THE
BOMBAY NATURAL HI STORY
SOCIETY
VOL. 93, No. 1
April 1996
BOARD OF EDITORS
Executive Editor J.C. DANIEL
M.R. ALMEIDA F.V. BOLE
M.K. CHANDRASHEKARAN B.F. CHHAPGAR B.V. DAVID R. GADAGKAR ANIL GORE
A.J.T. JOHNSINGH AJITH KUMAR A.R. RAHMANI J.S. SAMANT E.G. SILAS J.S. SINGH R. WHITAKER
Assistant Editor K.P. SHIRODKAR
>
INSTRUCTIONS TO CONTRIBUTORS
1. Papers which have been published or have been offered for publication elsewhere should not be submitted.
2. Papers should be submitted in duplicate, typed double space. Preferably an additional copy should be submitted on a floppy diskette (3.5" or 5.25"), using Word Star, Word Perfect. MS Word or in MS DOS.
3. Trinomials referring to subspecies should only be used where identification has been authentically established by comparison of specimens actually collected.
4. Photographs for reproduction must be clear, with good contrast. Prints should be at least 9x12 cm and on glossy glazed paper. Text - figures, line drawings and maps should be in Indian ink, preferably on Bristol board.
5. References to literature should be placed at the end of the paper, alphabetically arranged under author's name, with the abridged titles of journals or periodicals in italics and titles of books or papers in roman type, thus:
Aluri, Raju J.S. & C. Subha Reddi (1995): Ecology of pollination in two cat-mint species. J. Bombay not. Hist. Soc. 92(1): 63-66.
Prater, S.H. (1948): The Book of Indian Animals. Bombay Natural History Society, Bombay.
6. Each paper should be accompanied by an abstract, normally not exceeding 200 words, and 6-8 key words. Key Words should include the scientific names of important species discussed.
7. 25 reprints will be supplied free of cost to authors of main articles. In the case of new descriptions, reviews and miscellaneous notes, authors will be sent a free copy of the Journal.
8. The editors reserve the right, other things being equal, to publish a members's contribution earlier than a non-members's.
Hornbill House,
Shaheed Bhagat Singh Road, Bombay 400 023.
Editors,
Journal of the Bombay Natural History Society
VOLUME 93 (1): APRIL 1996
Date of Publication: 01-4-1996
CONTENTS
NIGHT ROOSTING AND LUNAR PHOBIA’ IN INDIAN FALSE VAMPIRE BAT LYRA ( With seven text -figures)
By R. Subbaraj and J. Balasingh 1
RELATIONSHIP BETWEEN DIVE AND POST-DIVE PAUSE WHILE FORAGING IN TWO DIVING DUCKS OF LAKE M ANSAR (With three text-figures )
By Y.R. Malhotra, N. Dcskyong and PS. Pathania 8
NOTES ON THE FEEDING AND BREEDING BEHAVIOUR OF GYMNOPLEURUS GEMMATUS HAROLD AND GYMNOPLEURUS M1LIAR1S ( F.) (COLEOPTERA: SCARABAEIDAE)
( With a text-figure)
By K. Veenakumari and G.K. Veeresh 13
NOTES ON LONG-EARED HEDGEHOG HEMIECHINUS A UR IT US (GMELIN) (With a text-figure)
By Satish Kumar Sharma 20
ROOST SELECTION BY INDIAN PEAFOWL (PAVO CR1STATUS) IN GIR FOREST, INDIA
By Pranav Trivedi and A.J.T. Johnsingh 25
TAXONOMIC AND NOMENCLATURAL STATUS OF MYRIONEURON R.BR. EX HOOK. F. (RUBIACEAE)
By D.B. Deb 30
SOME ASPECTS OF BIRD/MAMMAL ASSOCIATIONS: CONTRIBUTIONS FROM THE INDIAN PLAINS AND THE ZIMBABWE PLATEAU
By D. Ewbank 34
FISH FAUNA OF PERIYAR TIGER RESERVE
By V.J. Zacharias, A.K. Bhardwaj and PC. Jacob 39
PRELIMINARY OBSERVATIONS ON THE IMPORTANCE OF A LARGE COMMUNAL ROOST OF WINTERING HARRIERS IN GUJARAT (NW. INDIA) AND COMPARISON WITH A ROOST IN SENEGAL (W. AFRICA)
By Roger Clarke 44
STUDIES ON AMPHIPODS OF VISAKHAPATNAM COAST (With three plates)
By P. Somanadha Rao, K. Hanumantha Rao and K. Shyamasundari 51
YELLOWTHROATED BULBULS AT HORSLEY HILLS (With a text-figure)
By S. Subramanya and J.N. Prasad 55
NEW DESCRIPTIONS
A NEW SPECIES OF AGIOMMATUS CRAWFORD (HYMENOPTERA: PTEROMALIDAE) — AN EGG PARASITE OF A CUTWORM ON MULBERRY IN BANGALORE (KARNATAKA)
(With five text-figures)
By PM. Sureshan and T.C. Narendran 59
CROSSOCHEILUS PERIYARENSIS, A NEW CYPRINID FISH FROM THANIKKUDY (THEKKADY), KERALA, INDIA (With a plate)
By A. G.K. Menon and PC. Jacob 62
ADDITION OF THREE NEW SPECIES OF CHALCOLEPIS CANDEZE (COLEOPTERA, ELATERIDAE: HEMIRRHIPINAE) TO THE INDIAN FAUNA (With twelve text-figures)
By Punam, L.K. Vats and M.S. Saini 64
FIVE NEW SPECIES OF THE GENUS MACROCHELES LATREILLE (ACARINA: MACROCHELIDAE) FROM EASTERN INDIA (With twenty-nine text-figures)
By R.K. Roy
68
OBITUARY
C.V. KULKARNI (191 1-1995) (With a plate) By B.F. Chhapgar
REVIEWS
74
1. A NATURALIST IN KARBI ANGLONG
Reviewed by Philippa Mukherjee 77
2. ORN1THOBOTANY or Indian weaver birds
Reviewed by Zafar Futehally 77
MISCELLANEOUS NOTES
MAMMALS
1. Meal eating by lion-tailed Macaque,
Macaco silenus (Zimmermann)
By G. Uniapathy and A. Prabhakar 79
2. Panther, Pantlicra pardus (Linnaeus) with guinea worm infection
By Raza H. Tehsin 79
3. Some notes on Himalayan palm civet,
Panama larva la (Hainilton-Smith)
(Carnivora: Viverridac)
By M. L. Narang 80
4. Occurrence of the wolf (Canis lupus Linnaeus) in Rewa District (M.P.)
By A.M.K. Bharos 81
5. A record of the Nilgiri Marten
(Maries gwalkinsi Horsfield) in Upper Bhavani
By V. Gokula and N.K. Ramachandran 82
6. Back riding and possible aerial mating amongst the large fruit bat,
Ptempus giganleus (Brunnieh)
By A.M.K. Bharos 82
7. Interbreeding between grizzled giant squirrel, Ratufa macroura (Pennant) and Malabar giant squirrel, li. indica (Erxleben)
By Justus Joshua 82
8. Unusual feeding behaviour of squirrel,
Funanibulus spp.
By D.N. Haril 84
9. Toxicity of wax blocks against rodents in laboratory and field conditions
By Y. Saxena 84
10. Fish oils as alternative to river dolphin,
Plalanista gangetica (Lebeck) oil for fishing catfish Clupisoma garua in the river Ganges. India By R. S. Lai Mohan and
K.V. Mohammed Kunhi 86
BIRDS
1 1 . Large cormorant Phalacrocorax carbo sinensis (Shaw) breeding in the Nilgiris
By Manoj V. Nair 89
12. Record of the painted spurfowl Galloperdix lunulata (Valenciennes) in Ramgarh Sanctuary of district Bundi. Rajasthan
By Shantanu Kumar 89
13. Painted spurfowl, Galloperdix lunulata (Valenciennes) in Rajasthan
By Ashok Kumar Sharma 90
14. First record of fresh water crab Paralelphusa spp. in the barn owl’s Tylo alba (Scopoli)
diet in Tranquebar Taluk, Tamil Nadu, South India By R. Kanakasabai 90
15. Nesting in anchor-pipe by Brahminy myna.
Slurnus pagoda rum (Gmelin)
By Satish Kumar Sharma 91
16. Intraspecific brood parasitism in the common myna, Acridotheres Irislis (Linn)
By Salwant K. Dhanda and Manjit S. Dhindsa 91
17. Sideways leap-frogging by the large grey babblers, Turdoides nialcobni (Sykes)
By A.M.K. Bharos 93
18. Pipit (Antlius sp.) preying on leeches
By Manoj V. Nair 93
19. On some new breeding records of walerbirds from the Delhi region
By Abdul Jamil Urfi 94
20. Recent additions to the bird list of the Lakshadweep islands
By V. Santharam, D.N. Mathew. George Mathew and Tara Gandhi 95
REPTILES
21. The keeled box turtle Pyxidea nwulwlii Gray on
the north bank of the Brahmaputra — a new record By Anwaruddin Choudhury 97
22. Reproductive behaviour of Indian mugger
( Cmcodylus palustris) at Bhor-Saindan Crocodile Sanctuary in Haryana
By R.C. Gupta and C.S. Bhardwaj 97
23. Records of water snakes (Homalopsidae: Serpentes) from Gujarat state
By Raju Vyas 101
AMPHIBIA
24. Comments on the note “First record of
Microhyla rubra (Jerdon) (Amphibia: Anura)
from Maharashtra” by Kainble and Ghate
By C. Radhakrishnan 101
FISHES
25. Puntius dukai Day (Pisces: Cyprinidae) —
A new record from Uttar Pradesh hills
By K.D. Joshi and PC. Joshi 102
26. Record of new fishes from Periyar Tiger Reserve
By L.K. Arun, C.P. Shaji and PS. Easa 103
INSECTS
27. Biology and Altica coerulea (Oliv.)
(Chrysomelidae: Coleoptera) —
A potential bio-control agent against Jussiaea repens L.
By U.K. Sar, C.K. Sar and K.K. Sar 104
28. Study of male Genitalia of some species genus of Pheropsophus (Branchinini: Carabidae:
Coleoptera) and its taxonomic importance
By S.P. Singh 106
29. Ciliate infestation on the phytal Halacaridae (Acari) from the Kovalam Beach (Kerala Coast)
By Tapas Chatterjee 107
30. Delphinium dltissimum Wall. (Ranunculaceae)
— a new record for Sikkim Himalaya
By S.Z. Lucksom 108
3 1 . Pycnoplinthopsis bhutanica (Hara) Jafri (Brassicaceae): A new record from West Himalaya
By D.S. Rawat, L.R. Dangwal.and R.D. Gaur 109
32. Kosteletzkya vitifolius (Linn.) Comb. Nov.
By Neelam Patil and M.R. Almeida 1 1 1
33. Some new records of legumes from Garhwal Himalaya
By L.R. Dangwal, D.S. Rawat and R.D. Gaur 113
34. Stomata on seed of Bauhinia purpurea L. (Leguminosae: Caesalpinioideae)
By S. Bandyopadhyay and K. Thothathri 115
35. SEM studies on the testa surface pattern of some species of Bauhinia (Leguminosae: Caesalpinioideae)
By S. Bandyopadhyay and
K. Thothathri 116
36. On the occurrence of Gentiana infelix Clarke (Gentianaceae) in Garhwal Himalayas
By D.S. Rawat and R.D. Gaur 1 1 8
37. SEM studies on the testa surface pattern of two species of Bauhinia (Leguminosae:Caesalpinioideae) By S. Bandyopadhyay
and K. Thothathri 120
38. Kaempferia siphonantha King ex Baker (Zingiberaceae) in the Andaman Islands
By P.S.N. Rao and B.K. Sinha 121
39. On the occurrence of Dimeria kanjirapalliana K.C. Jacob (Poaceae) in Andhra Pradesh
By M.S. Gayathri and T. Pullaiah 122
40. Two new records of orchids for Andaman Islands By L. N. Ray, P.V. Sreekumar and
PM. Padhye 123
ANNUAL REPORT OF THE BOMBAY NATURAL HISTORY SOCIETY 1 26
STATEMENT OF ACCOUNTS OF THE BOMBAY NATURAL HISTORY SOCIETY 143
MINUTES OF THE ANNUAL GENERAL MEETING
161
JOURNAL
OF THE
BOMBAY NATURAL HISTORY
SOCIETY
April 1996 Vol. 93 No. 1
NIGHT ROOSTING AND ‘LUNAR PHOBIA’ IN INDIAN FALSE VAMPIRE BAT
MEGADERMA LYRA 1
R. SUBBARAJ2 AND J. BALASINGH3 (With seven text-figures )
Key words: Megadenna lyra, moon light intensity, night roosts, foraging, lunar phobia
‘Night roosts’ of Indian false vampire bats Megadenna lyra include cow-sheds, unoccupied buildings, ranging from small store rooms to large country houses. Observations of adult females occupying the night roosts suggest that the duration of occupancy of night roosts and the duration of foraging bouts vary depending on the phases of the moon and the reproductive conditions. Moon light avoidance (reflected by the duration of occupancy of night roosts) is significantly higher during breeding season than during non-breeding season in females. The behaviour of ‘lunar phobia’ in male M. lyra follows the patterns exhibited by the females. It is possible that in M. lyra ‘lunar phobia’ is probably an adaptation for reducing losses to nocturnal predators that are at least visually oriented.
Introduction
“Night roosts” of bats include places used to ingest food transported from nearby feeding areas, used by “sit and wait predators” and calling roosts as part of leks. They may also serve as centres for information transfer about the location of food patches and facilitate social interaction (See review Kunz 1982). Bats roost for short intervals in the night roosts to consume prey that they have captured in flight or on the ground. This behaviour seems most common in those bat species that
'Accepted May 1994.
department of Animal Behaviour and Physiology,
School of Biological Sciences, Madurai Kamaraj University, Madurai-625 021.
department of Zoology, St. John’s College,
Palayamkottai, Tirunelveli-627 002.
take relatively larger prey. For example the intermittent returns and departures of Antrozous pallidus at night roost (Orr 1954, Beck and Rudd 1960, O’Shea and Vaughan 1977) commonly involve the transport of large insects. The selection and the duration of occupancy of night roosts may be influenced directly or indirectly by lunar periodicity. For instance, some desert bats apparently use more protected shelters during brighter lunar periods than during darker ones (Hirshfeld el al. 1977)
We have gathered data concerning the foraging behaviour of M. lyra from radio-tracking studies (Doris et al. 1991). The foraging bouts of these bats in relation to different phases of the moon. In this present study the inhibitory effects of moon light on the foraging is related to the “night roosting” behaviour in these bats.
2
JOURNAL BOMBAY NATURAL HIST. SOCIETY. Vol. 93(1996)
TIME ( h )
Fig. 1 . Foraging activity of tagged M. lyra relative to the phase of the moon. Each horizontal solid line ( )
represents the time spent in the “night roost” by a single female. Dotted lines ( ) cover those hours of the
night when the moon is either set or not yet risen.
The time spent in the night roosts ( ) by M. lyra is plotted subsequently just above ( ) “dark hours”
of the night.
Abbreviations : NM — New moon; IQ — First Quarter; FM — Full moon; LQ — Last Quarter.
Materials and methods
The night roosts of M. lyra were mostly located 50 m to 500 m away from the diurnal temple roost (Tirunelveli, lat. 8° 44' N; long. 77°42' E, South India). They include cow-sheds, unoccupied buildings, ranging from a small store to large country houses. Bats were banded with plastic collars fitted with beads of different colours and combinations to enable individual identification of bats while night roosting (Balasingh et al. 1992). Weekly visits covering 36 nights representing all the lunar phases were devoted completely from dusk to dawn for observing banded M. lyra at night roosts. Interestingly one banded female M. lyra night roosting continuously in an unoccupied house was observed for 12 nights during the breeding season (February, March and April 1989) and for 12 nights during non-breeding season (September, October, and November 1 989). Another banded maleM. lyra, night roosting in a small two chambered temple was
observed for 12 nights during the months of February, March and April 1989. Bats were observed from a distance of less than 5 m with a red filtered lamp or night viewing device (Litton Precision Noctovision Sniperscope). The time spent by the bat in the night roost was recorded with a stop-watch. Timings of moon rise and moon set were obtained from the tables of Ephemeris Nautical Almanac published by the Director of Observatories, Calcutta and were adjusted for longitude, latitude and Indian Standard Time (1ST).
Results
M. lyra night roosted singly and not in groups. Night roosts tend to be favoured places situated 50 m to 2 Km from the day roost. During breeding seasons most of the lactating females night roosted 50 m to 500 m from the day roost. During rainy months most of the night roosts were closer to day roost while during the long summer, these night
Time (h)
NIGHT ROOSTING AND 'LUNAR PHOBIA' IN MEGADERMA LYRA
3
Fig. 2. Time duration of night roost occupancy by female M. lyra (ordinate) in relation to different phases
of moon.
NM — New moon; IQ — First Quarter;
FM — Full moon; LQ — Last Quarter.
I Breeding; | | Non-breeding.
roosts were temporarily vacated by the bats.
An adult female bat continuously occupied the night roost for more than 6 months. Variations in the patterns of fdraging by this single female M. lyra were correlated with the phases of the moon. During new moon nights the bats engaged in prolonged
Fig. 3. Time duration of night roost occupancy by male M. lyra (ordinate). Other details as in Fig. 2.
foraging bouts and as a result the duration of occupancy in the night roosts were greatly reduced. During first half of the “bright moon” nights (first quarter moon) the bat suspended foraging activity and prolonged the stay in the night roost (Fig. 1). From the first quarter to full moon, despite the presence of a relatively bright moon at sunset, the M. lyra female left the day roost at the usual time but stayed away long enough to complete one or two
4
JOURNAL BOMBAY NATURAL HIST. SOCIETY, Vol. 93 (1996)
Figure 4 illustrates the duration of stay (I bout) in the night roost by the female. The time spent in the night roost was relatively higher during “bright moon” hours of the first quarter moon and full moon nights. The results are also in agreement with the data collected for subsequent bouts in the night roosts (Figs. 5, 6 & 7). Even though there were variations in the time spent during different bouts in the night roost by the female during breeding and non- breeding seasons, the results of cumulative data collected for all the rest of the bouts suggest that during breeding season the moon light avoidance is significantly higher than during non-breeding season.
Discussion
Fig. 4. Time duration ol 1 bout night roost occupancy (ordinate) in breeding (|g ) and non-breeding (Q] ) female. Other details as in Fig. 2
feeding passes before returning to the night roost.
During full moon nights, even though there was an occurrence of two or three feeding bouts they were of shorter duration. The duration of occupancy in the night roosts was significantly higher.
During most of the nights, at moon-set this female M. lyra re-emerged from the night roost and foraged till dawn. These data on the influence of moon phases on the foraging activity of the female M. lyra differs significantly during breeding and non- breeding seasons. Fig. 2 shows clearly that the time spent in the night roost by the female M. lyra was significantly higher during breeding season compared to the non-breeding season during all phases of the moon. During last quarter, however, the female spent significantly lesser time in the night roost during non-breeding season than in the breeding season (Fig. 2).
Figure 3 illustrates the dependency of the duration of the night roost occupancy in an individual male on the lunar periodicity. The pattern follows those exhibited by the female. During “bright moon” hours the male returned to the night roost and remained there for a long time.
Bats are “faithful” to individual night roost as long as the nearby area remains resourceful. During rainy months most of the night roosts are closer to day roosts since the ponds and the neighbouring fields are flooded with water yielding rich food resources. During long summer months bats make long commuting flights between foraging grounds and night roosts, hence several of the night roosts were temporarily shifted to distant places which are rich food resources.
The lactating females “night roosted” close to day roost during breeding season because the mother bats carried their young to the night roost and left them in the night roost while they foraged. Since they have to carry the extra baggage the mother bats preferred to night roost closer to the day roost during breeding.
The first indications of moon light avoidance behaviour were given in the observations of Tamsitt and Valdi vieso (1961), Villa ( 1 966), Wimsatt ( 1 969), Schmidt et al. (1971) and Crespo et al. ( 1972) who could catch less number of foraging vampire bats and phyllostomids in their nets at moonlit nights than they could before the moon had risen or after it had set. A direct proof of the inhibitory effects of moon light was, however, obtained only after months of recording the flight activity of several captive bats such as Artibeus jamaicensis and Phyllostomus discolor under natural lighting conditions (Erkert
NIGHT ROOSTING AND ‘LUNAR PHOBIA ’ IN MEGADERMA LYRA
5
CD
E 3
2 a ^ a
Fig. 5. Time duration of II bout night roost occupancy (ordinate) in breeding (Hi ) and non-breeding (| | )
female. Other details as in Fig. 2.
1974). The findings of the studies have since been confirmed with a number of techniques, including the radio-tracking of A. jamaicensis (Morrison 1978a, b), bat detector recordings of the activity of various microchiropterans (Fenton et al. 1977) and simulation experiments on A. lituratus and P. hastatus in artificial light-darkness cycle in the laboratory (Haussler and Erkert 1978).
In addition, the differential sensitivity of bat activity patterns to moon light could also serve to reduce the direct interspecific competition among bats specialising on particular food resources by temporal separation of foraging activity based on the lunar cycle. Such a mechanism was proposed, for
(ordinate) in breeding ( H ) and non-breeding ( ) female. Other details as in Fig. 2. example, by Owings and Lockard (1971) for two rodent species of Peromyscus with differential responses to moon light intensities.
The moon light avoidance behaviour of M. lyra is not cued simply to ambient light level. M. lyra left the day roost after sunset even on nights when a bright moon was already present. At sunset, hunger may be an over-riding factor, causing the bats to emerge for a short bout of feeding despite the illumination from the full moon. Furthermore, the data of Lockard (1978) from field work measuring activity of kangaroo rats throughout the full range of naturally occurring conditions do clearly show that light intensity alone is not the cue, for much
6
JOURNAL BOMBAY NATURAL HIST. SOCIETY, Vol. 93(1996)
Fig. 7. Time duration of IV bout night roost occupancy (ordinate) in breeding (m ) and non-breeding ([^] ) female. Other details as in Fig. 2.
activity also occurred about the time of civil twilight (sun 6° below the horizon) when the illumination was about 10 lx. Whereas the full moon was on the order of 0.5 lx. Later, when there was much less illumination but a luminous disk was in the sky, activity was inhibited. Thus any of the three following hypotheses could account for the observed behaviour in M. lyra :
(1) Activity is inhibited by any reasonably conspicuous luminous disk in the sky.
(2) Activity is inhibited by a luminous disk in
the sky that provided the ambient illumination which is above some threshold point.
(3) Activity is inhibited by an endogenous clock running on lunar time.
. The limited data available in the present study permitted to analyse only the influence of the lunar periodicity on the night roosting behaviour in A/, lyra. In this context, it is of interest that moon Ph ases influence the activity of several bats directly or indirectly acting through changes in behaviour, abundance or availability of prey (Turner 1975).
We do not have complete data on the prey abundance at different moon phases over the seasons. However, it is possible that in M. lyra similar to other bat species “lunar-phobia” is probably an adaptation for reducing losses to nocturnal predators that are at least visually oriented (Lockard and Owings 1974, Lockard 1978, Morrison 1978a; Barclay 1985a, b; Fleming and Heithaus 1986).
Acknowledgement
We thank Prof. M.K. Chandrashekaran for critically reading the manuscript.
References
Balasingh, J., S. Suthakar Isaac & R. Subbaraj (1992): A convenient device for tagging bats in the field. Bat Research News. 33: 6.
Barclay, R.M.R. (1985a): Long-versus short-range foraging strategies of hoary ( Lasiurus cine reus ) and silver-haired (Lasionycteris noctivagans) bats and the consequences for prey-selection. Can. J. Zool. 63: 2507-2515.
Barclay, R.M.R. (1985b): The echolocation calls of hoary ( Lasiurus cinereas) and silver-haired ( Lasionycteris noctivagans) bats and adaptations for long-versus short-range foraging strategies and the consequences for prey-selection. Can. J. Zool. 64: 2700-2705.
Beck, A.J. & R.L. Rudd (1960): Nursery colonies in the pallid bat. J. Mammal. 41: 266-267 .
Crespo, R.L., S.B. Linhart, R.J. Burns & G.C. Mitchell (1972): Foraging behaviour of the common vampire bat related to moon light. J. Mammal. 53: 366-368.
Doris, A., G. Krull, G. Marimuthu, S. Sumithran & J. Balasingh (1991): Foraging behaviour of the Indian false vampire bat Megaderma lyra (Chiroptera: Megadermatidae). Biotropica
23: 63-67.
Erkert, H.G. (1974): Der Einfluss des Mondlichtes auf die Aktivitaetsperiodik nachtaktiver Saugetiere. Oecologia 14: 269-287.
Fenton, M.B., N.G.H. Boyle, T.M. Harrison & D.J. Oxley (1977): Activity patterns, habitat use and prey selection by some African insectivorous bats. Biotropica 9: 2860-2867.
Fleming, T.H. & E.R. Heithaus (1986): Seasonal foraging behaviour of the frugivorous bat Carollis perspicillata. J. Mammal. 67: 660-671.
Haussler, U. &H. Erkert (1978): Different direct effects of light intensity on the entrained activity rhythm in Neotropical bats (Chiroptera: Phyllostomidae). Behav. Process. 3: 223-239.
Hirshfeld, J.R., Z.C. Nelson & W.G. Bradley (1977): Night roosting behaviour in four species of desert bats. South West. Nat. 22: 427-433.
Kunz, T.H. (1982): Roosting ecology of bats. pp. 1-55, In: Ecology of bats (T.H. Kunz, ed.), Plenum Press, New York. 425 pp.
Lockard, R.B. (1978): Seasonal changes in the activity pattern
NIGHT ROOSTING AND LUNAR PHOBIA’ IN MEGADERMA LYRA
7
of Dipodomys spectabilis. J. Mammal. 59: 563-568. Lockard, R.B. & D.H. Owings (1974): Moon-related surface activity of bannertail ( Dipodomys spectabilis) and fresno (D. nitratoides ) kangaroo rats. Anim. Beliav. 22: 262-273. Morrison, D.W. (1978a): Lunar phobia in neotropical fruit bat, Artibeus jamaicensis (Chiroptera: Phyllostomatidae). Anim. Behav. 26: 852-856.
Morrison, D.W. (1978b): Influence of habitat on the foraging distances of the fruit bat, Artibeus jamaicensis. J. Mammal. 59: 622-624.
Orr, R.T. (1954): Natural history of the pallid bat, Antrozous pallidas (Le conte). Proc. Calif. Acad. Sci. 28: 165-246. O’shea, T.J. & T.A. Vaughan (1977): Nocturnal and seasonal activities of the pallid bat, Antrozous pallidus. J. Mammal. 58: 269-284.
Owings, J.F. & R.B. Lockard (1971): Different nocturnal activity
patterns of Peromyscus californicus and Peromyscus eremicus in lunar lighting. Psychonom. Sci. 22: 63-64.
Schmidt, U., A.M. Greenhall & W. Lopez-Forment (1971): Okologische Untersuchngen der Vampirfledermause (Desmodus rotundas ) in State Puebla, Mexiko. Z. Saugetierkd. 36: 360-370.
Tamsitt, J.R., & D. Valdivieso (1961): Notas sobre activiades nocturnas y estados de reproduccion de algunos Quiropteros de Costa Rica. Rev. Biol. Trap. 9: 219-225.
Turner, D.C. (1975): The vampire bat. Johns Hopkins University Press, Baltimore and London. 145 pp.
Villa, R.B. (1966): Los murcielagos de Mexico. Inst. Biol. Univ. Nac. Auto. Mexico City. pp. 491.
Wimsatt, W.A. (1969): Some problems of reproduction in rela- tion to hibernation in bats. Bull. Mus. Comp. Zool. 124: 249- 267.
RELATIONSHIP BETWEEN DIVE AND POST-DIVE PAUSE WHILE FORAGING IN
TWO DIVING DUCKS OF LAKE MANSAR1
Y.R. Malhotra2, N. Deskyong3, P.S. Pathania3
( With three text-figures)
Key Words: dive, pause, ducks, relationship, diving
Relation between dive and post-dive pause while foraging is quantified in the two diving duck species: Common pochard (Aythya ferina) and Tufted duck (A fuligula) that winter in Lake Mansar, Distt. Udhampur, J&K. A total of 1641 dive cycles (98 1 dive cycles for Tufted duck and 660 for Common pochard) were observed during the winter 1992-93. In both the species positive relationship between dive time and post-dive pause has been analysed by Karl Pearson’s Co-efficient of correlation (r) method. However, such positive relationship is more in Common pochard (r=0.88; t>0.05) than in Tufted duck (r=0.65; t>0.05). This has been correlated with the diet difference of the two ducks. Common pochard which is largely a vegetarian shows increase in pause time with dive time as compared to Tufted duck which feeds on sessile or slow moving benthic prey.
Long dives (21 sec. and above) are observed more in Tufted duck than Common pochard. This difference in dive time is influenced by diet difference and foraging decisions made while underwater.
Introduction
Foraging style of diving birds, that dive from surface of water and after spending some time underwater, returns to the water surface to breathe is well known (e.g., Johnsgard 1965, Wallace and Mahan 1975, Ali and Ripley 1978, Lessells and Stephens 1983, Ydenberg 1986, Woakes and Butler 1986). Similarly considerable literature on physiology of diving is also available. Butler and Jones (1982) in their review “Comparative physiology of diving in Vertebrates” listed almost one thousand references. In contrast, publications of diving behaviour and particularly on the relationship between dive and pause is very scarce.
Diving behaviour of birds hold great fascination not only among ornithologists but also among naturalists and this can be summed up from a Scottish physician and naturalist J.M. Dewar’s (1924) statement: “Among the problems surrounding the life of birds, none is more fascinating than the underwater activities of diving fowl”.
'Accepted June 1993.
-Vice-Chancellor, University of Jammu, Jammu (J&K)-1 80 004.
3Post-Graduate Department of Bio-Sciences,
University of Jammu, Jammu (J&K)-180 004.
In this paper, we describe briefly some of our studies on the comparative relationship between dive length and post-dive pause when freely foraging, in the two Aythya species, i.e. Tufted duck A. fuligula and common pochard A. ferina, that winter in lake Mansar. The starting point is the common place observation that although some dives by birds are made during courtship or to escape predators, most are made to capture food.
Study Area
Lake Mansar (32°42* N and 75° E) is a heart shaped sub-oval water body, 65 km to the east of Jammu city (J&K), and is located at an elevation of 710 m above msl. The lake is 37 metres deep at the centre, and has a circumference of 3.294 Km. It is primarily fed by surface run off, and has some submerged spring sources. It is classified as a fault basin, non-drainage, type of lake without any distinct regular inflow or outflow channel.
The lake is utilized by a number of migratory aquatic birds in winter.
Methods
This study was conducted during the winter of 1992-93 in a lake Mansar. Birds were watched
RELATIONSHIP BETWEEN DIVE AND POST-DIVE PA USE IN DIVING DUCKS
9
and observation recorded for the duration of their sequential dives and surface pause after re- emergence using a stop-watch. Total of 1641 dive cycles (98 1 for Tufted duck and 660 for Common Pochard) were observed. Only one bird (of either of the two diving duck species) was observed at a time. During the present work only those dives were taken into consideration which were made for foraging. Divers make repeated foraging excursions from the surface to which they must return to breathe (Ydenberg 1986), though some distance away from where they dive. Thus the underwater time and the post-dive pause which is spent on the surface, completes a dive cycle. During a dive cycle, the bird under observation may get disturbed, by one way or the other on quite a few occasions, particularly during emergence at surface, thus either lengthening the post-dive pause or forcing the bird to dive. All such observations were deleted from the data so that we have a data set purely of free foraging dive cycles.
To compare the post-dive pause time to its preceding dive length time, all the observations were grouped that were made under similar dive length time (in seconds) and then the mean ± S.D. of all the post-dive pauses were calculated for each group of similar dive length (Table 1).
To analyse any correlation between dive length and post-dive pause, statistical method of Karl Pearson’s co-efficient of correlation (r) was applied to the data-
r= Ixy
v Xx2 x ly2
Where x = (X-X) and y = (Y-Y)
X is the dive length,
Y is the mean post-dive pause time,
X is the mean dive length of the data,
Y is the mean pause time of the data,
To test the significance of the observed Co- efficient of Correlation (r), t-test has been applied as follows:
t = X
Where ‘n’ is the d.f.
Regression analysis has also been worked out to estimate values of pauses (Y) at independent values of dive (X).
Regression equation of Y on X is expressed as follows:
Y = a + bX
X (dive) is the independent variable and ka’ and ‘b’ are constant having values of
a=7.17 and b= 0.26 in Tufted duck. a=3.6 and b= 0.49 in Common pochard.
Result and Discussion Table 1 and Fig. 1 summarizes comparative data on dive length and post-dive pause relation-
Table 1
COMPARATIVE ACCOUNT OF POST-DIVE PAUSE LENGTH TO ITS PRECEDING DIVE LENGTH IN THE TWO DIVING DUCK SPECIES IN LAKE MANSAR
|
Dive time (in seconds) |
Pause time in Tufted duck Mean ± S.D |
Pause time in Common pochard Mean ± S.D |
|
6 |
7.33 ±2.51 |
|
|
7 |
6.66 ± 2.08 |
|
|
8 |
7.66 ± 2.08 |
|
|
9 |
7.00 ±3.82 |
|
|
10 |
8.75 ± 3.59 |
|
|
11 |
9.00 ± 0.00 |
10.25 ±4.99 |
|
12 |
10.00 ±0.00 |
13.33 ±4.04 |
|
13 |
— |
10.50 ±4.94 |
|
14 |
9.50 ± 1.00 |
11.00 ±4.21 |
|
15 |
12.75 ±2.62 |
12.33 ±4.21 |
|
16 |
12.50 ±3.80 |
10.00 ± 1.41 |
|
17 |
9.54 ± 2.25 |
10.12 ±5.20 |
|
18 |
12.40 ±4.30 |
12.80 ±2.28 |
|
19 |
14.98 ±4.97 |
12.50 ±3.50 |
|
20 |
15.78 ±7.62 |
15.11 ± 1.83 |
|
21 |
13.53 ±6.77 |
13.22 ± 1.39 |
|
22 |
12.03 ±3.31 |
14.84 ±2.70 |
|
23 |
12.90 ±2.25 |
17.14 ±4.52 |
|
24 |
15.59 ±5.30 |
14.00 ±3.81 |
|
25 |
13.27 ±4.09 |
14.33 ±4.16 |
|
26 |
13.23 ±4.62 |
13.60 ±3.13 |
|
27 |
13.90 ±3.58 |
14. 10 ±3.69 |
|
28 |
13.36 ±3.86 |
15.00 ±4.69 |
|
29 |
11.35 ±2.89 |
15.80 ± 1.78 |
|
30 |
14.18 ±4.35 |
22.16 ±2.40 |
|
31 |
19.75 ±4.11 |
25.00 ± 6.85 |
10
JOURNAL BOMBAY NATURAL HIST. SOCIETY, Vol. 93 (1996)
ship in the two diving ducks Aythyafuligula and A. ferina in Lake Mansar. In both the species positive relationship has been worked out, i.e. r=0.88 in Common pochard and r=0.65 in Tufted duck, both of which are significant at 5% level (i.e. t > 0.05).
Fig. 1. Dive-pause relationship in the two diving ducks: Cp — Common pochard and Td — Tufted duck. Each point is the mean of 10 to 132 dive cycles.
Similar to our observation, Dewar (1924) has also shown positive relationship with significant increase in pause time with dive time in diving ducks and other divers (including loons, grebes, cormorants and auks). Positive relationship between dive and pause duration has also been worked out by Forbes (1985) in Western Grebe Aecmophorus occidentalis, and Stonehouse (1967) in cormorants Phalacrocorax melanoleucos and P.carbo. Houston and McNamara (1985) developed a general theory of central place foraging for single prey loaders that takes account of the fact that longer dives are followed by longer pauses. Data from a variety of species show that pause duration is directly related to the length of the preceding dive which strongly suggests that this is, at least, partly, recovery time during which respiratory gases are exchanged (Butler and Woakes 1979). Comparative data on dive and surface times suggest that different species, depending on their foraging ecology, allocate the recovery time from
dives in different ways, such as, divers whose prey may escape or hide between dives (e.g. fish) may postpone recovery to dive more frequently until a series of dives has been completed. Divers whose prey are sedentary (e.g. shellfish) seem to complete much more of the recovery after each dive (Ydenberg 1986). Diving is energy expensive (Woakes and But(er 1983, 1986) and divers spend much time underwater. Time spent on surface is used for recovery from physiological effects (partial asphyxiation) which may be a consequence of diving and underwater life as also suggested by Ydenberg ( 1 986) and Ydenberg and Forbes (1988). Alongside, body heat lost while underwater may be regained while on the surface after a dive as also suggested by Mac- Arthur (1984).
DIVE LENGTH (seconds)
Fig. 2. A comparison of the dive-pause relationship in Common pochard (Cp) and Tufted duck (Td) by Regression analysis.
Of the two dfving ducks, Common pochard shows more increase in pause time with increase in dive time than in Tufted duck, i.e. regression line for pause time is steeper in case of Common pochard than in Tufted duck (Fig. 2). This difference can be related to the difference in diet of the two species.
RELATIONSHIP BETWEEN DIVE AND POST-DIVE PA USE IN DIVING DUCKS
DIVE LENGTH (seconds)
Fig. 3 Percentage of dives made at a frequency of
That Common pochard is more a vegetarian than the Tufted duck, is well documented by Ali and Ripley (1978). Ydenberg (1986) also explains that difference in observation of Dewar (1924) where steepest slope is for diving ducks as compared to other divers like mergansers, loons, grebes, cormorants, etc., is suggested to be due to different diets of the latter from former. The diving ducks generally feed on sessile or slow moving benthic prey such as shellfish and crustaceans (Nilsson 1972, Pehrsson 1976, Ali and Ripley 1978) whereas other species (mergansers, loons, grebes, cormorants) which capture fish in active pursuit. Ydenberg (1986) explains that short pauses are advanta- geous because prey may escape between successive dives.
When dive time of the two ducks is compared for same time (2 1 seconds and above), we find Tufted duck diving for this duration on 77.5% of the total
DIVE LENGTH (seconds)
5 seconds in (A) Tufted duck; (B) Common pochard.
observation (Fig. 3, A), whereas Common pochard does so for 52% of the total observations only (Fig. 3, B). The difference in dive duration is influenced by diet difference in the two species as already stated and also by the foraging decisions (Stephens and Krebs 1986) made while underwater, namely which prey to eat and which to neglect or how many prey to capture and how long to continue a search before surfacing. Houston and McNamara (1985) concluded that, for a diving bird, the decision policy, for accepting and rejecting prey items, is in favour of that which maximizes the rate of energy gain which makes the bird less and less selective as the dive progresses, because rejection becomes more and more costly.
Conclusion
Data collected and analysed thus shows positive relationship between dives and post-dive
12
JOURNAL, BOMBAY NATURAL HIST. SOCIETY, Vol. 93 (1996)
pause, i.e. with increase in dive time, post-dive pause also increases. Such relationship in dive-pause is well supported by reports of other investigators also. Dive-pause relationship is more in Common pochard than in Tufted duck and is well explained by the difference in diets of the two species. Further it is observed that on an average Tufted ducks go for longer duration of dives and on more occasions than Common pochard do, which can be related to
their differences in diet and foraging decisions made while underwater.
Acknowledgements
This work was carried out at the Post-graduate Department of Bio-Sciences, University of Jammu. We are grateful to the staff of the Department and especially to Prof. O.P. Sharma for providing good working facilities.
References
Ali, SAlim & S.D. Ripley (1978): Handbook of the birds of India and Pakistan. Vol. I. Oxford University Press, London, New York.
Butler, P.J. & D.R. Jones (1982): Comparative physiology of diving in vertebrates. In: Advances in physiology and biochemistry. Vol. 8. D.E. Lowenslein (ed.). Academic Press, New York, pp. 179-364.
Butler, P.J. & A.J. Woakes (1979): Changes in heart rate and respiratory frequency during natural behaviour of ducks, with particular reference to diving. ./. Exp. Biol. 79: 283-300.
Dewar, J.M. (1924): The bird as a diver. H.F. & G. Witherby, London. 173 pp.
Forbes, L.S. (1985): The feeding ecology of Western Grebes breeding at Duck Lake, British Columbia. Unpublished M.Sc. thesis, Univ. of Manitoba, Winnipeg. 72 pp.
Houston, A.l. & J.M. McNamara (1985): A general theory of central place foraging for single-prey loaders. Them: Popul. Biol. 28: 233-262.
Johnsgard, P.A. (1965): Handbook of waterfowl behaviour. Cornell Univ. press. New York.
Lessells, C.M. & D.W. Stephens (1983): Central place foraging: Single prey loaders again. Anim. Behav. 31: 238-243.
Mac-Artiiur, R.A. (1984): Aquatic Thermoregulation in the Muskrat ( Ondatra zibethicus): Energy demands of swimming
and diving. Can. J. Zool. 62: 241-248.
Nilsson, L. (1969): Knipans Buceplwla clanyula beteende under Vinterhalvaret. Var Fagelvarld 28: 199-210.
Pehrsson, O. (1976): Food and feeding grounds of the golden eye Buceplwla clausula (L.), on the Swedish Coast. Ornis Scan. 7: 91-112.
Stephens, D.W. & J.R. Krebs (1986): Foraging theory. Princeton Univ. Press, Princeton.
Stonehouse, B. (1967): Feeding behaviour and diving rhythms of some New Zealand Shags, Phalacrocoracidae. Ibis 109: 600-605.
Wallace, G.J. & H.D. Mahan (1975): An introduction to Ornithology. MacMillan Pub. Co., Inc., New York. Collier MacMillan Publishers, London.
Woakes, A.J. & P.J. Butler (1983): Swimming and diving in Tufted ducks, A ythya fuligula with particular reference to heart rate and gas exchange. J. Exp. Biol. 107: 311-329.
Woakes, A.J. & P.J. Butler (1985): Respiratory, circulatory and metabolic adjustments during swimming in the Tufted Duck, Aythya fuligula. J. Exp. Biol. 120: 215-218.
Ydenberg, R.C. (1986): Foraging by diving ducks. Congressus internationalis ornithologici (19th: 1986: Ottawa, Ont.).
Ydenberg, R.C. & L.S. Forbes (1988): Diving and foraging in the Western Grebe. Ornis. Scand. 19: 129-133.
NOTES ON THE FEEDING AND BREEDING BEHAVIOUR OF GYMNOPLEURUS GEMMATES HAROLD AND GYMNOPLEURUS MILIARIS (F.) (COLEOPTERA: SCARAB AEIDAE) 1
K. Vfjenakumari2 and G.K. Veeresh3 (With a text-f igure)
Key Words: Gymnopleurus gemmcitus, G. miliaris, nesting, competition
Field studies on the feeding, mating and competitive behaviour of Gymnopleurus gemmatus and G. miliaris were conducted in Bangalore. Both the species were diurnal and fed both at the pat and on dung balls that they fashioned, transported and buried before feeding. Competition was intense both for dung balls and mates within the species and for dung balls alone between the species. Species belonging to the genera Onthophagus and Caccobius were found frequently as kleptoparasites in the brood balls of these beetles.
Introduction Material and Methods
Gymnopleurus , a dung rolling coprophagous genus of beetle, is widely distributed in Asia, Europe and Africa (Arrow 1931). The fashioning and transportation (by rolling) of dung balls, by these beetles not only reduces congestion at the resource site but could also give the rollers a competitive edge over the dung burying groups like the Coprini, Onthophagini, Onitini, etc., which compete for food and burial space beneath and around each dung pat (Halffter and Matthews 1966).
It was Fabre (1897), the famed french naturalist, who made the first systematic studies of the three dung rolling genera Sccirabaeus, Gymnopleurus and Sisyphus. Subsequently, several studies on the ball making, rolling and burial behaviour of Gymnopleurus have been made (Hingston 1923, Honda 1927, Prasse 1957a, 1957b, 1957c, 1958a and 1958b).
In India, however, after Hingston (1923), the behaviour of these beetles (in particular Gymnopleurus ), has gone largely unnoticed. A study on the field behaviour of two commonly occurring species of Gymnopleurus , namely G. miliaris and G. gemmatus was therefore undertaken.
‘Accepted February 1994.
2P.B. No. 43 1. Junglighat, Port Blair, Andaman Islands, 744103. 'Vice Chancellor, University of Agricultural Sciences, Bangalore
560065.
The feeding and breeding behaviour of G. miliaris and G. gemmatus were studied in grazing fields at two locations (Hebbal and Allalsandra) on the outskirts of Bangalore (12° N lat. and 77° E long., 916 m alt.) in S. India. The study sites are situated at about 7 and 1 1 km north of Bangalore, respectively. The rainy season which commences here in late April continues till the end of September during which period these beetles are active. During the period of study (1984-1986) the mean maximum and mean minimum temperatures were 29.8° C and 18.2° C while the total annual rainfall amounted to 548.3 mm.
Observations were made on the following elements of beetle behaviour, namely a) approaching food, b) feeding, c) ball making, d) ball rolling, e) mating, f) intra- and inter-specific competition, and g) kleptoparasitism.
The rollers, G. miliaris and G. gemmatus were identified by Dr. R. Madge of the British Museum (Natural History), London.
Results and Discussion
The beetles commenced activity after the first rains in late April. They are diurnal, with their period of activity usually extending from 0700 to 1 830 hrs. Light showers did not make them cease activity.
Approach to food: The beetles always located
14
JOURNAL BOMBAY NATURAL HIST. SOCIETY, Vol. 93 (1996)
their food by flying low, in a zigzag manner and alighting at a mean distance of 8.65 cm (G. gemniatus) and 8.85 cm (G. miliaris) from the food source. Whenever they landed on their backs they used their mid pair of legs as pivots to right themselves. A few alighted directly on the dung pat. Having detected the presence of food, the beetles, with antennae waving feverishly, walked briskly towards it. On reaching the food source, they walked all over it to finally select a suitable spot to commence feeding.
Feeding: Both the species of Gymnopleurus were attracted to human faeces, sheep excrement and cow dung. It was visually apparent that the beetles preferred human faeces of the three sources of excrement.
The beetles commenced feeding at the dropping after having selected a suitable spot on. the dung mass. When feeding on sheep-pellets, it was noticed that the beetles fed exclusively in the inner core by boring into the pellet. This brief period of feeding was followed by ball making.
Balls of excrement were fashioned for various purposes, namely i) for food, ii) as nuptial gifts, and iii) for rearing brood.
Both the species of Gymnopleurus made food balls, rolled, buried and fed on them unlike G. mopsus Pallas andG. geojfroyi Fuessly (Prasse 1957a, 1957b, 1958a) which never made any food balls. Hingston (1923) also reports food ball preparation in G. miliaris.
Prasse (1957a) has reported that species of Gymnopleurus undergo a ‘Reifungsfrass’ period of 3-3.5 months. In the present study the beetles which emerged with the onset of the rains were found mating on the first day and brood balls were made a week after. So the Reifungsfrass period is definitely shorter for these two species of Gymnopleurus.
Ball making: After selecting a suitable spot on the faecal mass the beetles carved out circular segments of dung from the rest of the mass using their front tibiae and clypeus. During the process of fashioning, bits of dung were added from the main mass whenever the initial mass was found to be inadequate.
The sphere was finally detached from the main mass by the beetle moving down the side of the ball to its point of attachment at the base. The beetle then pushed against the ball with its middle and hind pairs of legs till the ball got detached. It was then rolled to the edge of the pat.
Perched on the ball they patted it into a smooth sphere using their fore tibiae. Once again, any deficiency in the quantum of dung making up the sphere was made up by adding from the main mass and if in excess, the extra material was cut away with the clypeus and discarded. Each beetle took about 11.07 ± 1.03 min. (G. gemniatus) and 10.10 ± 1.23 min. (G. miliaris) to construct and fashion one ball. In those cases where the partner was chosen during the period of ball construction, the new partner also helped in ball making.
When sheep pellets were used as raw material for making balls, a different technique was adopted. The beetles then broke open a number of sheep pellets to collect sufficient material from the soft inner core. These were then used sequentially in the fashioning of a ball of requisite proportions.
Ball making was observed at temperatures between 23.3°C and 29.4°C.
Ball rolling: The belli of dung once detached was rolled away from the dung pat to a distance of about 10-20 cm, where the beetle was generally noticed to finally shape the ball. Sitting on the ball, the beetle patted the ball into shape using its forelegs, as well as by intermittently pressing the ball with its clypeus. On completion of this final fashioning of the ball into a compact spheroid the beetle began rolling the ball. The average diameter and weight of the balls rolled by a lone beetle was 1.0 ± 0.13 cm and 0.34 ± 0. 1 1 g (G. gemniatus), 1 .03 ± 0. 1 cm and 0.36 ± 0.09 g (G. miliaris). On the other hand, when pairs rolled the ball, it was 1 1 . 1 5 ± 0. 1 1 cm and 0.36 ± 0.06 g (G. gemniatus), 1 . 1 3 ± 0.3 1 cm and 0.48 ± 0.03 g (G. miliaris).
As the ball was rolled along, it acquired a thin coating of soil. If a single beetle was engaged in rolling, it pushed the ball with its middle and hind pairs of legs while it had its forelegs on the ground. In 90% of the cases it was the male that made the
FEEDING AND BREEDING BEHAVIOUR OF, GYMNOPLEURUS GEMMATUS AND G. M1LIARIS
15
ball; to be later joined by the female. As Thornhill (1983) suggests, the “material benefits” — here the dung ball — are provided by males probably to reduce the loss of energy in the female in making a ball as she has to spend a lot of energy on other activities like — rolling, burying and brood ball making, egg laying, etc.
However, it was noticed that the maker of the ball, irrespective of its sex, did not readily accept as a partner the first individual that came up to the ball. If a male was the ball maker it fought and chased away all approaching males; while it accepted an approaching female, only after a brief combat of about 1 5-20 seconds. If the ball maker was a female it too chased away all females but accepted a male after a brief combat. To ascertain the sexual identity of each rolling pair, over 50 pairs of both G. gemmatus and G. miliaris were dissected. Almost every pair, it was found, consisted of a male and a female. In only two instances members of the same sex were found rolling the ball.
This was the result of intra-sexual combat in which individuals of the same sex fought each other in a bid to steal and gain possession of the ball. In fact, females of both the species were rolling a ball. This behaviour has been reported earlier for Sccirabcieus sacer (Fabre 1897) and G. miliaris (Hingston 1 923). In transporting the ball, males and females took up characteristic stances in all cases. The female always stood behind the ball, pushing with the last two pairs of legs, while the male pulled the ball from the front using its last two pairs of legs.
This contradicts Hingston’s ( 1 923) observation that ‘as a rule’ the male pushes the ball while the female pulls it. Observations similar to that in the present study were made by Honda (1927) for G. sinuatus (Ol.) and Prasse (1957b, 1958a) for G. mopsus and G. ge affray i.
During the course of rolling the male was unable to keep pace with the female. He would tumble off but the female would continue rolling, apparently unconcerned, while the male hastened to catch up. On the other hand, in cases where the female fell, the male waited with the ball for her to rejoin him. He would permit her to resume her role
only after a short skirmish lasting 2 to 3 seconds. If, for some reason, the female failed to return, the male abandoned the ball after a period of waiting. In one instance the male waited for 3 min. 40 sec. and in another for 5 min. during which time they met new females with whom they continued the activity of rolling. Unlike males the females continued rolling the ball and buried it even when the males had deserted.
In some cases, the male was found sitting on the side of the ball instead of pulling it, which contradicts Hingston’s (1923) report that this is found only in Scarabaeus sacer and never in G. miliaris. Nevertheless Prasse ( 1 957b) has reported the same for the two species of Gymnapleurus. In such cases the male would get off the ball when the female needed help in surmounting an obstacle.
The beetles never rolled the ball in a straight line. Most were found rolling the ball in a haphazard manner, sometimes crossing the same spot several times and finally burying their balls close to the starting point, even though the balls were rolled for much greater distances than the shortest distance between the food source and the burial site. This once again contradicts Hingston’s ( 1 923) observations that the pellets must be rolled strictly in straight lines.
Whenever the beetles encountered an obstacle they adopted any one of the following three strategies, i) crossed over the obstacle, ii) took a detour, or iii) buried the ball at the base of the obstacle.
The average distance rolled by a pair of beetles was 11.71 m, n = 15 (G. gemmatus ) and 22.22 m, n=15 (G. miliaris). The average distance rolled per minute was 105.10 cm, n=15 (G. gemmatus) and 1 1 9.53 cm, n = 1 5 (G. miliaris). When a single beetle rolled a ball it often stopped rolling, climbed on the ball, checked all around the ball and got back into position to continue rolling the ball. On the other hand when a pair were engaged in ball rolling, the female stopped periodically to inspect the surface of the ball. On encountering the male on the other side she often tried to butt him off, only to recognize him later and then continue rolling. The average time taken for rolling was 1 6 min., n = 1 0 (G. gemmatus)
16
JOURNAL BOMBAY NATURAL HIST. SOCIETY. Vol. 93 (1996)
and 20 min., n = 10 (G. miliaris).
Ball burying: In burying the ball, individuals and pairs adopted different strategies. In the case of pairs, it was always the female who selected the burial spot. She walked away from the ball, examined a certain area and then returned. When individual beetles were involved, they tested and selected sites while holding onto the balls with their hind legs.
Selecting a suitable burial site involved the prior rejection of a number of sites. Four such rejected sites were examined by digging. Strangely, the kleploparasites Onthophag us pygmae us (SchaU.) and
O. centricornis (F.) were found to be present beneath the soil surface.
When single beetles buried balls they came out of their pits several times during the process of digging to check for the presence of the ball. Having made a small pit. the beetle dragged the ball into the pit and disappeared beneath it to continue digging as a result of which the ball disappeared into the soil.
In case of pairs, it was always the female which took to digging while the male stood guard. After making a small pit, she rolled the ball with her hind pair of legs. Sometimes the male helped her in pushing the ball into the pit. During the period of burying the male either sat on the ball or walked around the pit. Sometimes he held the ball with his front two pairs of legs while he stood at the rim of the pit. When 75% of the ball disappeared into the soil, the male also entered the soil to join the female. The average time required for burying the ball was 14.36 ± 4.06 min., n = 10 (G. gemmatus) and 13.45 ± 2.08 min., n = 15 (G. miliaris).
Digging up twenty marked burial sites after a day or two revealed the following.
1 . During the early part of the season, most of the lone beetles used the balls for feeding and only frass and faecal matter were found in the burrow.
2. Later in the season, lone females used the ball for raising brood.
3. Balls rolled by the pairs used them both for enticement and raising brood.
Mating: Both species generally mated in the space between the side of the dung ball and the pit and sometimes above the ball when it had fully
descended into the pit. The male then clasped the female in the copulatory position, and kept strumming on the female’s elytra with its forelegs. The frequency of strumming increased whenever other insects moved in the vicinity. The female generally fed on the dung ball while engaged in mating, while at times she stood still doing nothing. On completing mating which took about 13 min., n = 4 (G. gemmatus ) and 19 min. 30 sec., n = 5 (G. miliaris ), the male stayed with the female for a further 4-5 sec., after which time it either flew away or stayed on to guard the female when he suspected the presence of other males. The female continued digging beneath the dung ball. Some pairs of beetles (3 of G. miliaris and 2 of G. gemmatus) were seen mating even before the pit was dug, such beetles abandoned the ball and Hew away.
Occasionally males were found trying to mate on the dung pat, but were not successful.
Brood ball construction: After mating, the female proceeded to make a slanting tunnel which terminated in a brood chamber. The ball was torn apart and refashioned into a pear shaped ball. An egg was laid in the egg chamber constructed at the lop end of this fashioned brood ball. The ball rested on its broad end in a pear shaped brood chamber located at an average depth, of 5. 1 cm (G. gemmatus) and 7.5 cm (G. miliaris). The average length and breadth of the brood balls of G. miliaris and G. gemmatus were 1 .45 ± 0.23 cm and 1 .2 ± 0. 1 8 cm (n = 12); and 1.34 ± 0.31 cm and 1.1 ± 0.08 cm (n = 1 0), respectively. Their respective weights were 1.12 ± 0.2 g (n = 12) and 0.99 ± 0.18 g (n = 10).
Competition: Both the species of
Gymnopleurus under review exhibited inter- and intra-specific competition.
Intra-specific competition: There was intense competition among the males to gain possession of ready made dung balls. This was observed from the very first day of activity.
Ball rolling males had to repeatedly fend off males that challenged the right to ownership of their ball. On sensing the arrival of a rival, the owner stood on the ball with its head towards the ground. The strategy of the rival male was always to move around
FEEDING AND BREEDING BEHAVIOUR OF GYMNOPLEURUS GEMMATUS AND G. MILIARIS
17
the ball trying to get a foot hold. The owner of the ball would keep moving on the ball so that the rival was always on the ground without access to the ball. If the rival came too close, the owner tossed it away using its clypeus. But if the rival managed to out manouvre the owner and gel atop the ball, then it in turn would toss the owner away.
Whenever a rolling pair was challenged by the arrival of a rival male it was only the male that fought him while the female remained passive. They butt and toss each other in combat. If the period of waiting was too long the female abandoned the fighting males with the ball, in search of a new partner.
Whenever a pair of rollers were rolling a ball, they were followed by males of their own species as well as other species. The males followed the ball either by walking or flying. Whenever the pair stopped rolling, the male that was following behind hid under grass or leaves in the vicinity. In this way, males followed the ball till the pair started burying the ball. Then the males in hiding attempted to reach the digging female, but retreated whenever the ‘owner’ male noticed and chased them. It was also observed that when the owner male succeeded in getting at the smaller intruder males, he sat atop them and started drumming on them with his clypeus and forelegs after which the looser ran away and took cover in the grass.
It was also noticed that whenever a male sitting and waiting on a ball, saw another male sitting or rolling a ball nearby it immediately chased the other male and took possession of that ball. In one such case a male took poosession of new balls on three successive occasions.
Competition was seen even after the ball was buried. Males of G. miliaris and G. gemmatus were seen landing and walking straight into mounds of soil where pairs of G. miliaris and G. gemmatus were found with dung balls. Males of one species sneaked into the burrows of other species.
Interspecific combat: Various species of Onthophagus and Caccobius also compete for access to dung balls in addition to the competition that has been noticed between the two species of the Gymnopleurus under study. Three species of
Onthophagus, namely O. centricornis. O. pygmaeus and O. ludio Bouc. and one species of Caccobius , namely C. meridionalis Bouc., tried repeatedly to gain entry into the brood balls of both G. miliaris and G. gemmatus.
Competition was noticed between the males of G. miliaris and G. gemmatus for the balls, and was similar to that explained in intraspecific competition. In one instance combat was observed between two females of Gymnopleurus spp. for the ball. The ball was pulled out by G. gemmatus and during the process of combat the ball was being rolled along.
The presence of kleptoparasites was noticed right from the ball construction stage to even after the egg had been laid in the brood ball. But they proved to be more persistent attackers once the beetles commenced rolling the balls. The kleptoparasites followed the rolling pairs on the wing, alighted in the vicinity of the ball and sought a hasty entry into it whenever the pairs paused for some reason. One pair of G. miliaris that was observed, had to face attacks from G. gemmatus (once), O. centricornis (eleven times), O. pygmaeus (four times) and O. ludio (once), while having to traverse a distance of 17.10 m.
If detected the Gymnopleurus spp. butted and tossed away the sneaking kleploparasitic species. Even those kleptoparasites that had managed to enter the ball unnoticed were detected in a short while and extricated from the ball after it had been pried open by Gymnopleurus. Along with the kleptoparasitc a small amount of dung was lost. The ball was then refashioned once more and the process of rolling continued.
It was also noticed that those males with the ball abandoned by the female never extricated the kleptoparasites when they entered the ball but Hew away abandoning them.
The kleptoparasites that escape detection convert the brood balls of Gymnopleurus into brood masses for their own off-spring, O. centricornis was observed to have made four brood masses out of one brood ball in one instance. In two other cases they had converted it into 2 or 3 brood masses. In the
18
JOURNAL. BOMBAY NATURAL HIST. SOCIETY. Vol. 93 (1996)
latter case the brood masses weighed 160, 1 10 and 1 18 mg.
Predation: Analysis of the stomach contents of the garden lizard Calotes sp. revealed fragments of Gymnopleurus.
Ants of the genus Camponotus were also found attacking ball rolling individuals of G. gemmatus. When attacked, the beetles abandoned the balls and Hew away.
G. gemmatus and G. miliaris invest a considerable quantum of their time and energy in the fashioning ( 1 1 .08 ± 1 .36 min., 1 0.40 ±1.14 min. respectively) and burial (14.36 ± 3.66 min.; 1 3.45 ± 2.8 min. respectively) of dung balls. The evolution of this behaviour not only enables these species, like the other dung rollers, to rapidly acquire the necessary resources in ball form, but it also provides the females ample opportunity to choose a more fit male. Hence, while giving these species an edge in the competition for food and burial space over the other burying groups like the Coprini, Onthophagini, etc., which are competing beneath or around each dung pat (Halffter and Matthews 1966) this behaviour simultaneously ensures rigorous epigamic selection (Huxley 1938). However, the greater amount of time spent on the surface of the ground while rolling exposes these beetles to the hazards of parasitism and predation.
Observations made during the current study indicate that while pre-mating female choice docs occur in both the species, female choice during and after mating are also possible.
Pre-mating female choice occurs during the three stages of fashioning, transportation and burial of the ball.
i. Ball fashioning: To increase their reproductive success by attracting females, males of G. miliaris and G. gemmatus have to fashion relatively larger balls of excrement. As there is very low correlation between the size of the male and the size of the female (r=0.08) the female does not seem to be choosing males. The weak correlation between the male size and ball size indicates that large individual size does not necessarily result in large ball size (r=0.2). All this combined with the fact that
female size and ball size are relatively highly correlated (i-0.57) points to the possibility of cryptic female choice (Fig. 1).
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FEEDING AND BREEDING BEHAVIOUR OF, GYMNOPLEURUS GEMMATUS AND G. MILIARIS
19
Instead of choosing the larger males (1-O.O8) females are choosing larger balls (r=0.57).
ii) Ball transportation: Having initially chosen a male capable of making a larger ball, the female further assesses male capability/fitness by:
a) watching how soon the male she has chosen can ward off the intruder males; b) choosing the winning male in the event of a skirmish. Males which take either too long to win a fight or are ousted in battle are abandoned by the females.
iii) Ball burial: In this phase males face the danger of losing females to other competing males or to having their effort wasted by intruding kleptoparasites which occupy the dung balls and destroy the eggs of Gynmopleurus. Intraspecific male competition is thus intense during these phases.
iv) Mating: During mating the female can exercise mate choice by regulating the mating duration. However, since the number of mating individuals observed during the study was small (n = 4 for G. gemmatus, forC. mi liar is n = 5) no definite conclusions could be arrived at in this regard (cv for
Refer
Arrow, G.J. (1931 ): The Fauna of British India including Ceylon and Burma. Coleoptera, Lamellicornia, III (Coprinae). Taylor and Francis, London.
Fabre, J.H. (1897): Souvenirs Entomologiques. Vol. V. Paris (translation by de Mattos, A.T. (1918) The Sacred Beetle and Others, London).
Halfeter, G. & W.D. Edmonds (1982): The nesting behaviour of dung beetles of the subfamily Scarabaeinae. Folia Entomologica Mexicana 12-14: 1-312.
Halffter, G. & E.G. Mathews (1966): The natural history of dung beetles of the subfamily Scarabaeinae. Folia Entomologica Mexicana 12-14: 1-312.
Hingston, R.W.G. (1923): A naturalist in Hindustan. H.F. & G. Witherby, London.
Honda, H. (1927): Intersting instincts of Gymnopleurus sinuatus (01.). Proceedings Imperial Academy Tokyo, 684- 686.
Huxley, J.S. (1938): Darwin’s theory of sexual selection and the data subsumed by it in the light of recent research. Am. Nat. 72: 416-433.
Prasse, J. (1957a): Nahrungserwerb koprophager Pillenwalzer ( Sisyphus scliaefferi L. und Gymnopleurus ge affray i Fuessl.). Wissenschaftliche Zeitschrift der Martin-Luther
G. gemmatus was 1 6.6, while for G. miliaris it was 21.7).
v) Post-mating: The female makes her final choice of the male by deciding whether to mate again or not. Having mated once, females have been observed to mate again. The presence of a horse- shoe shaped spermatheca also points to the distinct possibility of sperm precedence (Halffter and Edmonds 1982) as the sperm of the last mated male is most likely to fertilize the ova in spermatheca shaped thus. This could be the reason for the male to follow rolling bisexual pairs, waiting for an opportunity to be the final one to copulate with a female before she lays eggs.
Acknowledgements
We are grateful to Dr. R. Madge, British Museum (Natural History) for identifying the specimens. We wish to express our gratitude to Mr. Prashanth Mohanraj for having gone through the script and Mahalingappa for his assistance in field work.
ences
Universitat Halle Wittenberg 6, 439-444, cited in Halffter and Matthews (1966).
Prasse, J. (1957b): Das Brutfursorgeverhalten der Pillenwalzer. Sisyphus scliaefferi L. und Gymnopleurus gepffroyi Fuessl. (Col. Scarab.). Wissenschaftliche Zeitschrift der Martin- Luther Universitat Halle Wittenberg, 6, 589-614, cited in Halffter and Matthews (1966).
Prasse, J. (1957c): Die Entwicklung der Pillenwalzer Sisyphus scliaefferi L. und Gymnopleurus gepffroyi Fuessl. (Col. Scarab.) inder Brutbirne. Wissenschaftliche Zeischrift der Martin-Luther Universitat Halle Wittenberg, 6, 1033 — 1044, cited in Halffter amd Matthews (1966).
Prasse, J. (1958a): Verhaltensweise des Pillenwalzer Gymnopleurus mopsus Pall. (Col. Scarab.). Biol. Zbl. 77: 714-723.
Prasse, J. (1958b): Diekample der Pillerwallzer Sisyphus scliaefferi (L.) and Gymnopleurus gepffroyi Fuessl. (Col. Scarab.). Wissenschuftliche Zeitschrift der Martin — Luther Universitat Halle Wittenberg 7, 89-92, cited in Halffter and Matthews (1966).
Thornhill, R. (1983): Cryptic female choice and its implications in the scorpion fly Harpobittacus nigriceps. American Naturalist 122: 765-788.
NOTES ON LONG-EARED HEDGEHOG HEMIECHINUS AURITUS (GMELIN)1
Satish Kumar Sharma2
( With a text-figure )
Key Words: hedgehog, sex ratio, body-weight, nurseries, litter-size
The Long-eared Hedgehog Hemiechinus auritus (Gmelin) is a nocturnal mammal found in the arid and semi-arid areas of Rajasthan. A study was conducted at World Forestry Arboretum. Jaipur, on the biology and behaviour of H. Auritus between June 1989 and June 1991. Male-female sex ratio in adults was nearly 1:1. The mean weight of the adult females (306.2 gms) was slightly more than that of adult males (256.6 gms). Their breeding coincided with the rainy season and the Young are born in burrows. Litter size varies from 1-2. The female alone takes care of the young. The young are born blind and without spines. Later the mother escorts her babies during night rounds. The young become independent before commencement of summer. Hedgehogs are destroyed in various ways in Rajasthan. We can save them only by creating awareness among the public.
Introduction
The Long-eared Hedgehog Hemiechinus auritus (Gmelin) is a nocturnal spiny small mammal, widely distributed and known in the slate of Rajasthan by vernacular names such as ‘Jhaoo- chuha’, ‘ Jhaoo-musa’ , ‘Jhaoo’, ‘Jhawla’, kBhuinthda\ ‘Dhuan-dhuan’, ‘Gaoo-ghota’, etc. It is mostly confined to arid and semi-arid portions of the state. Very little is known about the breeding biology and other habits of this animal. In this paper, these aspects of this species are described.
Study area
The present study was mainly conducted at the World Forestry Arboretum (Part A), Jaipur, confined to an area of 1 .45 sq. km. The arboretum is situated on the outskirts of Jaipur city between National Highway (Bye Pass) No. 8 and Jhalana hills.
This place receives an annual rainfall of 548 mm. The maximum rainfall is received during July and August. The average relative humidity is 58.6 percent. The temperature ranges from 2° C to 44° C (av. 23°C). Eastern part of the arboretum is hilly but aeolian (soil transported by wind) sand deposit is confined to the foothill zone with Leptadenia
'Accepted August 1995.
:Range Forest Officer, Aravalli Afforestation Project, Jhadol (F.), Udaipur 313 702.
pyrotechnica, Calotropis procera, Saccharum bengalense, Mytenus emarginata, etc. The sandy tract of the arboretum provides an ideal habitat to the Long-eared Hedtiehog.
Materials and methods
Observations were taken from June 1989 to June 1991. Long-eared Hedgehogs were captured during night time. Two of our night-watchmen rendered their services for the purpose. From July to October, intensive night surveys were made to capture the animals. All the captured animals were sexed, marked and freed after weighing. To give a particular code number, tips of spines of cephalic and lumber regions were cut using scissors. Each year a fresh marking was practiced.
Pit digging and planting are two major operations in the forest areas during rainy season. While doing these operations in the arboretum, all the officials and labourers were requested to note and inform about nurseries of Long-eared Hedge- hogs, when encountered.
Foraging individuals were captured after dusk and before dawn. No animal was found naturally wandering in the day time. Bagged animals were weighed on a portable dial balance in the field to avoid any loss in body weight of animals due to exertion and starvation.
NOTES ON LONG-EARED HEDGEHOG
21
Results
Breeding: Data on animals caught arc given in Table 1 . It suggests that the breeding period of the animal coincides with the rainy season, i.e. June to September. It can also be concluded from Table 1 that during winters, to escape severe cold, the animals probably hibernate. During the months of October and November, when cold is not severe, few wandering animals were observed in the night; but, from December to February, which are the severest months of the winter, not a single animal was observed in the field. This suggests that they probably hibernate, become lethargic during the severely cold months. Prater (1980) has suggested this possibility. It can also be deduced from Table 1 that 1 to 2 young are found in a litter. Interestingly the litter size is very small in comparison to the five pairs of teats present. Early in fant mortality may be one of the reasons, for the recorded litter size.
At the time of birth, the young are blind and lack spines. During infancy, babies have flexible spines on their dorsal and lateral sides. As the young grows, the spines stiffen. Initially a fairly good number of spines seemed white in colour but gradually became shiny black. Except light coloured whiskers on snout, the head, chin, throat, limbs, ears and ventral side generally lacked hairs. The bare skin of these parts are pinkish in colour. Pinnae were proportionally smaller in infants. As they grow, size and erectness increases in pinnae and a fold develops on lower edge of each pinna at adulthood. When ‘ball’ formation takes place, this fold helps the animal to adjust the long ears in the minimum of space.
Though adult individuals have a pointed pig- like snout, the newly born do not have this character. Their snouts are almost blunt having roughly equal sized upper and lower jaws while in adults the upper jaw projects far beyond the lower jaw. The blunt snout of the baby hedgehog could be an adaptation for suckling. As the young grows, the snout gradually tapers and ultimately becomes pointed in adulthood.
Parental Care: The young are born in burrows. It was general observation shows that
wherever the nursery of a female was disturbed, it abandoned its burrow with its new born as early as possible. The young are transported by the mother at night only. She carries them by gripp- ing the loose skin of the side of neck or body
Pig. 1. A female Hedgehog transporting her young. (Fig. 1).
When the young are separated from their mothers, they make a shrill sound. When a foraging mother returns to her burrow and senses something unusual (like the presence of an observer in a hide near the burrow) she may wait among the thickets till she is assured that there is no danger near the burrow. She may even remain away from her young for a whole day and a night or even more, as is evident by the behaviour of the female C-6-1990. The activities of the female are given in Table 2.
During September 1990, an adult female and two small sized young were found run over in the morning on the bye-pass road bordering the arboretum. They had been run over during the night and their bodies had became Hat. All the carcasses were roughly in a straight line, parallel to the road. Heads of all the three individuals were facing south. Interestingly, corpses of both the young ones were behind the mother. All three dead animals were confined to a 40 cm x 20 cm area. An almost similar incidence was observed in October 1990. These observations led to the assumption that after the eyes are open, the young hedgehogs follow the mother at
Table 1
MONTH-WISE DATA ON LONG-EARED HEDGEHOGS IN WORLD FORESTRY ARBORETUM. JAIPUR.
22
JOURNAL BOMBAY NATURAL HIST. SOCIETY. Vol. 93 (1996)
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NOTES ON LONG-EARED HEDGEHOG
23
Table 2
DEPARTURE AND RETURN TIMINGS OF DISTURBED FEMALE HEDGEHOG C-6-1990
|
Date |
Time of departure from burrow (hrs) |
Time of return to burrow (hrs) |
Time of stay in burrow (minutes) |
Time of slay outside burrow (minutes) |
Observations |
|
14.9.90 |
1905 |
203 1 |
— |
86 |
|
|
2035 |
- |
4 |
- |
Carried one young. |
|
|
15.9.90 |
— |
— |
Not returned to burrow at all. |
||
|
16.9.90 |
— |
0615 |
— |
2020 |
— |
|
0618 |
- |
3 |
- |
Carried another young. |
night. During night surveys ‘Caravans’ of Hedgehog families eaptured in groups of 1 + 1 or 1+2, with the mother leading the juveniles, substantiates this fact (Table 1 ).
During November 1990, three solitary young of the current year were captured wandering at night. One solitary young was also found dead in a water tank. Before June and after December no female was captured with young. These observations suggested that the young hedgehogs become independent before the summer of the same year.
Sex ratio and body weight: A total of 35 adults and 1 3 young ones were weighed in the field. The mean weight of adult females (306.2 gm, n= 1 7) was slightly more than that of adult males (256.6 gm, n= 1 8). The adult sex ratio was nearly equal. The blind young ones about to open the eyes in burrows weighed only about 60.1 gm (n=10) while the independent young hedgehogs weighed 125.2 gm (n=3)
Destruction of hedgehogs: A considerable number of Long-eared Hedgehogs are destroyed in Rajasthan every year. It is believed by rural people that hedgehogs are rat-destroying animals, hence they are transported to houses from fields. Due to their removal from the fields, many mothers get separated from their new born. It is obvious that the young cannot survive without their mothers. Similarly, there is a practice among rural people to fumigate the Cucurbitaceous climbers, like Luffa acutangula, L. cylindrica, Benincasa hispidci , etc., by burning the skin and spines of Hedgehogs to induce heavy fruiting. Hedgehogs are killed for medicinal purposes also. During epidemics of foot and mouth disease of cattle, it is a general practice
in many parts of Rajasthan, specially in the southern part, to bury a live hedgehog in a pit in front of each house. Cattle are then passed over the pit.
During nights, wandering Hedgehogs may stumble in wells and tanks that arc without parapet walls (Sharma 1 993). Many are runover by vehicles (Sharma 1988, 1992).
Conservation measures: Hedgehogs play an important role in destruction of insects and even rats and mice (Sinha and Ray 1983), hence they should be protected. A considerable number of hedgehogs are killed in rural areas. “Gram-Sewak” — an ‘agricultural guide' in rural areas can educate farmers and agricultural labourers by teaching them about the role played by the hedgehogs and other wild animals in an agro-ecosystem. Agriculturists should be advised not to bring the hedgehogs, specially females, to the home, atleast during breeding season. The children’s science books may also have a chapter on these animals which are beneficial to our agriculture.
Raising of parapet walls around wells and tanks on agricultural land avoids the falling of stumbling hedgehogs and other nocturnal animals into them. Vehicle drivers should be careful of animals crossing the roads.
Conclusion
It can be concluded from the present study that female hedgehogs are slightly heavier than males. Sex ratio among adults is nearly 1:1. Young ones are given birth in burrows during rainy season when
24
JOURNAL BOMBAY NATURAL HIST SOCIETY, Vol. 93 (1996)
grass cover and plenty of insects are available. Litter size varies from 1 to 2. The female alone takes care of the young ones. If a nursery of a female is disturbed, she transports her young one by one to another safer place during the night. Babies are transported by her by griping the loose skin of the neck or body as members of the Cat family do.
A large number of hedgehogs are destroyed in Rajasthan every year for domestic uses. These animals cannot be saved without creating awareness in public.
Acknowledgements
I am grateful to the night watchmen of the arboretum, Jaipur, for helping me in hedgehog collection during night surveys. I am thankful to all the labourers and staff, specially to Mr. Vijay Saxena, Forester, for having given me information about nurseries of hedgehogs. Thanks are also due to Mr. Subhash Bhargava for accompanying me for field photography.
References
Prater, S.H. ( 1 980): The book of Indian animals. Bombay Natural History Society. Bombay.
S harm a, S.K. ( 1988): Wild animals and road accidents. Vijnana Parishad Anusandlmn Patrika 31(1): 43-53.
Siiarma, S.K. (1992): Accidents of wild animals on roads at outskirts of Jaipur city. Vijnana Parishad Anusandlmn
Patrika 35( I ): 47-60.
Siiarma, S.K. (1993): Cemented tanks in forest areas and wildlife management. Indian Forester 119(10): 849- 852.
Sinha, N.K. & P. Ray (1983): New Records of Hedge-hogs from Madhya Pradesh. Cheetal 25 ( I): 36-37.
ROOST SELECTION BY INDIAN PEAFOWL (PA VO CRISTATUS) IN GIR FOREST, INDIA1
Pranav Trivedi2 and A.J.T. Johnsingh3
Key Words: roost, riverine forest, preference index, selection, structure, floristics,
predation
A study was carried out on roost selection of Indian peafowl ( Pavo cristatus) in Gir forest, Gujarat.
The results revealed that all the roosts were located in the narrow riverine forest bells. Peafowl selected tall trees growing on steep river banks with thorny undergrowth and climber thickets in the canopy for roosting.
This clearly indicates that roost selection is chiefly influenced by the risk of predation from nocturnal mammalian predators such as leopard. Trees of Pongamia pinna to and Holoptelia integri folia showed more than expected use. However, it could not be confirmed whether a choice at species level does exist at all. Roost selection appeared to be a hierarchial process with structure at first and floristics at second level affecting the choice.
Introduction
Roost selection is a vital component of the overall habitat selection process. Therefore information on roost selection by a species carries immense importance for assessing its conservation needs. Gadgil and Ali (1975) attempted to explain the communal roosting habits of Indian birds based on the existing hypotheses which include reduced heat loss, information sharing, assessment of population and reduced risk of predation. Though, Indian peafowl (Pavo cristatus ), a common bird of India is known to roost in the trees at night, no information exists on roost selection by the bird. In a strict sense, it is neither a communal nor a solitary rooster (Trivedi 1993).
This paper attempts to provide information on roost selection by Indian peafowl in a wild landscape. The following results were obtained during a study carried out from November 1992 to April 1993 (Trivedi 1993) on habitat selection by peafowl in Gir forest.
Study Area
The study was carried out in Gir National Park (N.P.) and Sanctuary [(both collectively hereafter
'Accepted January 1994.
2WWF-India, Ahmedabad Division Office, ‘Sundarvan’, Jodhpur Tekra, Ahmedabad 380 015.
’’Wildlife Institute of India, P.O. Box 18, Dehradun-248 001.
referred to as Gir Protected Area (PA)] located in Gujarat, India. Gir PA (1412 sq. km) is the only remaining large, contiguous, forested tract in the Saurashtra peninsula of Gujarat. The PA is covered with tropical dry deciduous forests, thorn forests and riparian forests. The chief floral elements include Tectona grand is, Diospyros melanoxylon, Wrightia tinctoria, Zizyphus mauritiana, Ficus bengalensis, Morinda tinctoria, Phyllant/uis emblica, Bauhinia racemosa, Holoptelia integrifolia, Boswellict serrata and Lannea coroniandelica.
The PA is the last stronghold of the Asiatic lion (P anthera leo persica) and apart from lion the vertebrate fauna includes leopard (Panthera pardus), jackal (Canis aureus), jungle cat (Felis chans) and crested hawk eagle (Spizcietus cirrhatus) as potential predators of peafowl. Checklist of mammals is available in Spillett (1968). Nearly 250 species of birds have been recorded.
The ‘Maldharis’ who are local pastoralists and have changed their nomadic lifestyle to a settled one, reside inside the Sanctuary in their settlements called ‘ness’. However, N.P. is free from all human activities. Buffalo grazing, tourism, grass harvesting, fireline burning and non-wood forest produce collection are the chief human influences.
Methods
Eight different localities in three study sites (Sasan, Chhodavdi, Dodhi) were surveyed for roost
26
JOURNAL BOMBAY NATURAL HIST. SOCIETY, Vol. 93 (1996)
Table I
PREFERENCE INDICES FOR SOME PHYSICAL FEATURES OF ROOST TREES USED BY PEAFOWL
|
Slope category |
Distance to water (in m) |
Height (in in) |
Height of first branch (in m) |
||||
|
Class |
FI |
Class |
PI |
Class |
PI |
Class |
PI |
|
Very steep |
1.45 |
0-25 |
LOO |
0-10 |
0.08 |
0-2 |
0.61 |
|
Steep |
0.81 |
26-50 |
0.87 |
11-15 |
3.38 |
2.1-4 |
1.73 |
|
Gradual |
0.90 |
5 1 -75 |
1.08 |
16-20 |
6.25 |
4.1-6 |
2.57 |
|
Flat |
0.60 |
76-100 |
0.73 |
21-25 |
8.10 |
6.1-8 |
2.69 |
|
— |
- |
>100 |
0.73 |
>25 |
8.09 |
8.1-10 |
3.44 |
PI= Preference Index.
tree use by peafowl. Both, direct and indirect methods were used to locate and identify the roost trees. The former involved walking along the riparian areas during late evening or early morning, to flush the roosting birds and locate the trees. The latter involved searching for droppings below potential roost trees to identify actual roost trees. When a roost tree was located, GBH, height, of the first branch, tree height, slope category of the site where the tree was standing (rated qualitatively as very steep, steep, gradual and flat), distance from water (or water body), canopy and understorey characteristics were recorded. The same data were collected on the ten nearest trees from the roost tree to get availability information. In this manner, use and availability of the trees was determined. A widely used method given by Neu et cil. ( 1 974) was employed for analysing the availability-use information. Preference index (PI) which is expressed as a ratio of per cent usage to per cent availability was calculated for the structural parameters of roost trees.
Results
1 034 trees were quantified as described earlier. Of these 128 were roost trees, which reflected the use, and the remaining gave an idea about the availability. All the roosts were located in riparian
areas. Roosts were found to be either continuously spread along the riverine areas or located at the confluence of two streams which is normally a steep area.
Structure: There were differences in the use of trees with and without certain structural features. There was a significant difference between the use of trees with and without thickets of climbers in the canopy (x2= 10.62, df=l, p<0.01). Similarly, there was also a significant difference between the use of trees with and without thorny undergrowth (x2= 24.6 1 , df= 1 , p< 0.00 1 ). In both the cases, trees with thickets of climbers in the canopy and with thorny undergrowth were used more often for roosting. Preference indices (PI) for various structural features are presented in Table 1. Trees on very steep and steep river banks received a higher usage followed by gentler slopes. Trees growing on flat areas were least used for roosting. All tree height categories above 15 metres were highly used, while category <10 metres was used least. Nearly ninety percent of the trees were within 75 metres from water. Trees with 8-10 m high first branch were used more and the use went in a decreasing order towards 0-2 m height.
Floristics: Twenty one plant species were identified as roosts (Appendix 1 ) of which twenty were trees and one was Dendro calciums strictus (i.e. bamboo). Table 2 shows the availability and use of
ROOST SELECTION BY INDIAN PEA F OWL
27
Table 2
ROOST TREE PREFERENCE OF PEAFOWL [Using Neu et al. (1974) technique]
|
Tree species |
Relative availability |
Expected Observed use use |
Confidence intervals |
|
|
Holoptelia integrifolia |
0.051 |
6.554 |
18 |
0.058-0.223** |
|
Tectona grandis |
0.124 |
15.846 |
14 |
0.035-0.184 |
|
Pongamia pinnata |
0.198 |
25.370 |
41 |
0.209-0.431** |
|
Syzygium rubicunda |
0.131 |
16.717 |
13 |
0.030-0.173 |
|
Tamarindus indica |
0.181 |
23.168 • • |
6 |
0.000-0.097* |
|
Diospyros melanoxylon |
0.033 |
4.224 |
2 |
0.000-0.045 |
|
Others |
0.282 |
36.096 |
34 |
0.161-0.371 |
Indicates that the species was used less than availability. ** Indicates that the species was used more than availability.
Rest were used in proportion to availability.
(Z=2.6899, X2= 45.36)
major roost trees by peafowl. Holoptelia integrifolia, Tectona grandis, Pongamia pinnata, Syzygium rubicunda, Tamarindus indica and Diospyros melanoxylon were the commonest tree species available and used as roost by peafowl. Rest of the species were in meagre numbers and therefore these were clumped and collectively called ‘others' for analysis. Availability-use analysis of these six species and others showed (Table 2) that only H. integrifolia and P. pinnata were used more than expected; T. grandis, S. rubicunda, D. Melanoxylon and others were used in proportion to availability whereas T. indica was used less than its availability.
Discussion
The five most striking features of the roost trees selected by peafowl were; they had climber thickets in the canopy, possessed thorny undergrowth, were on steep river banks, were tall
and had a higher first branch. All these features indicate that while selecting a roost tree, the most important aspect is of reducing the risk of predation. In Gallinaceous birds, predation is a major population regulatory mechanism (Lack 1954, Hill and Robertson 1 988) and therefore it is likely to influence habitat selection significantly. Selection of trees with the above mentioned features is obviously an antipredatory strategy against nocturnal mammalian predators such as leopard and jungle cat which can climb trees and capture peafowl. In Gir, trees with such features are available only in riverine areas and therefore these forests become crucial for peafowl. The location of roosts al the confluence of two streams was due to the fact that this region is steep and therefore predators would find it difficult to approach from below.
It is pertinent to point out that the height of first branch does not seem to be of significance in roost selection. Similarly, distance from water carries secondary importance as all the roosts are located in the riverine area and one hardly finds a roost >100 m from water. However, trees growing right along the bank with overhanging branches above the river provide ideal roosts as birds are safe from the predators due to water. The vital features, therefore appear to be height of the tree, steepness of the bank/ slope on which the tree is situated and the presence of thickets in the undergrowth and in the canopy. It was realized that height alone can be sufficient for selection if the tree is > 1 6 m. But, if it is shorter than that, the other tree features play a crucial role. In a semi-arid and deciduous forest system such as Gir, trees hardly attain a height of over 15 m and therefore it is the presence of thickets and steepness of the slope that should be of significance in the selection process.
Peafowl in semi-urban and rural landscapes often use unusual substrates as roosts, like electric pylons. Palmyra trees ( Borassus flabellifer) are commonly used in the Southern districts of Tamil Nadu (pers. obs.). This flexibility probably reflects a synergistic effect of the absence of predation pressure and a low availability of good quality roosts.
28
JOURNAL. BOMBAY NATURAL HIST. SOCIETY. Vol. 93 (1996)
Interpretation of the data suggests that it is the strueture whieh is the unit of seleetion at a broader seale, but at a finer scale, the selection can be for species. Any tree which satisfies the structural requirements for avoiding predators should be selected by the birds. Structure undoubtedly appears to be the first step in roost selection process. It is possible that only certain tree species possess the necessary structural features of an ideal roost tree which means the choice can be at the level of species. The situation seems to be one of a hierarchial selection as described by Svardson (1949), Hilden (1965) and Wiens (1985). However, this is just a logical speculation and no experimental evidence is available to test it. Peafowl (Genus Pavo) are regarded as the terminal lineages of peacock pheasants (Geist 1977). Pavo left their original rain forest habitat and started exploiting the productive forest-water ecotone (Geist 1977). They gradually advanced to human dominated landscape also, but were always lied to riverine habitats. Roosting on riverine trees might have evolved at the time of their dispersal from climax forests to more xeric environments, because in these habitats only riverine forests can provide good quality roosts.
One more important feature which influences roost selection is the occupancy of trees by other species. It was observed on ten occasions that peafowl did not use particular trees (even when these were ideal for roosting) because common langurs ( Presbytis entellus ) were roosting there. This brings in the question of competition between taxa for a crucial resource, as langurs also roost to avoid predation by leopard. Langurs too, like peafowl arc distributed along the riverine areas in Gir (Joslin 1973) and they too roost in riverine forest. However,
R E F E
Gadgil. M. & S. Ali (1975): Communal roosting habits of Indian birds. J. Bombay not. Hist. Sac. 72(3): 716-727.
Geist, V. ( 1977): A comparison of social adaptation in relation to ecology in gallinaceous bird and ungulate societies. Ann. Rev. Ecol. Systematic s 8: 193-207.
Hilden, O. ( 1965): Habitat selection in birds: A review. Ann. Zool. Fenn. 2: 53-75.
Hill. D.A. & P.A. Robertson (1988): The pheasant: Ecology,
the magnitude of such potential competition might not be significant. Only one roost tree of chicks was identified which was short (c. 10 m) and had extensive thorny thickets wrapped around the stem. With the exception of four trees (out of 128), no roost tree was located close to the road presumably to avoid the disturbance caused by the vehicles.
Peafowl alone with common langur are important buffer prey which facilitate the niche separation of leopard and lion in the Gir PA (Ravi Chellam 1993). Both the prey species need to be conserved. Both need roost trees in the riparian areas. In Gir, there is hardly any disturbance to the trees in riverine areas, but incidences of repeated fire can destroy the undergrowth thickets substantially thereby reducing the availability of good quality roosts. At present it is safe to conclude that the population of peafowl in Gir does not face any imminent danger. Our data on roost selection can be used to predict and confirm the use of roosts in other such deciduous forest ecosystems in a wild landscape.
Acknowledgements
We extend our thanks to the Forest Department, Gujarat for granting us the permission to work in Gir and for providing the necessary infrastructure. Qamar Qureshi and Ravi Chellam of Wildlife Institute of India (W.I.I.) commented upon the earlier draft of this paper and gave valuable suggestions. Diwakar Sharma of W.I.I. provided encouragement and support. We thank our field assistants at Gir without whose help it would not have been possible to obtain this data.
E N C E S
Management and Conservation. BSP Professional books, Oxford. 281 pp.
Joslin, P. (1973): Behaviour and ecology of the Asiatic Lion ( Panthera leo persica). Ph.D. Thesis, University of Edinburgh. Lack, D. (1954): The natural regulation of animal numbers.
Clarendon Press, Oxford. 343 pp.
Neu, C.W.. C.R. Byers & J.M. Peek (1974): A technique for analysis of utilization-availability data. WilclI. Manage. 38:
ROOST SELECTION BY INDIAN PEAFOWL
29
541-545.
Ravi Chellam (1993): Ecology of the Asiatic Lion ( Panthera leo persica). Ph.D. thesis. Saurashtra University, Rajkot. Spilu-tt, J.J. (1968): A report on wildlife surveys in South and West India, November-Deceinber 1966.7. Bombay not. Hist. Soc. 65: 1-46.
Svardson, G. (1949): Competition and habitat selection in birds.
OikosL 157-174.
Trivedi, P. (1993): Habitat selection by Indian peafowl (Pavo cristatus Linn.) in Gir forest, India. M.Sc. Dissertation, Saurashtra University, Rajkot. 78 pp.
Wiens, J.A. (1985): Habitat selection in variable environments: shrub steppe birds. In Cody, M.L. ed. Habitat selection in birds. Academic press, Inc., New York. 227-252 pp.
APPENDIX 1
LIST OF SPECIES USED FOR ROOSTING BY PEAFOWL
1 . Holoptelia integrifolia
2. Tectomi grandis
3. Pon garni a pinnata
4. Syzygium rubicunda
5. Tamar Indus indica
6. Dio spy ms melanoxylon
7. Tenninalia belle rica
8. T. tomentosa
9. Manilkara lie.xandra
10. Syzygium at mini
1 1 . Ficus glome rata
12. F. bengalensis
13. Miliusa tomentosa
14. Mitragyna parviflora
15. Ga ruga pinnata
16. Sterculia urens
17. Acacia Senegal
18. Anogeissus latifolia
19. Phoenix sylvestris
20. Dendrocalamus striclus
2 1 . Sapindus emarginatus
TAXONOMIC AND NOMENCLATURAL STATUS OF MYRIONEURON R.BR. EX
HOOK. F. (RUBIACEAE)1
D.B.Deb2
Key words: plant taxonomy, Rubiaceae, Myrioneuron, generic status, nomenclature
Taxonomic and nomenclatural status of Myrioneuron R. Br. ex Hook.f. (Rubiaceae) is discussed. The generic status is upheld. Nomenclature is clarified. Lectotypes of the genus and the type species are selected.
Introduction
The genus Myrioneuron (Rubiaceae) is not yet included in Index Nominum genericorum (Plantarum). R.C. Bakhuizen (1975: 26, 29) treated Myrioneuron R. Br. as synonymous with Mycetia Reinw., and those of authors, non R. Br. as synonymous with Keenania Hook. f. Van Steenis ( 1 987 : 1 06) treated Myrioneuron R. Br. ex Kurz 1 870 = ? Keenania. Myrioneuron spp. (in Herb.) = Keenania. He (1 . c.) further included this name as a synonym of Mycetia. Robbrecht (1988: 244) treated Myrioneuron R. Br. ex Kurz, nomen = Keenania ?
Diane Bridson of Kew Herbarium (in lit.) drew my attention to the situation on examining some Indian material determined by me as Myrioneuron nutans. She further observed “neither Dr. Brummitt, nor I find any problem in accepting Myrioneuron R. Br. ex Hook. f. 1 873 as valid (assuming Myrioneuron R. Br. ex Kurz 1 870, nomen)”. Dr. Dan H. Nicolson, Nomenclatural Editor, Taxon , in response to my letter in this connection, advised me to publish a note.
History of Nomenclature
Nathaniel Wallich, the then Superintendent of the Botanic Garden, Calcutta, in 1828, took to London, all the specimens so far accumulated in CAL and brought out those stored in the East India Company’s India Museum , London (which was dispersed in 1879), with a view to sort out the specimens and name them. He sought help and
'Accepted June 1994.
2Central National Herbarium, Indian Botanic Garden, Howrah, India.
assistance from contemporary botanists, who were interested in tropical plants.
‘A numerical list of the dried plants in the East India Company’s Museum ” (1832), commonly known as Wallichian Catalogue (Wall. Cat.), more correctly, Wallich num. List is the result of that effort. Robert Brown of British Museum named many plants of the Rubiaceae. Wall. num. List No. 6225 in page 211, 1832 names Myrioneuron R. Br., under which M. nutans R. Br. is named for two gatherings: 6225a, collected from Sillet, in 1821, by Francis de Silva and 6225b from Gualpara, Assam, on 27th June, 1808 collected by Buchanan (later Francis) Hamilton. The latter was named by Hamilton as Bertiera nutans Ham. in Scheda (nom. nud.)
Robert Brown postulated generic status for Myrioneuron to accommodate these two gatherings and used the specific name given by Hamilton on the herbarium specimens, i.e. Myrioneuron nutans R. Br. as is evident in Wall. Num. List (1. c.). The taxon remained in name only until J.D. Hooker, in Benth. & Hook. f. Gen. p 1 . 2: 69. 1873 validated the generic name with a description. He did not name any species therein. Art. 37 of ICBN (1988) clearly states that prior to Jan. 1, 1958, for validity of publication of a new genus, it was not essential to name the type species.
Kurz ( 1 877:55) validated M. nutans R. Br. with a specific description for the material collected from Chittagong by C.B. Clarke, working as the first Curator of the Herbarium, Royal Botanic Garden, Calcutta, Kurz must have studied the Wallichian specimens of M. nutans R. Br. extant in CAL. Citation of R. Br. as the author of the species evidently supports this contention. Thus M. nutans
TAXONOMIC AND NOMENCLATURE STATUS OE M YRIONEURON
31
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32
JOURNAL BOMBAY NATURAL HIST. SOCIETY, Vol. 93(1996)
R. Br. ex Kurz (1877) is the validating description of the species which is selected as the lectotype of the genus. Wall. Num. List No. 6225a (the left hand specimen) collected by Francis de Silva from Sillet, extant in CAL is selected as the lectotype of the species.
Taxonomic status: The genus Myrioneuron R. Br. ex Hook. f. is apparently allied to Keenania Hook. f. and Mycetia Reinw. Characteristics of these 3 allied genera are given below:
Keenania Hook. f. FI. Brit. Ind. 3: 101. 1880.
Small subherbaceous shrubs. Leaves membranous; stipules somewhat recurved, membranous. Flowers sessile, in terminal solitary shortly peduncled involucrate heads; bracts imbricating, concave, coriaceous, unequal, outer ones orbicular, inner linear-oblong or spathulate; bracleoles in pair, spathulate, coriaceous, equalling the flowers. Flowers unisexual. Hypanthium fleshy, shortly oblong; calyx lobes 5 or 6, imbricate, unequal, erect, oblong or spathulate, coriaceous, concave. Corolla about as long as the calyx lobes; corolla tube inflated, glabrous, with a ring of stiff hairs at the throat; lobes 5, valvate, short, orbicular ovate, apiculate, papillose externally. Stamens 5, epipetalous at the base of broad lobed disc; filaments short; anthers small, linear. Ovary 2-loculed; style short; stigmas 2, flat, ovate; ovules numerous on globose placenta, adnate to the membranous septum. Fruit a capsule with hard endocarp.
Type species: Keenania modesta Hook. f.
Distribution: 5 species in India (Chachar, Assam) and SE. Asia.
Mycetia Reinw. in Bl. Bijdr. 986. 1 826 & Sylb. Ratisb. 2: 9. 1928. Syn. Ade nos acme Wall, ex Endl. Gen. 1: 552. 1838.
Shrubs; branches with a conspicuous spongy swollen corky bark. Leaves membranous, stalked glandular at the margin; stipules tardily caducous, oblong or lanceolate, stalked glandular at the margin, sometimes toothed, and bifid above. Flowers pedicelled, bisexual, heterostylous, sometime bi or tri-formous, in axillary or terminal peduncled paniculate or corymbiform, often with stalked glands in floral parts. Hypanthium globose or hemispherical;
calyx lobes 4-6, persistent, stalked glandular. Corolla tube cylindric; lobes 4-6, valvate in bud. Stamens 4- 6, inserted in the tube at different positions; filaments short; anthers linear-oblong, dorsifixed. Disc annular. Ovary 2 or 3-6 loculed; style slender or thickened above; stigmas 2, linear; ovules numerous on fleshy peltate placenta. Berry globose, white, spongy or not, indehiscent or irregularly dehiscent, 2-6 loculed. Seeds many, minute, angled; testa dotted; endosperm fleshy; embryo minute.
Type species: Mycetia cauliflora Reinw.
Distribution: About 25 species, India to S. China, Vietnam, W. Malesia.
Myrioneuron R. Br. ex Hook. f. in Benth. & Hook. f. Gen. PI. 2:69, 1873.
Shrubs; branches stout, with a conspicuous spongy swollen corky bark. Leaves large, coriaceous or subcoriaceous; stipules large, coriaceous, bifid above. Flowers on short stout peduncle in terminal or rarely axillary capitate or corymbose cymes, erect or nodding; bracts involucrate, rigid, coriaceous; pedicels very short, one bracteolate. Hypanthium ovoid; calyx tube very short, lobes persistent, rigid. Corolla cylindric, 5-toothed, valvate in bud, densely villous inside, shorter than the calyx lobes. Stamens 5, adnate to the corolla tube. Disc cushion like. Ovary 2-locular; style short; stigmas 2, lanceolate, cohering; ovules many on hemispherical placenta. Berry ovoid or globose, dry, sometimes fleshy, 2-coccous; cocci horny, slowly dehiscent. Seeds black, many, minute, angled, flat; testa pitted, albumen fleshy; embryo minute.
Type species: Myrioneuron nutans Wall, ex
Kurz
Distribution: 3 species in E. India, Bangladesh (Chittagong) and Myanmar.
Discussion and conclusion
Keenania Hook. f. was described as a monotypic genus on the basis of a gathering by R.L. Keenan from Duarbund, Cachar, Assam. The type specimen of the type species Keenania modesta Hook. f. of the genus was based on a specimen with sterile anthers and it did not have any fruit. This
TAXONOMIC AND NOMENCLATURE STATUS OF MYRIONEURON
33
species has never been recollected. Species described under this genus subsequently from SE. Asia have flowers apparently heterostylous, and the heterostyly is probably combined with dioecism (Bremekamp 1947: 191 ). The fruit is a two-loculed capsule with hard endocarp; flowers are sessile, apparently unisexual and heterostylous; the inflorescence is in terminal heads; leaves, stipules, bracts and calyx lobes are without stalked glands; branches are without spongy swollen corky bark. These characteristics keep this genus distinct from others.
The genus Mycetia Reinw. with about 25 species is more widely distributed. The ranges of vegetative and reproductive characters are much more variable than those in the other two. Leaves, stipules, bracts and calyx lobes are with stalked glands at margins; the inflorescence is terminal or axillary paniculate corymbiform or capitate cymes; flowers are bisexual, heterostylous; the fruit is fleshy or not, 2 or 5-6-loculed. These characters distinguish Mycetia Reinw. from others.
Short pedicelled bisexual, isostylous flowers,
2 coccous fruits with horny cocci; bifid stipules; leaves, stipules, bracts, calyx lobes, etc., without marginal stalked glands, etc., distinguish Myrioneuron R. Br. ex Hook. f. from the other two.
In the branches with a conspicuous spongy
Refer
Bakhuizen von den Brink, R.C. (1975): Thai For. Bull. Bot. 9: 15-35.
Bremekamp, C.E.B. (1947): A monograph of the genus Pomazota Ridley. J. Am. Arb. 28: 186-203.
Endlicher, S.L. (1838): Genera Plantarum. pail 8. Wien.
Hooker, J.D. (1873): Bentham, G. & J. D. Hooker, Genera plantarum. Vol. 2. London.
Hooker, J.D. (1880): The Flora of British India, Vol. 3, London. Kurz, S. ( 1 877): Forest Flora of British Burma. Caleutta.
Lindley, John ( 1 847): The Vegetable Kingdom, ed. 2.
swollen corky bark and in the ranges of vegetative and reproductive characters sometimes extending to such an extent that it appears that Mycetia is more akin to Myrioneuron and farther away from Keenania.
Myrioneuron R. Brown ex J.D. Hooker in Benth. & Hook, f., Gen. PI. 2: 69. 1873 (T. non designatus).
Myrioneuron R. Brown, nom.nud., in Wallich, Num. List 21 1, No. 6225. 1832; Steud., nom. 2: 174. 1841; Walp., Rep. 2: 525. 1843; Endl., Gen. 566. 1838; Lindl., veg. Kingdom 765. 1847 (g. Cinchonaceae). Lectotype selected here: M. nutans R. Brown ex Kurz.
M. nutans R. Brown ex Kurz, For. FI. Brit. Burma 2: 55. 1877. Lectotype selected here: Sillet, 1821, Francis cle Silva s.n. ex Wallich, Num. List No. 6225a (the left-hand specimen) in CAL is selected here as the lectotype.
Acknowledgements
I am thankful to Mrs. Diane Bridson (K) for drawing my attention to the uncertainty of the generic status, and to Dr. Dan H. Nicolson, Nomenclatural Editor, Taxon, Smithsonian Institution, Washington D.C., USA, for his views on nomenclature and suggestion to publish this note.
ENCES
Robbrechr, E. (1988): Tropical Woody Rubiaceae. Oper. Bot. Belz. I. Belzium.
Van Steenis, C.E.C. (1987): Checklist of generic names in Malesia Botany.
Steudel, E.G. (1841): Nomenclator botanicus, Vol. 2. Stuttgart, Tubingen.
Wallich, Nathaniel (1832): A numerical list of dried plants in the East India Company’s Museum. London.
Walper, W.G. (1843): Repertorium botanices systematicae. Vol. 2. Leipzig.
SOME ASPECTS OF BIRD/MAMMAL ASSOCIATIONS: CONTRIBUTIONS FROM THE
INDIAN PLAINS AND THE ZIMBABWE PLATEAU1
D. Ewbank2
Key words: bird/mammal association, Indian plains, Zimbabwe plateau
Some species of birds associate with mammals to feed on insects living either near or on them. This paper attempts to quantify the size of these aggregations with mammals and compare them with unassociated birds on the plains ol India and the plateau of Zimbabwe. In India four species of bird were observed associating: two of them occurred in larger groups in such associations and preferred cattle to water buffalo. In Zimbabwe in a drought situation, with a much lower density of mammals, a much higher proportion of observed birds were in such associations. Egrets are known to obtain more food in such associations but this is not proved for the other species involved.
Introduction
All organisms need a competitive edge of some sort for survival. One such advantage is feeding associations between birds and mammals which are a common sight in the tropics. Some birds gain significant amounts of food from such associations, e.g. in Africa oxpeckers Buphcigus spp. are obligate feeders on ticks (Ixodidae) and other ectoparasites and have lost range in southern Africa with the decimation of large mammal populations (Brooke 1984).
Another example of conservation significance concerns the endangered Kirtland’s Warbler Dendroicci kirtlandii in the United States, whose range has recently been invaded by a nest parasite the cowbird Molothurus ater. This bird was previously an associate of the American Bison Bison bison and has now expanded its range with the arrival of cattle populations in the range of the warbler and now reduces its breeding success by over a half (McFarland 1981, Robisnon and Bolen 1989).
There have been few quantitative studies of such associations and even fewer of a total community. To the best of my knowledge there are none published for India. Hence while travelling around northern India by train mainly through peasant owned farmland between April and August 1982 covering a distance of 3000 km, an attempt was made to record all associating birds and domestic
'Accepted March 1993.
274 High St., Landbeach, Cambridge, U K.
mammals and quantify their associations. No attempt was made to record all Common Mynas Acridotheres tristis because they are liable to be overlooked when perched in trees. The superabundance of the House Crow Corvus splendens led to my recording all sightings for only a portion of the total distance. Other species recorded associating with domestic mammals were Cattle Egret Bubulcus ibis and the Black Drongo Die rums macrocercus.
A similar study was performed over a number of years in Zimbabwe by vehicle through European farms and peasant occupied communal areas in the late eighties covering a distance of some 5000 km recording all birds associating and their hosts. The birds involved here were mainly Cattle Egrets again and Forktailed Drongo Dicrurus adsimilis, which is considered a different species from the Indian bird. There are published studies of Cattle Egret associations in Southern Africa (Blaker 1969, Siegfried 1978), but not of any other species.
Methods
A herd was defined as any group of animals in a finite area seperated by a larger area from other such groups. A bird was regarded as being in association with a mammal if it was within two metres of that mammal regardless of its perch. Where mixed groups of mammals were observed these were divided into species and the corresponding bird species associating with them were recorded. The number of mynas observed on Dal Lake, Kashmir
BIRD/MAMMAL ASSOCIATIONS: CONTRIBUTIONS FROM INDIAN PLAINS AND ZIMBABWE PLATEAU 35
in June 1982 were used for the comparisons of the number of birds in association and not associating. Statistical analyses were performed on the results using students t-tests to determine significant differences. The percentage of all birds (excluding mynas for reasons given above) observed associating, out of the total number observed is also presented. An attempt to see if the number of birds in association with a group increased with increasing group size failed because virtually all groups observed were small (less than five animals). Animal and bird masses are taken from Lander (1949) and Ali and Ripley (1968-1974) respectively. These animal masses are probably larger than the peasant owned stock observed here. These have been used as an estimate of the relative size of the animals and birds concerned.
A similar study was done in Zimbabwe travelling between Bulawayo and West Nicholson. Statistics were not performed for this data but otherwise methods were similar except that for drongos the distance required for association was increased even on occasion to the other side of the road as this bird chases Hying insects which are flushed by hosts.
Results
Eight types of domestic mammals were observed in bird/mammal associations in India, but of these donkeys, sheep, pigs, camels, and horses contributed only a very small proportion of such associations and are not further discussed. A total of about 650 groups each of cattle and water buffalo and nearly 300 groups of goats, giving a total of between 5000 and 6000 head each, were observed.
Table 1 presents the frequency of bird/ mammal associations observed on the plains of India. For each of the three mammals the percentage of groups observed with each avian species is presented. Further, the percentage of each bird (except the myna) observed in such associations is also presented. Table 2 presents the mean number (± the standard deviation) of birds in association with each mammal and those not in such associations
Table 1
FREQUENCY OF BIRD/MAMMAL ASSOCIATIONS ON THE PLAINS OF INDIA (EXPRESSED AS THE PERCENTAGE OF GROUPS WITH ASSOCIATING BIRDS AND THE PERCENTAGE OF BIRDS OBSERVED ASSOCIATING OUT OF THE TOTAL NUMBER OF BIRDS OBSERVED)
|
Bird Species |
House Crow |
Common Myna |
Cattle Egret |
Black Drongo |
|
mass |
300 gm |
1 1 0 gm |
360 gm |
45 gm |
|
cattle (mass 360 kg) |
1% |
7% |
6% |
5% |
|
water buffalo (mass 630 kg) |
3% |
13% |
10% |
7% |
|
goat (mass 70 kg) |
1% |
1% |
1% |
5% |
|
% of birds in association |
1% |
— |
16% |
20% |
are presented. Data are too meagre for such calculations with goats. All these birds are believed to be residents preforming at most local movements in the area. At this time of the year, all four species would be breeding (Ali and Ripley 1968-74): though some species may be represented by nonbreeders for instance the Cattle Egret.
The number of Common Mynas in mammal associations is significantly larger than those not in such associations (t= 2.812, p<0.05). This assumes that flock sizes in Kashmir (altitude 2000 m) are the same as those on the plains. The number of egrets associating and not associating is not significantly different; but the numbers associating with cattle is significantly larger than those associating with water buffalo (t= 1.813, p<0.05). Black Drongos, which are the smallest of the four avian species considered here, is the only one not to show a lower association with the goat groups. The number of drongos in associations is significantly larger than the number of those not in such associations (t=2.116, p<0.05). All species show a preference for the heaviest animal (water buffalo) in terms of the number of groups
36
JOURNAL BOMBAY NATURAL HIST. SOCIETY, Vol. 93(1996)
Table 2
NUMBERS (MEANS ± STANDARD DEVIATIONS) OF BIRDS WITH AND AWAY FROM HOSTS ON THE PLAINS
|
OF |
INDIA |
|||
|
• |
House Crow |
Common Myna |
Cattle Egret |
Black Drongo |
|
Water buffalo |
1.1 ±0.7 |
1 ,6±0.6 |
1 .5±0. 1 |
l.4±0.8 |
|
cattle |
l.3±0.4 |
1 ,6±0.9 |
3.5±2.4 |
1.1 ±0.4 |
|
total bird/mammal |
1 ,4±0.9 |
1.6±0.9 |
— |
1.3 ±0.6 |
|
away from mammals |
1.2±0.6 |
1.3±0.5 |
4.6±2. 1 |
1.1 ±0.4 |
associated with. This makes sense if we assume that the largest animal flushes the most insects or supplies the best perch.
Thirty one records of two bird species and one record of three species associating with the same group of mammals (mostly water buffalo) were observed.
Table 3
FREQUENCY OF BIRD/MAMMAL ASSOCIATION ON THE ZIMBABWE PLATEAU (AS TABLE I )
|
Cattle |
Forktailed |
|
|
Egret |
Drongo |
|
|
Cattle |
6% |
11% |
|
goats |
5% |
0.5% |
|
% of birds |
70% |
40% |
Table 4
NUMBERS OF BIRDS WITH AND AWAY FROM THEIR HOSTS (AS TABLE 2)
|
Cattle Egret |
Forktailed Drongo |
|
|
cattle |
3.5 |
1.2 |
|
goat |
7.2 |
1.0 |
|
total bird/mammal |
4.8 |
1.2 |
|
away from mammals |
1.4 |
1.1 |
Tables 3 and 4 present similar data from the Zimbabwe plateau. A total of 579 herds of cattle and 262 herds of goats were observed giving a lower density for each species and a much lower overall biomass of domestic animals due to the absence of water buffalo. In contrast to India the egret is a passage migrant through the area: passing quickly through around April and not associating much and again in Sept. /Nov. when it does associate. The drongo is a resident but like Indian birds undergoes local movements when not breeding: in Zimbabwe this is believed to be into more open areas (Irwin 1981). Like the Cattle Egret (Blaker 1969 and this study), they associated more in the dry season. The egret appears to aggregate in larger flocks with goats than with cattle, though two large flocks of 16 and 30 were seen with goals in April, a time of year when they occur in larger groups and do not usually associate. No such flocks were seen with cattle at this time of year. The drongo appears to show no variation in numbers with either animal or away from animals.
No records of more than one species associating with the same group were obtained despite the increased rate of association. Does this imply that in India some groups are more attractive to associating birds than others? While the drongo is well known as an aggressive species, it is difficult to believe that any territorial exclusion occurs between the two species and no cases of agression were observed. In any case the drongo takes flying insects whereas the egret takes mainly grasshoppers and they are presumed not to be in competition for food.
Discussion
Dean and Macdonald (1981) have divided bird/mammal associations into those which compete with their host for food and those which use the mammal to obtain their food. The myna and the crow are omnivorous taking some vegetable food and are thus potentially in competition with their hosts. The other two species are purely insectivorous (Ali and Ripley 1968-74). Another possible aspect of these
BIRD/MAMMAL ASSOCIATIONS: CONTRIBUTIONS FROM INDIAN PLAINS AND ZIMBABWE PLATEAU 37
associations is disease transmission but this has not actually been demonstrated (Dean and Macdonald 1981). It is possible that the birds also gain some protection or concealment from predators from these associations. It is not always clear what the mammal gains from these associations though the myna has been recorded as eating ticks in Southern Africa, where it has been introduced (Dean and Macdonald 1981) and presumably do so in India. Another suggestion is that the egret (at least) lakes herbivorous insects, which compete with cattle for food (Dinsmore 1973).
All these bird species feed on insects attracted to or flushed by the mammals for instance, coprophagous beetles. Some use the mammal as a perch though egrets only do so in long grass (Siegfried 1978) and I have never observed it. Animals in woodland are less likely to have attending birds. Drongos will also use a perch on a tree or a fence near the herds for the same purpose. The other species tend to forage on the ground behind the animals though I have seen mynas preening while perched on a mammal.
I doubt if the House Crow gains significant amounts of food from these associations except perhaps by raiding grain bins. Its very ubiquity appears to lead it to perch on or around the mammals especially in the absence of other suitable perches. On the Calcutta Maiden, in the absence of the other suitable mammals, they were observed attempting to perch on goats. The goats however actively attempted to dislodge these birds by running and swinging their bodies until the crows dropped off. Unless the crows were trying to glean ectoparasites, which was not observed, I can only interpret this behaviour as a type of play (McFarland 1981).
Indian Elephants Elephas maximus were observed behaving similarly to dislodge mynas, which appeared to have become too numerous and too noisy, in Mudumalai Reserve in Tamil Nadu.
The crows are highly adaptable: in Calcutta zoo they were observed associating with African mammals, e.g. Eland Tauretragus oryx. Cattle Egret occur in larger groups with cattle but they occur with a larger percentage of groups of the bulkier water
buffalo but the actual numbers of egrets associating with each species are similar. This is in contrast to the situation in Australia where water buffaloes are preferred (Siegfried 1978). This might also be a function of the more aquatic habits of the water buffalo, which appears likely to be the original host in Asia. In Africa the original host is believed to be the buffalo Syncerus coffer, which also prefers flooded grasslands (Siegfried 1 978). The same flock sizes with and away from hosts is in contrast to the South Africa situation where flocks are larger and away from their hosts (in this case only cattle) (Blaker 1 969). Cattle Egrets acquire more prey with less effort when associating with cattle (Dinsmore 1973). Smith (1971) has demonstrated this for two species of anis ( Crotophaga spp.)
The Zimbabwe situation in the Cattle Egret is different again with same size of flocks with cattle as in India, an increased use of goats and much higher percentage of birds in associations. This is believed to be a reflection of very poor feeding conditions due to the succession of drought years which occurred in Zimbabwe during the study. The same flock size in both areas may be a reflection of territorial behaviour.
Benson (1964) reported that drongos were seen associating with five groups of cattle on a journey of 1 50 km in a nearby area of Zimbabwe. The former author never observed such associations by drongos in 30 years in Zambia and Malawi, which have a higher rainfall than the southern areas of Zimbabwe where these birds were observed. There is another published record from Banket (Parnell 1962). Most of these records come from low rainfall areas (less than 600 mm per year). This makes it difficult to understand why drongos in Zimbabwe did not increase in numbers when with their hosts or why they made a much lower use of goats in an area with much lower domestic mammal densities. Yet they doubled their use of cattle groups and the proportion of birds with hosts as compared with India. This suggests that the drongos were regulated by other factors like territorial behaviour. I have also observed a drongo perched on a small shrub near a chicken and apparently associating with it. This bird has been
38
JOURNAL. BOMBAY NATURAL HIST. SOCIETY. Vol. 93(1996)
reported as associating with ostriches Struthio came lens (Dean and Macdonald 1981).
I also observed Glossy Starling Lamprotornis sp. and a Pied Crow Corvus albus once each associating with cattle. It is striking that these four birds represent the same families as the birds seen in such associations as in India. Further in Britain a starling Sturmis vulgaris and a crow Corvus corone are also commonly observed in fields with domestic animals most frequently sheep. Cattle egrets and drongos do not occur in Britain. Are these birds/ families somehow preadapted to such associations more than other families? Or are they birds who have adapted well to farming habitats? The crows at least are believed to have increased with human numbers. Certainly the Cattle Egret has vastly increased its
Re
Ali, S. & S.D. Ripley (1968-1974): Handbook of the birds of India and Pakistan. Oxford University Press, Delhi. Benson, C.W. (1969): Birds associating with ungulates. Auk 8J: 436.
Blaker, D. (1969): Behaviour of the Cattle Egret Arcleola ibis. Ostrich 40: 75-129.
Brooke. R.K. (1984): South African Red data book-birds. S.
Afr. Nat. Scient. Prog. Report No. 10.
Dean, W.R.J. & l.A.W. Macdonald! 1981 ): A review of African birds feeding in association with mammals. Ostrich 52: 1 35- 155.
Dinsmore, J.J. (1973): Foraging success of Cattle Egrets Bubulcus ibis. Am. Midi. Nat. 89: 242-246.
Irwin, M.P.S. (1981): Birds of Zimbabwe. Quest Publishers,
range due to its association with cattle in the wake of their range being increased to include the Americas and Australia (Siegfried 1978).
Bird/mammal assoiations have been very little studied in the tropics though the tremendous increase in the range of Cattle Egrets has generated interest in this species. It is hoped that this contribution will stimulate others to study such associations more closely.
Acknowledgements
Thanks to Mr. M.P.S. Irwin and Dr. K. Hustler for comments on an early draft, Mr. K. Ratcliffe for assistance with statistical analysis and to Mr. T. Ewbank for the use of a computer.
ERENCES
Harare.
McFarland, D. (Editor) (1981): The Oxford companion to animal behaviour. Oxford University Press, Oxford.
Lander, P.F. (1949): The feeding of farm animals in India. Macmillan, London.
Parnell, G.W. (1962): Birds following cattle. Honeyguide 39: 1-2.
Robinson, W.C. & E.G. Bolen (1989): Wildlife ecology and management. Collier McMillan Publishers, London. Siegfried, W.R. (1978): Habitat and modern range expansion of the Cattle Egret. National Audubon Report No. 7.
Smith, S.M. (1971): The relationships of grazing cattle to foraging rates in anis. Auk 88: 876-880.
FISH FAUNA OF PERIYAR TIGER RESERVE1
V.J. Zacharias, A.K. Bhardwaj and P.C. Jacob2
Key Words: fishes, Periyar Tiger Reserve, Kerala, India
The status and distribution of fishes in the rivers and the lake of Periyar Tiger Reserve, Kerala was studied and 35 species belonging to 7 orders and 1 1 families were recorded. The family Cyprinidae contained maximum number of species ( 1 3). Thirteen species of fishes collected during the study are endemic to Southern Western Ghats. Two new species, namely Lepidopygopsis typhus (Schizothoracinae) and Cmssocheilus pehyurensis (Cyprinidae) were recorded from Periyar while another, Echathalakanda ( Barbus ) opliiocephalus (Cyprinidae) was rediscovered from Periyar river.
Introduction
The rivers in Kerala once harboured a rich fish fauna according to earlier investigators like Pillay ( 1 929 ), John ( 1 936), Hora (1941 a,b), Raj (1941 a,b), Chacko (1948), Silas (1951a, b). But very little information is available on the present status of the freshwater fishes of Kerala, which are threatened by over exploitation, introduction of exotic fishes, habitat destruction and pollution. In the midlands and lowlands of Kerala many fishes have become locally extinct and are disappearing fast. A limited number of them remain in the hills; in protected areas.
Periyar is known to support several interest- ing and important fishes. Some preliminary studies were done on the fish fauna of Periyar (Raj 1 94 1 a,b; Chacko 1948, Silas 1951a, b). Very little is known about the current status offish fauna in this reserve. Chacko, made an attempt to make a survey of the indigenous fish fauna in 1 946, with a view to develop the fishery. He listed 35 species of fishes in the lake. Raj (1941a) has described a small scaled schizothoracine, Lepidopygopsis typus Raj, from the Periyar river and Hora (1941a) has des- cribed, from Mr. Jone’s collection a Homalop- terid loach, Travancoria j one si, from Travan- core.
Menon and Jacob (1991) have more recently 'Accepted August 1995.
-Periyar Tiger Reserve. Thekkady, Kerala 685 536.
described a small scaled Barbel, Cmssocheilus periyarensis and rediscovered a Cyprinid fish Barbus ( Puntius) opliiocephalus (Raj) from the Periyar river adding two more species of fishes to the fish fauna of Periyar Tiger Reserve.
The purpose of this paper is to present a status report of the fish fauna of Periyar Tiger Reserve for making comparative studies in future. It will also be helpful to identify the conservation problems and recommend management measures.
c
Study area
Periyar Tiger Reserve lies between 9° 16' and 9° 40’ N. lat. and between 76° 55’ and 77° 25' E. long. It is bordered by Kottayam and Pathanamthitta districts in the west and south, Peermade Taluk of the Idukki District in the north and Madurai district of Tamil Nadu in the east. The elevation of the Reserve ranges from 800 to 2019 m. Several peaks rise above 1600 m the prominent peak being Vellimala (2019 m).
River, Periyar which originates from Chokkampetti-Kallimalai side, about 58 Km from Thekkady with its various tributaries form the main drainage of the area. The lake, which was formed as a result of the construction of the dam has an area of 26 sq. km. Maximum depth of water at highest watei level is 46 m. Two other rivers, Pamba and Azhutha also flow along the border of the reserve in the Vallakkadavu range.
40
JOURNAL, BOMBAY NATURAL HIST. SOCIETY, Vol. 93(1996)
Table 1
FISH FAUNA OF PERIYAR TIGER RESERVE
|
Name of (he Species |
Local/Tribal Name |
Locality |
Relative abundance |
General distribution |
|
Order CYPRINIFORMES Family Cyprinidah Subfamily cyprininae 1. Cyprians carpio communis |
||||
|
(Linn.) |
Goldfish |
Lake only |
Common |
Exotic species |
|
2. Punt ius melanostigma (Day) |
Kudukunda |
River & Lake |
Very common in lake |
India; Hill streams of Kerala, Nilgiris. |
|
3. Pun tins melanampyx (Day) |
Kudukunda |
River & Lake |
Very common in lake |
India; Hill streams of Kerala, Nilgiris. |
|
4. Punt ms (impliibius (C. & V.) |
Kooral |
Azhutha, Pamba |
Common |
India; Peninsular India, Sri Lanka. |
|
5. Punlius nuthecoUt (Val.) |
Karuva/Paral |
n n |
ii |
ii |
|
6. Puntius filameiitosus (Valenciennes) |
- |
11 11 |
ii |
|
|
7. Echatlialakanda opliioceplialus (Raj) |
Eetilakanta |
River above Rare Mlappara Ummikkuppanthodu |
India; Kerala Hill stream. Rediscovered from Mlappara. |
|
|
8. Tor khudree (Sykes) |
Kuyil |
River & Lake |
Common |
India; Deccan and P. India; Sri Lanka. |
|
9. China mullya (Sykes) |
Kallemutti |
River & Reservoir Common |
India; Hill streams of P. India. |
|
|
10. Garni gotyla steiiorliynclius (Jerdon) |
ii |
n ii |
ii |
India; Western Ghats, Cauvery &. Krishna drainages. |
|
1 1 . Hypsilobarbus kurali (Menon & Rema Devi) |
Kooral |
Stream & Lake |
ii |
India; Dakshina Kannada to Travan core Hills. |
|
12. H. periyarensis (Raj) 13. Crossoclieilus periyarensis |
Kariyan Karimbachy |
Fast flowing rivers River above |
ii |
India; Kerala, Periyar. |
|
(Menon & Jacob) |
Thannikudy |
Rare |
ii |
|
|
Subfamily schizothoracinae |
||||
|
1 4. Lepidopygopsis typus Raj |
Brahmana Konda |
Periyar above river |
India; Kerala, Periyar river system. |
|
|
Subfamily rasborinae |
||||
|
15. Parluciosoma daniconius (Ham.) |
Kannanjon |
Lake, Azhutha stream |
Very common |
India; Sri Lanka. Pakistan, Nepal, Bangladesh, Myanmar (Burma), Thailand. |
|
16. Barilius bendelisis (Ham.) |
Pavukan |
Thannikudy, Lake, River |
Common |
India; Pakistan, Nepal, Bangladesh. |
|
17. B.bakeri Day |
ii |
India; Kerala, W. Ghats. |
||
|
18. B. I'atensis (C. & V.) |
i i |
ii |
ii |
Peninsular India, W. Ghats. |
|
19. Danio aequipinnatus (McClell.) |
Azhutha, Pamba |
Very common |
India; Nepal, Bangladesh, Myanmar, Thailand. |
|
|
Family Homalopteridae |
||||
|
20. Travancoria jonesi Hora |
Kallotty |
Rivers above Thannikudy |
Rare |
India; W.Ghats, High Ranges of Kerala. |
|
Family Cobitidae |
||||
|
21. Lepidocephalus thermalis (C. & V.) |
Ayira |
Lake/River |
Common |
P. India; Sri Lanka. |
|
22. Noemaclieilus botia |
Ayira |
Small streams |
Rare |
Hill stream of Travancore. |
|
23. N. triangularis |
n |
ii |
” |
India; W. Ghats. |
FISH FAUNA OF PERIYAR TIGER RESERVE
41
Table 1 (Contd.)
|
Name of the Species |
Local/Tribal |
Locality |
Relative |
General |
|
Name |
abundance |
distribution |
||
|
24. N. evezardi |
Ayira |
Small streams |
Rare |
India; W. Ghats and Madhya Pradesh. |
|
Order SILURIFORMES Family Heteropneustidae 2 5 . Heiewpneustes fossil is |
Kary (Bloch) |
Lake |
Very Common |
India; Pakistan, Sri Lanka, Nepal. |
|
Family Siluridae 26. Ompok himaculatus (Bloch) |
Chottavala |
Lake |
India; Pakistan, Nepal, Bangladesh, Myanmar, Thailand, Java. Sumatra, Borneo. |
|
|
Family Sisoridae 27 .Glyptothomx madnispatanwn |
Parayotti (Day) |
Thannikudy |
Uncommon |
India; W. Ghats. |
|
Order ANGUILLIFORMES Family Anguillidae 28. Anguilla bengalensis |
Mlanjil |
Azhutha |
Uncommon |
India; Pakistan, Sri Lanka, |
|
Myanmar. |
||||
|
Order ATHERINIFORMES |
||||
|
Family Cyprinodontidae 29. Aplocheilus lineatus lineatus |
Poonjan |
River & Lake |
Very Common |
Peninsular India. |
|
Order FERCIFORMES Family Cichlidae 30. Oreochmmis mossambica (Peters) |
Thilapi |
Lake |
Very common |
Introduced. |
|
Family Channidae 31. Channa striata (Bloch) |
Varal |
Lake |
Rare |
India; Sri Lanka, Pakistan, Bangladesh, Nepal, Myanmar, Malaya, Malaya Archipelago, Thailand up to Philippines. |
|
32. C.orientalis (Bloch & Schn.) |
Vatton |
Lake & Rivers |
11 |
India; Iran, Afganistan, Nepal, Pakistan, Sri Lanka, Bangladesh, Myanmar, Thailand, Yunan, Malaya, Malay Archipelago, Hainan, and Taiwan. |
|
33. C. main tins |
Cherumeen |
Azhutha, Pamba |
n |
India; Pakistan, Sri Lanka, Bangladesh, Nepal, Myanmar, Thailand, Sumatra, Borneo, China. |
|
Order MASTACEMBELIFORMES Family Mastacembelidae 34. Mastacembelus anuatus (Lacep.) |
Aaron |
River, Lake |
Common |
Pakistan, Sri Lanka, Nepal, Thailand, India, Myanmar, Malaya to South China. |
|
35. Macro gnat bus oral |
ii |
Vazhukkappara Rare |
India; Pakistan, Sri Lanka, |
|
|
(Bloch & Schn.) |
stream |
Vietnam, Bangladesh, Nepal. Myanmar, Thailand, Laos, Malaya and East Indies. |
Materials and Methods Fish samples were collected from January, 1992 to December, 1994 from different localities in the rivers and lakes while conducting wildlife studies.
The collections were made from the Periyar river, Mullayar river, and their tributaries and different areas of the reservoir; boat landing, near dam, Mullakkudy, Manakkavala, Swamikkayam and
42
JOURNAL BOMBAY NATURAL HIST. SOCIETY, Vol. 93(1996)
Padikkayam. Fishes were also collected from rivers Pamba and Azhutha.
Fishes were collected by gill-nets, cast-nets, hooks and bait. Worms, grasshoppers, small fishes, fruits of some trees, boiled tapioca and even rice paste were used as baits. For collecting small fishes, a special method called ‘Watty” was used. Some were collected from local fisherman. Fishes were preserved in 5% formalin and identified in the laboratory.
Results and Discussions
Thirtyfive species of fishes, representing 21 genera and 1 1 families were collected and identified (Table 1). Morphological particulars of these species are available in Day (1876-88), Flora ( 1 94 1 a,b), Silas ( 1 95 1 a,b), Munro (1955), Talwar and Jhingran (1991). Out of the 33 species mentioned in the list of Chacko (1948), species YikeMystus cavasi us (Ham.), M. vittcitus (Bloch), Notopterus notopterus (Pallas), etc., were neither found in the reservoir nor in the tributaries of Periyar and Pamba during this survey.
The species Parluciosomci dan icon ins (Ham.), Puntius melanamphyx (Day), Hypsilobarbus kurcdi (Menon & Rema Devi), Garni mullya (Sykes) were collected from almost all localities of the reservoir. Among these 35 species, about 1 3 species are usually found in upper streams and are adapted to lotic torrential waters. They are Travancoria jonesi (Hora), Garra mullya (Sykes), Garni gotyla stenorhynchus (Jerdon), Tor khudree (Sykes), Hypsilobarbus periyarensis (Raj), Crossocheilus periyarensis (Menon & Jacob), Barbus ophiocephalus (Raj), Lepidopygopsis typus (Raj), Noe mac he i l us evezardi (Day), Glyptotliorax madraspatanum (Day), Barilius bakeri (Day), B.bendel is is (Ham.) and B. gatensis (Cuvier & Val.). Some of them are adapted to cling to the substratum by some attachment devices, e.g. Garra, Travancoria and Glyptotliorax.
Fishes like Heteropneustes fossilis (Bloch), Ompaok bimaculatus (Bloch), Channel striatus (Bloch), C.orientalis (Bloch & Schn.), C.marulius
(Ham.), Cyprinus ccirpio communis (Linn.), Oreochromis mossambica (Peters), etc., are restricted to lentic waters of the reservoir.
Among the hillstream fishes Crossocheilus periyarensis (Menon & Jacob), a rare species was a new discovery from Periyar. Another fish which was believed to be extinct, Barbus ophiocephalus (Raj) was rediscovered. Lepidopygopsis typus (Raj), Hypsilobarbus kurcdi (Menon & Rema Devi), Tor khudree (Sykes), Travancoria jonesi (Hora) are some of the endemic species of the Southern Western Ghats. Preliminary studies on the food habits of Tor (through stomach content examination) have indicated that this species took a variety of fruits.
Conservation and Management
“Mannan” and “Paliyans” (local tribals) above forty years of age speak of abundant fish in all the rivers, especially in Periyar and in the reservoir in the past. According to them, this abundance was due to undisturbed conditions of Periyar, Mullayar and their tributaries. But now all these areas are disturbed by indiscriminate fishing, deforestation, hunting, etc. In addition, new exotic fishes which were introduced to the reservoir, namely Oreochromis mossambica (Peters) and Cyprinus carpio communis compete with native species of fishes for food and habitat.
Recommendations
1. Fishing activities in Periyar should be controlled.
2. Fishing during monsoon, which is the breed- ing season of most of the fishes should be banned.
3. Research should be conducted for assessing population density and habitat requirements ol fishes in the rivers and lake.
4. Remove the introduced fishes from the reservoir and restock with fingerlings of species like Tor khudree (Sykes), which has sport value.
5. Sport fisheries could be developed to cater to
FISH FAUNA OF PERIYAR TIGER RESERVE
43
the needs of tourists in the tourist zone which could generate revenue for the Government.
6. Puntius melanampyx could be used as an aquarium fish.
7. Plant trees on the lake edges for providing food for species like Tor.
Refer
Chacko, P.l. (1948): Development of Fisheries of the Periyar lake.
./. Bombay not. Hist. Soc. 48: 191-192.
Day, F. ( 1 886- 1 888): Fishes of India, London.
Hora, S.L. (1941a): Homalopterid fishes from peninsular India. Rec. hul. Mus. 43: 230-23 1 .
Hora, S.L. ( 1941b): The Freshwater fish of Travancore. Rec. hul. Mus. 43: 234-256.
John, C.C. (1936): Freshwater fishes'and fisheries of Travancore.
./. Bombay nut. Hist. Soc. 38: 702-783.
Mf.non, A.G.K. & PC. Jacob (1996): Crossoclieihis perivarensis, a new Cyprinid fish from Thannikudy (Thekkady), Kerala,
S. India. ./. Bombay nut. Hist. Soc. 93( I ): 62-64.
Monro, I.S.R. (1955): The marine and fresh water fishes of Ceylon. Canberra (Dept. External Affairs), pp. 351 .
Pillay, R.S.N. ( 1929): A list of fishes taken in Travancore from
Acknowledgements
Wc arc grateful to Dr. A.G.K. Mcnon for his suggestions and constructive comments on an early draft of this paper. We thank Dr. M.P. Thobias for his suggestion and help. Sri C.P. Shaji and Sri P. Muhammed Jafer assisted in the collection of specimens.
[•: n c f s
1901-1915. J. Bombay uat. Hist. Soc. 33: 347-379.
Raj B.S. (1941a): A new genus of Schizothoracine fishes from Travancore, South India. Rec. hul. Mus. 43: 209-214.
Raj, B.S. ( 1941b): Two new Cyprinid fishes from Travancore. South India, with remarks on Barbus (Puntius) micro/ioyon. Cuv. & Val. Rec. hul. Mus. 43: 375-386.
Silas E.G. (1951a): On a collection of fish from the Anamalai and Nelliampathi Hills ranges (Western Ghats) with notes on its zoogeographical significances. ./. Bombay not. Hist. Soc. 49: 670-681.
Silas E G. (1951b): Fishes from High Range of Travancore. J.
Bombay uat. Hist. Soc. 50: 323-330.
Talwar, PK. & A.G. Jiiingran (1991): Inland Fishes of India and adjacent countries: Vol. 1 & 2. Oxford & IBH Publishing Co. Pvt. Ltd.
PRELIMINARY OBSERVATIONS ON THE IMPORTANCE OF A LARGE COMMUNAL ROOST OF WINTERING HARRIERS IN GUJARAT (NW. INDIA) AND COMPARISON
WITH A ROOST IN SENEGAL (W. AFRICA)1
Roger Clarke2
Key words: Gujarat, locust, Montagu’s Harrier, Pallid Harrier, pellets, roosting
The largest communal roost of wintering harriers in the world reported in recent times occurred in the grassland at Velavadar National Park, Gujarat, north-west India. It consisted of up to 2000 birds, mainly Montagu's Harriers Circus pygargus (about 75%) and Pallid Harriers Circus macrourus (about 20%) and has been known since the mid 1980s. Pre-roosting, roosting and post-roosting behaviours are described. About 30% of the harriers present were adult males, but only one dark morph Montagu’s Harrier was recorded. Foraging behaviours are described; the Montagu’s Harriers fed mainly on locusts in shrublands and cotton fields and male Pallid Harriers were observed hunting small birds in grassland. Of 134 pellets collected at the roost, 60% contained locust remains, principally of the Tree Locust Anacridiuni rubrispinum. It was calculated that the harriers attending the roost probably consumed more than 1 .5 million locusts each winter. The rest of the prey remains in the pellets were mainly of small birds, principally larks, although the remains of a few mammals and reptiles also occurred. The roost is compared to one found in Senegal, West Africa.
Introduction
More than 1 500 Hamers were counted roosting at Velavadar Blackbuck Sanctuary and National Park, Gujarat, north-west India in November 1991 (W.S. Clark in lift.). This appears to be the largest roost of harriers recorded in the literature since nineteenth century observations of thousands of Montagu’s Harriers roosting after the breeding season and before migration at a marsh in the west of France (Barbier Montault 1838). The great majority of harriers at Velavadar were Montagu’s Harriers Circus pygargus (about 75%) and Pallid Harriers C. macrourus (about 20%), but very few Marsh Harriers C. aeruginosus and one or two ringtail Hen Harriers C. cyaneus were also present. This paper details observations during two visits to Velavadar, on 1-6 February 1992 and 25- 31 January 1993. On my first visit I collected 134 pellets from two settling areas which were attracting about 500 and 300 birds respectively at the time. The results of my analysis of the pellets and observations of foraging and roosting behaviour are given below and compared with observations on harriers and
'Accepted June 1993.
’New Hythe House, Reach, Cambridge CB5 OJQ, U K.
the results of analysis of 1 1 3 pellets collected at a roost of about 1000 Montagu’s Harriers in late December 1988 and early January 1989 near M’bour, Senegal, West Africa (Cormier and Baillon 1991).
Roost Catchment Areas
The Indian roost site (22° North) is in a remnant of grass plain occupying the northern half of 17.38 sq. km of land preserved as a National Park in 1976 to conserve the Blackbucky4/zf//oy?c cervicapra. The Park is situated in a semi-arid area of alluvial plain known as the ‘Bhal’ (reputed to mean ‘forehead’, i.e. a bare, open landscape), on the western shore of the Gulf of Khambhat (Arabian Sea). Some land between the Park and the Gulf of Khambhat (20 km away) is a saline wasteland irregularly inundated by the sea in the monsoon, but much of the surrounding plain consists of shrublands of Mesquite Prosopis chilensis and large arable fields, in winter, mainly growing a special strain of cotton not requiring irrigation. A high proportion of the fields were ploughed at the time of my visits and cotton was being harvested. In the day, many Montagu’s Harriers were observed hunting over the cotton fields.
LARGE COMMUNAL ROOST OF WINTERING HARRIERS IN GUJARAT
45
The African roost site (14° North) is situated close to the Atlantic coast in fairly flat, open savannah with abundant ground vegetation and a few trees. The harriers hunted around brackish lanoons, in savannah and dunes, not over crops.
Roost Sites
The Indian Harriers roosted in aDichanthium- dominated grass community about 40 cm tall in large, totally open fields which had been used as hay plots and were now maintained just by shrub clearance, although bordered by hedges of shrubs, especially P l o s op is cli Hen s is .
The African roost was in ground vegetation in an area of savannah with some trees, principally Ziziphns niciuritiana, and with scattered humps of old termite mounds 1-1.5 m high. Domestic grazing herds, mainly of goats and cattle, crossed the site by day, but the ground vegetation was still quite dense.
Pellet Analysis Methods
For the purposes of this paper, the term ‘locust’ includes large grasshoppers. Pellets from India containing only locust or a mix of locust and reptile remains were dissected dry. Pellets at least partly made up of bird or mammal remains were dissected wet and the remains were washed and allowed to dry. Pellets of pure locust remains were very fragile and liable to break in two on the ground or when collected. The collection comprised 46 part pellets, which were counted as 23 whole pellets, and 1 1 1 whole pellets to give the equivalent of 1 34 pellets. The locusts were identified from the manual issued by the Centre for Overseas Pest Research (1992) and specimens in the collection of the Bombay Natural History Society. The number of locusts in each pellet was taken as the highest number represented cither by mandibles or by ovipositor valves. Care had to be taken to identify dorsal and ventral pairs of ovipositor valves to arrive at the correct number of female locusts that they represented. Counts of locusts from mandibles were based on the highest number of right or left mandibles in the pellet. Bird remains were identified by matching against reference collections. Larks were counted by
means of bill parts or hind claws. Pigeons and doves were recognised from their white down and feather fragments. Sparrows were recognised and counted by the palatal thickenings from their bills. Bird remains which could not be identified were counted as one bird per pellet, but were few in number. Reptiles could not be enumerated or identified since remains were almost entirely loose scales. One pellet contained a lizard jaw. Mammals were identified from their fur and teeth. Indian Bush Rat Golunda ellioti and Gerbil Tatera indica teeth were identified from the early, but accurate illustrations in Blanford (1888). Mice and rats were identified from hair, jaws or incisors. Lagomorph fur was confirmed by microscopic examination of the medulla (Koppikar and Sabnis 1976).
Observations
Roosting, pre-roosting and post-roosting behaviour: The number of harriers using the Velavadar roost fluctuated during each winter. In both 1991 and 1992, large counts ( 1500-2000) were made in November/December. Later in each winter, numbers dropped in January /February (800 birds in early February 1 992 and 600 declining to 300 in late January 1993). The full seasonal pattern is yet to be established, but it seems likely that the roost is reduced in size from midwinter because birds pass south before coming back on return migration. Further large roosts are known or suspected south to Andhra Pradesh (southeast India) and research is required to ascertain whether they peak later (A. Mulchandani, pers. comm.). Migration of harriers from the direction of Gujarat through the Western Ghats has been observed in the past (Khacher 1 977).
The Indian harriers were using at least two pre- roosting areas of bare, flat, dried mud, separated from the night-roost grassland by Prosopis chilensis thickets. Up to 126 harriers (26 January 1993) were counted on the largest of these. Only a proportion of the harriers seemed to be using them, since a constant stream of harriers passed by, heading for the grassland night-roost. The pre-roosting harriers stood facing into the wind, well spread out in loose groups. The mud was generally very flat, but many individual
46
JOURNAL BOMBAY NATURAL HIST. SOCIETY. Vol. 93 (1996)
harriers were noted perching on the slightest lumps projecting from the surface. Numbers built up until about sunset. After sunset, these pre-roosts would gradually disperse, individuals and small groups flying off to the night-roost.
Especially with a good breeze, aerial activity at the night-roost was on two levels. The number of birds ‘milling’ above the night-roost quickly built up into a mass of up to several hundred birds towering one hundred metres or more above the settling area. At the same time, a significant number was positioned much higher in the sky, using the warm air until well after sunset and then individuals folded in their wings to stoop down to lower levels, criss- crossing each other’s flight paths in the sky, to join the roost. Occasionally a separate tight ‘carousel’ of circling birds would form. The cause of these could not be ascertained at the time, but subsequent observations suggest that they are a mutual warning mechanism alerting birds to predators in the grass. On some evenings the harriers formed up to three groups of birds milling in the air, which roosted in separate parts of the grassland. Settling by hundreds of birds took place quite rapidly at about twenty minutes after sunset and with some hesitation over places, switching of places or displacement of one harrier by another.
The Velavadar harriers dispersed from the roost at an early dawn. By sunrise almost all had gone, principally in flight-lines south to the main post-roost and west to the main cotton fields and shrublands. Male Pallid Harriers (grey males only identified with certainty in the poor early morning light) left in the same direction as the other harriers, but on average a few minutes later than male Montagu’s Harriers, and generally at lower altitude.
The largest pre-roosting area was watched on the morning of 31 January 1993 and at its peak, 81 harriers post-roosted on the ground there. They stood around in exactly the same manner as at pre-roost, preening a little and eventually flying off before sunrise. A dispute over food between 3 or 4 individuals was noted, involving some chasing low over the ground.
The African harriers (Cormier and Baillon 1991) are described as arriving at the roost site in the one and a half to two hours up to dusk, the first ones continuing to hunt over the site and settling on mounds or trees to deal with the prey caught, then taking flight to settle a little later. Communal aerial activity is described as the formation of as many as two or three simultaneous carousels of hundreds of birds each night at about 1 5-20 minutes before dusk as they took flight in alarm, which subsided before the harriers generally took to the air prior to finally settling. No observations of morning dispersal are given.
Proportions of grey males and dark morphs: Cormier and Baillon (1991) recorded only 11% grey males in Senegal, but made the point that this was probably the result of differing preferences of males and females for certain wintering areas. At Velavadar,
I recorded an average of 30.5% adult grey males, Montagu’s and Pallids combined, from counts (n =
II counts, 397 birds in total) at the pre-roosts. Successive counts were made at each pre-roost to attempt to average out the effect of any difference in arrival and departure times of the sexes. For example, on 26 January 1993 at 1 807 hours there were no grey males out of 16 birds, but 17 out of 35 birds at 1841 hours. On the January 1993 visit, I noted that a few of the juvenile Montagu’s harriers present showed signs of moult into adult male plumage with grey heads, throats and upper breasts. To my knowledge, just one dark morph harrier has been seen at Velavadar — a totally dark brown female Montagu’s Harrier observed once in flight towards the roost in January 1993 (R. Naoroji, pers. comm.). This is in contrast to the situation in Senegal where 5% of the Montagu’s Harriers present were dark morphs (Cormier and Baillon 1991).
Foraging behaviour: The great majority of harriers observed foraging over the cotton fields close to Velavadar were Montagu’s Harriers. Each one hunted intensively and alone over a field or two, flying 3-4 m above the crop looking down into it for locusts.
through the crop intently to see any. Strike rate success was casually assessed as on average once every 10-15 minutes. Strikes were usually a feet-first descent with
LARGE COMMUNAL ROOST OF WINTERING HARRIERS IN GUJARAT
41
wings upraised, but shallow stooping was observed once. The rows of cotton plants were about Ini apart and the harrier would momentarily disappear amongst them. After a successful strike, they flew with the locust held firmly at each end, clearly visible in lowered talons and sometimes in tandem much in the manner of an Osprey Pandion haliaetus carrying a fish. To eat the locust, they flew to an open piece of ground such as a ploughed field or a trackway. Occasionally they fed on the locust in flight, bill and talons being brought together to meet, in the manner of a Hobby Fcilco subbuteo feeding on insects. There was a lull in hunting activity during the heat of midday, when the harriers tended to circle up in the sky in ones and twos.
The few harriers I saw on the Velavadar grassland during the day were adult male and juvenile Pallid Harriers. They mostly hunted earlier and later in the day, but also at midday if it was overcast. Whilst it was warm, they flew slowly into the wind with 3-4 shallow flaps between each glide, then turning to drift quickly downwind, and repeating. In early morning and evening I saw fast low-level flight, with agile swerves at small birds, reminiscent of the fast, low bird-hunting flight mode of the Hen Harrier (Wassenich 1968). One stoop at potential bird prey on the ground from a few metres height was observed (an unsuccessful strike). I saw one of the adult male Pallid Harriers in fast, determined level chases of small birds that he had flushed or missed on a strike, clearly with some expectation of success.
Pellet Analysis Results
The principal prey in the pellets from both continents (Tables 1 and 2) were locusts, in Senegal predominantly the Desert Locust Schistocerca gregaria, and in Gujarat the Tree Locust Anacridium rubrispinum. Locusts featured in 97% of African pellets and 60% of Indian pellets respectively. In the African pellets, Cormier and Baillon (1991) found that ovipositor valves of female locusts greatly outnumbered mandibles. They commented that the male sub-genital plates were difficult to detect and
Table I
PELLETS CATEGORISED BY PREY CLASSES
|
Gujarat n pellets |
% |
Senegal* n % pellets |
|
|
Locust only |
54 |
40 |
84 74 |
|
Locust & bird |
6 |
4 |
3 3 |
|
Locust & mammal |
8 |
6 |
21 18 |
|
Locust & reptile |
9 |
7 |
2 2 |
|
Locust, bird & reptile |
3 |
2 |
|
|
Locust, mammal & reptile 1 |
1 |
||
|
'Absence of locusts' |
3 3 |
||
|
Bird only |
32 |
24 |
|
|
Bird & mammal |
3 |
2 |
|
|
Bird & reptile |
6 |
4 |
|
|
Bird, mammal & |
|||
|
reptile |
2 |
2 |
|
|
Mammal only |
8 |
6 |
|
|
Mammal & reptile |
2 |
2 |
|
|
Total |
134 |
113 |
* Cormier & Baillon (1991).
so females greatly predominated in the analysis, although they give no hard figures. Only females could be identified in the Indian pellets, since no male sub-genital plates were evident. However, pairs of mandibles usually substantially outnumbered the count of females in pellets, based on ovipositor valves, contrary to the African results.
Cormier and Baillon (1991) concluded that the harriers in Africa often did not eat the heads of locusts, preferring the content of the abdomen. The predominance of mandibles in the Indian pellets suggests that this was not the case in India. However, remains collected from one ‘plucking’ place consisted mainly of wings representing about 6 locusts, 5 pronotums, 7 whole and 3 part hind-legs femurs (6 with the rest of the leg attached), and 3 heads. All remains were that of Tree Locusts apart from one Black-spotted Grasshopper Cyrtacanthacris ranacea hind- wing.
Of the bird prey, larks could not be identified to species. Pigeons and doves identified in the pellets included Blue Rock Pigeon Columba livia, Collared Dove Streptopelia decaocto, Little Brown Dove S. senegalensis and Green Pigeon Treron phoenicoptera.
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JOURNAL, BOMBAY NATURAL HIST. SOCIETY, Vol. 93 (1996)
Table 2
PREY IDENTIFIED IN INDIAN AND AFRICAN PELLETS
|
Gujarat n |
Senegal* n |
|
|
Desert Locust Schistocerca gregaria |
1355 |
|
|
Tree Locust Anacridiutn rub rispi mini Other locusts or grasshoppers India — Black-spotted Grasshopper |
614** |
|
|
Cyrtacanthacris ranacea |
68** |
|
|
Africa — (Acrida sp.) |
167 |
|
|
Other insects (mostly Coleoptera) Birds |
55 |
|
|
Larks |
47 |
|
|
Pigeons and doves |
7 |
|
|
Sparrows Passer spp. |
4 |
|
|
Unidentified |
18 |
5 |
|
Mammals |
||
|
Indian Bush Rat Golunda elliotti |
12 |
|
|
Rat Rattus spp. |
2 |
|
|
Mouse Mas spp. |
1 |
|
|
Indian Gerbil Tat era indica *** |
3 |
|
|
Indian Hare Lepits nigricollis |
1 |
|
|
Unidentified |
6 |
23 |
|
Reptiles |
||
|
Agamas Unidentified — number of pellets containing remains (unquantifiable as to |
2 |
|
|
number of individuals) |
23 |
* Cormier & Baillon 1991.
** Total count of 682 locusts from mandibles, etc. (see Methods) apportioned according to the ratio of the pronotums of the two species found in the pellets (209:23).
*** Possibly Indian Desert Gcrbil Meriones hurrianae.
Discussion
The Senega] roost materialised in response to the largest explosion in the population of the Desert Locust in the area for 20 years, with large swarms south of Dakar. In contrast, the Velavadar roost has been known since 1984 (S.Rooke, pers. comm.) and has recurred each winter. The economic value of such roosts can be measured firstly in terms of the number of locusts eaten and secondly in their wildlife tourism potential. A tentative calculation of the number of locusts taken from the surrounding fields by the Velavadar roost in each of the winters might be based
on an average of 750 harriers (mean from pattern of nil at beginning, 1 500 peak, nil at end) for 1 82 days (October to March) consuming 10 locusts on average (calculated from 682 locusts /1 34 pellets in my sample = 5 x say 2 pellets per day) = 1.365 million. This can be regarded as an underestimate, since the pellet analysis probably significantly undercounts the number of locusts eaten because some heads are discarded and locust pellets disintegrate faster probably reducing the number collected. Calculated another way, 750 harriers on average x 75% (Montagu’s Harriers) x 1 82 days x 2, 2-hour hunting sessions producing 8 locusts each (one every 15 minutes) = 1.64 million. Without further research these figures are crude estimates, but the true figure is probably in excess of 1.5 million. This is one measure of the worth of a protected grassland.
Five locusts were caught in the cotton fields close to Velavadar by M. Pai, three of which were Black-spotted Grasshoppers and two Tree Locusts. Despite the tiny sample, this was a surprising result in view of the scarcity of the Black- spotted Grasshopper in the harrier pellets and prey remains examined (about 10% of locusts). It begs the questions: 1. Were more of the boldly-patterned creamy-yellow and black Black-spotted Grasshoppers caught because they are more obvious to the human eye than the uniformly greyish-pink Tree Locusts? or 2. Do the harriers select Tree Locusts — cither for palatability or for ease of capture? Further research is required to answer these questions.
The broader food niche of the Velavadar roost is of course to be expected because of the range of harrier species there. Considering the Senegal results, it might be assumed that the pellets of locust remains at the Indian roost were mainly those of the Montagu’s Harriers present, and this could be largely correct. Observation of the birds leaving the roost in the morning emphasised the difference in the flight actions of at least the males of the two principal species. The Montagu’s Harriers were able to leave at an early dawn, intermittently flapping gently and gliding out towards the croplands on their relatively larger wings, whereas the Pallid Harriers left later and at lower altitude. Sehipper (1977) found that
LARGE COMMUNAL ROOST OF WINTERING HARRIERS IN GUJARAT
49
Montagu’s Harrier ranged further from the nest than other species of harrier sympatric with it in western Europe and obtained enough return for hunting effort from smaller prey. This appears to be because of its light wing loading (Nieboer 1973). Locust prey therefore suits Montagu’s Harrier well despite its small size and its quantity provides the biomass necessary to attract maximum number of birds to one area. This was not only the case at Velavadar and in Senegal, but ‘grasshoppers’ were also the principal prey when Montagu’s Harriers roosted in thousands in France in the nineteenth century (Barbier Montault 1838).
I suspect that the Pallid Harriers took the largest share of bird prey at Velavadar. The Pallid Harrier is larger than Montagu’s Harrier. It exhibits a greater degree of reversed sexual size dimorphism, has proportionately larger feet and claws, and shorter but proportionately more tapered wings (Nieboer 1973). These adaptations point to feeding on birds. Size dimorphism is generally greater in bird-eating raptors (Newton 1979) and larger feet assist in grasping manoeuvrable prey. The slimmer wing structure of the Pallid Hamer favours swifter flight for chasing birds. There is relatively little information on the diet of the Pallid Harrier on its breeding grounds in the steppes of western and central Asia. Early information indicated that the majority of breeding season food was small mammals (80% according to pellets analysed — Osmolowskaja in Glutz von B lotzheim et al. 1971), but birds have recently been found to constitute an important part of the breeding season diet, especially when rodents are scarce (Davygora and Belik 1994).
Apart from larks, the Velavadar grassland itself appeared to hold little prey for the harriers. This is in contrast to the situation in Andhra Pradesh (southeast India), where Rahmani and Manakadan (1986) found grasshoppers so abundant that they flushed a few at every step, and ‘during the day, fifty to sixty harriers... tirelessly quartering the grassland’ at a roost of 800- 1000 harriers (mainly Montagu's) in the 1985-86 winter. Similar findings were made by Satheesan and Rao (1990) who identified both large and medium- sized grasshoppers consumed.
The Marsh Harrier is the largest species of harrier, with the shortest wings and tail relative to body size (Nieboer 1973), and so some of the larger prey items in the Indian pellets, such as hare, might be attributed to them.
The occurrence of Hen Harriers at the Indian roost shows, for a few individual birds, that the species’ distribution extends south of that quoted in Ali and Ripley (1978).
Further work to be done on hairier roosts in India should include an assessment of pesticide ingestion by harriers, especially in view of the importance of such a large number of harriers to the Asian breeding population. It is possible that they acquire organochlorines in parts of India which affect their success on breeding grounds. Montagu’s Harrier is under threat in many areas of the western part of its breeding range. This makes the gaining of an understanding the eastern component of the world population all the more urgent. The status of the more easterly-biased world Pallid Harrier population is not well known, although recent information suggests that the east European breeding population has largely vanished and there may have been some shifting of the range of the Asian breeding population due to major losses of habitat to grazing and agricultural use (Davygora and Belik 1994).
Acknowledgements
I thank Bill Clark, Graeme Hewson, Anil Mulchandani, Manoj Pai, Rishad Naoroji, Steve Rooke, the Bombay Natural History Society (Asad Akhtar, S. Nayak and other staff), M.K. Shivbhadrasinhji, the Jamsaheb of Jamnagar and the Gujarat Forest Department for their assistance. Ray Symonds at the University of Cambridge Museum of Zoology provided access to bird specimens. The observations were made during expeditions for the Hawk and Owl Trust, the first sponsored by British Airways, and as part of my studies as a postgraduate student at Liverpool University. Further work is being carried out on the Velavadar roost by the Bombay Natural History Society, sponsored by the Hawk and Owl Trust.
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JOURNAL, BOMBAY NATURAL HIST. SOCIETY, Vol. 93 (1996)
References
Ale S. & S.D. Ripley (1978): Handbook of the Birds of India and Pakistan. Vol. 1 . Second Edition. Oxford University Press, Delhi.
Barbier Montault, M. (1838): Notice sur les moeurs du Busard Montagu, Falco cinerascens Temm. Revue Zoologique 1 : 221-223.
Blanford, W.T. (1888): The Fauna of British India. Mammalia. Taylor & Francis, London.
Centre for Overseas Pest Research (1992): The Locust and Grasshopper Agricultural Manual. Hobbs, Southampton.
Cormier, J. P. & F. Baill.on (1991): Concentration de Busards Cendres Circus pygargus (L.) dans la region de M’bour (Senegal) durant l’hiver 1988-89. Alauda 59: 1 63- 168.
Davygora, A.V. & V.P. Belik (1994): The Pallid Harrier ( Circus macrourus) as an endangered species in the Palearctic. pp. 93-96, in Mcyburg, B.-U. & R.D. Chancellor (eds.) Raptor, Conservation Today. World Working Group on Birds of Prey and Owls.The Pica press.
Glutz von Blotzheim, U.N., K. M. Bauer, E. Bezzel (1971):
Handbuch der Vogel Mitteleuropas. Bd. 4. Falconiformes. Frankfurt a. M. Akademische Verlagsgesellschaft.
Khacher, L. (1977): Migrating harriers. J. Bombay nat. Hist. Soc. 74: 355-356.
Koppikar, B.R. & J.H. Sabnis (1976): Identification of hairs of some Indian Mammals. J. Bombay nat. Hist. Soc. 73: 5-20.
Newton, I. (1979): Population Ecology of Raptors. Poyser, Berkhamsted.
Nieboer, E. (1973): Geographical and ecological differentiation in the genus Circus. Ph.D. Thesis, University of Amsterdam.
Rahmani, A.R. & R. Manakadan ( 1 986): A large roost of harriers in Andhra Pradesh, India. J. Bombay nat. Hist. Soc. 83: 203- 204.
Satheesan, S.M. & P. Rao ( 1 990): Roosting and feeding of harriers in Secunderabad, Andhra Pradesh. J. Bombay nat. Hist. Soc. 87: 143.
Schipper, W.J.A. (1977): Hunting in three European harriers (Circus) during the breeding season. Ardea 65: 53-72.
Wassenich, V. (1968): Durchzug and Uberwinterung der Kornweihe. Regulus 9: 214-225.
STUDIES ON AMPHIPODS OF VISAKHAPATNAM COAST1
P. SOMANADHA RAO, K. HANUMANTHA RAO AND K. SHYAMASUNDARI 2
( With three plates )
Key words: nine genera, nine species, planktonic amphipods, Visakhapatnam
During a survey of amphipod fauna of Waltair coast, some gammaridean amphipods were encountered. Nine species belonging to five families and nine genera are described. The order Amphipoda comprises four sub-orders the Gammariidea, Hyperiidea, Caprelliidea and Ingolfelliidea. According to Barnard (1969) the constant presence of atleast 6 pairs of thoracic appendage, five pairs of gills and four pairs of brood lamellae in females are distinctive characters of Gammariidea and Hyperiidea.
Some of the recent contributions on the taxonomy of amphipods are those of Bellan-Santini and Dauvin (1981), Dickinson (1982), Goeke and Heard, Jr. (1983, 1984), Andres (1985), Goeke (1985, 1987), Thomas and Barnard (1986) and Locke and Corey (1989).
Samples were collected twice a week by towing a plankton net for a definite time. Each sample was used for numerical estimation and identification in living condition. All collections were made in Lawson’s Bay, 1 km away from the coast. Samples collected during June, 1984 and May, 1987 form the material for the present study.
Family Ampeliscidae Genus Ampelisca Krgyer
AmpeSisca zamboangae Stebbing (Plate 1, Figs. 1-10)
Body transparent. The distinguishing character is the fifth peraepod (Fig. 1). The expanded second joint reaches beyond the third joint. The flagellum and antenna of female have three segments. Coxal plate of first gnathopod (Fig. 4) broader towards the distal end fringed with plumose setae, second joint of second gnathopod elongate, devoid of setae, fifth and sixth joints subequal and setose, sixth narrow distally. Peraeopod description is similar to that given by Sivaprakasam (1966). The outer ramus of the third uropod is longer than the inner ramus. There are four spines on the outer surface and three setae on the inner surface of the outer ramus. There is a deep notch in the telson, each lobe bearing a spine and four setae at its distal end.
1 Accepted June 1993.
department of Zoology, Andhra University, Visakhapatnam-530 003, India.
Length: 4.5 mm.
Occurrence: Lawson’s Bay — 3 males; 3 females.
Distribution: Philippines, Sri Lanka, East Indies, Arabian Sea, Red Sea and Bay of Bengal.
Family Haustoridae Genus Uwthoe Dana Urothoe ruber Giles (Plate 1, Figs. 11-14)
Remarks: Fairly common in plankton.
Head slightly reduced. A common form in the plankton. First antenna 5-jointed with a 2-jointed accessory flagellum. Second antenna longer than the body.
First gnathopod subchelate. Fifth joint longer than sixth. There are long setae on the second joint. In the second gnathopod, fifth joint is narrower than that of the gnathopod one. Sixth joint apically produced to form chela with dactylus. Long setae on 2nd and 3rd joints. The first two peraeopods are alike (Fig. 12). Fourth joint long, sixth joint club- shaped and bears stout spines. Seventh joint cannot be distinguished from the spines of the sixth joint.
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JOURNAL BOMBAY NATURAL HIST. SOCIETY, Vol. 93 (1996)
The joints of the third peraeopod and plumose setae. Fourth and fifth peraeopods have a flat joint.
Uropod 1 and 2 are alike. In the third uropod (Fig. 1 4) the rami are flattened and bear long plumose setae along the margins. The eleft telson has five distal spines.
Length: Male 3.3 mm; Female 4.4 mm.
Distribution: Bay of Bengal.
Genus Platyischnopus Stebbing
Platyischnopus herdmanii Walker (Plate 1, Figs. 15-17)
A number of males and a few females were seen in plankton. Rostrum oblong. Head fairly long, longer than the first four segments put together. First antenna with a two jointed accessory flagellum. Antenna extends beyond the body. Second antenna shorter than the first.
First gnathopod has adistally expanded second joint (Fig. 15). Third and fourth joints subequal in length. Sixth joint forms a chela with the seventh. Second gnathopod similar to the first. The structure of the peraeopods agree with the description given by Walker ( 1 904). First two peraeopods similar with a distal ly expanded fourth joint. Fifth peraeopod long and narrow. Telson cleft, each lobe with a strong stouter tooth and 2 distal setae.
In the third uropod the inner ramus is small with a pointed apex. The outer ramus is 2-jointed with a spine-like second joint.
Occurrence: Lawson’s Bay — throughout the
year.
Distribution: Sri Lanka, South Africa, Arabian Sea and Bay of Bengal.
Family Oedicerotidae Genus Perioculodes O. Sars
Perioculodes megapleon (Giles)
(Plate 2, Figs. 1 8-26)
Body uniformly broad, with a short rostrum. Pleon segments large. First antenna short, stout, with the peduncle bearing setae. Flagellum 9-jointed and fringed with fine hairs. The joints of the peduncle of
second antenna decrease in width. In male, the flagellum is longer than the body.
The two gnathopods are similar, subchelate. First gnathopod with second joint very long and slender. Palm slightly oblique, with fine teeth. Second gnathopod similar to first, but slightly longer. Palm more oblique than the first. Seventh joint forms a chela with the fifth.
First and second peraeopods similar. Fourth, fifth and sixth segments properly setose. Sixth segment with 3 spines, seventh indistinguishable from setae. In the third peraeopod, second segment is broader. Third is small, rest are slender; long hairy setae on every segment. Dactylus spiniform, with a fringe of hairy setae. Fourth peraeopod is slightly longer than the third. Fifth leg (Fig. 23) is longer than the preceding legs.
Uropods (Figs. 24-26) are similar in structure. Peduncle longer than the rami, with terminal spines. Rami fringed with setae on both margins. The length decreases successively from first to third uropod. Telson bilobed, with four setae.
Family Gammaridae Genus Hornellici Walker
Hornellia incerta Walker (Plate 2, Figs. 27-29)
Body has serrated pleon segments. Peduncle of first antenna fringed with short hairs. A long 20- jointed flagellum and 2-jointed accessory flagellum. Flagellum longer in males than in females. In the first maxilla, the second joint of the palp has small spines alternating with larger spines unlike that observed by Walker (1905).
Both gnathopods similar with triangular fifth joint. Palm border possesses spines (Fig. 27). Except the first two, all other peraeopods are long, with all segments spiny. Firt uropod (Fig. 28) is the longest and reaches slightly beyond the third uropod. Outer ramus of the first and second uropods shorter than the inner. Rami of third uropod (Fig. 29) unequal, inner border with a few plumose setae.
Length: 4.0 mm.
Occurrence: Lawson’s Bay.
J. Bombay nat. Hist. Soc. 93 Somanadha Rao etal.: Amphipods
Plate 1
Figs. 1-10. Ampelisca zamboangae Stebbing: 1. Mandible; 2. Maxilla; 3. Maxilliped; 4. Gnathopod 1;
5. Peraeopod 1; 6. Peraeopod 4; 7. Peraeopod 5; 8. Uropod 1; 9. Uropod 2; 10. Telson.
Figs. 1 1-14. Urotlioe ruber Giles: 11. Antenna; 12. Peraeopod 2; 13. Peraeopod 3; 14. Uropod 3 with telson. Figs. 15-17. Platyischnopus herclmanii Walker: 15. Gnathopod 1; 16. Peraeopod 5; 17. Telson.
J. Bombay nat. Hist. Soc. 93 Plate 2
Somanadha Rao et al.\ Amphipods
Figs. 18-26. Pcrioculodcs megapleon (Giles): 18. Antennule; 19. Antenna; 20. Gnathopod 1; 21. Gnathopod 2; 22. Peraeopod 1; 23. Peraeopod 5; 24. Uropod 1; 25. Uropod 2; 26. Uropod 3.
Figs. 27-29. Horndlia incerta Walker: 27. Gnathopod 2; 28. Uropod 1; 29. Uropod 3.
Figs. 30-31. Megaluropus agilis Hoek: 30. Uropod 3; 31. Telson.
Figs. 32-35. Pseuclotiron brevidcictylus Pillai: 32. Gnathopod 2; 33. Peraeopod 2; 34. Peraeopod 5; 35. Uropod 3.
Fig. 35a. Parcdasinoplms suluensis Stebbing: Gnathopod 2.
J. Bombay nat. Hist. Soc. 93 Somanadha Rao et ai: Ainphipods
Plate 3
Figs. 36-39. Pcirelasnwpiis suluensis Stebbing: 36. Uropod 1; 37. Uropod 2; 38. Uropod 3;
39. Telson. Figs. 40-48. FJasmopus pectenicms (Bate): 40. Maxilla 1; 41. Maxilla 2; 42. Maxilliped; 43. Gnathopod 1; 44. Gnathopod 2; 45. Peraeopod 1: 46. Peraeopod 5; 47. Uropod 1; 48. Uropod 3.
STUDIES ON AMPHIPODS OF VISAKHAPATNAM COAST
53
Distribution: Sri Lanka, Arabian Sea and Bay of Bengal.
Genus Megcilumpus Hoek
Megaluropus agilis Hoek (Plate 2, Figs. 30-31)
First antenna as long as the peduncle of second antenna with a 10-jointed flagellum and 2-jointed accessory flagellum. Second antenna 13-jointed. Gnathopods as described by Stebbing (1906).
The second segment in the third, fourth and fifth peraeopods is expanded. Fifth peraeopod longer than the body. Seventh segment spine-like with plumose setae. All appendages setose, rami of the third uropod (Fig. 30) highly expanded, bearing scanty plumose setae. Telson cleft with rounded distal lobes (Fig. 31 ).
Length: 4.2 mm.
Occurrence: Lawson’s Bay.
Distribution: Krusadai Island, Tamil Nadu, Port Blair, Andamans, Sri Lanka, South Africa, Mediterranean, North Sea.
Family Tironidae Genus Pseudotiron Chevreux
Pseudotiron brevidactylus Pillai
(Plate 2, Figs. 32-35)
This species was first erected by Pillai (1957). It has a compressed body with a square head. First antenna with short accessory flagellum. Mandible without a palp; accessory plate with seven barbed spines. Outer lobe of first antenna has 9 spines. Inner lobe small and bears 5 teeth and 1 seta. Inner lobe of the second maxilla broader and slightly setose. The inner lobe of the maxilliped truncate with seven stiff setae.
Two gnathopods alike, with hook-like dactylus (Fig. 32). Third, fourth and fifth peraeopods with long spines (Figs. 33 and 34).
First and second uropods with spines. Tips of all rami with three spines each. The rami of third uropod slightly flattened, with long apical and short outer spines and plumose setae on the inner border
(Fig. 35). Telson long, lanceolate, each lobe with a long apical spine.
Length: 3.5 mm.
Occurrence: Lawson’s Bay.
Genus Parelasmopus Stebbing Parelasmopus suluensis Stebbing
(Plate 2, Fig. 35a, Plate 3, Figs. 36-39)
Agrees in all essential details with the description given by Stebbing (1906). Pleon segments less massive. Fourth pleon segment bears large teeth, the lower border of third pleon segment has a single tooth. Male has a 3-jointcd accessory flagellum. Gnathopods weak, fifth segment of second gnathopod longer than broad. The palmar border of sixth segment serrated (Fig. 35a). The apical spines of the first and second uropods (Figs. 36-38) are longer than in Stebbing’s illustration. The apical lobe of the telson (Fig. 39) bears 3 spines.
Occurrence: Lawson’s Bay.
Distribution: Maidive and Laccadive Islands and Bay of Bengal, Sri Lanka, Red Sea, East Africa. Australia.
Genus Elasmopus Costa
Elasmopus pectenicrus (Bate)
(Plate 3, Figs. 40-48)
Antenna 1 is half the length of the ani- mal. Peduncle and flagellum of equal length. Peduncular joint subequal. Accessory flagellum 2-joined. Antenna 2 as long as the peduncle of antenna 1 .
Inner lip with very small inner lobes. Fine setae present on the anterior margin of both the lobes. First maxilla (Fig. 40) with an outer and inner lobes and a palp. Maxilla 2 (Fig. 41) with inner plate heavily setosed. Outer lobe bears a number of apical setae (8-10). Maxilliped (Fig. 42) with a narrower inner and a broader outer lobe bearing long and slender setae around them. Second joint of palm reaches well beyond the outer lobe. The last joint of palp bears apical ly long and hairy setae.
Gnathopod 1 (Fig. 43) smaller, second joint long, fifth and sixth segments subequal in length,
54
JOURNAL BOMBAY NATURAL HIST. SOCIETY, Vol. 93(1996)
sixth with parallel sides. Finger slightly curved to fit the palm. Palm slightly oblique with a row of weak spines. The whole appendage is heavily setose on the margins.
Gnathopod 2 (Fig. 44) bigger than the first, second joint elongated, fifth joint slightly cup- shaped. Bands of setae present on the sixth article. Palm oblique, sixth segment with a row of curved
Refer
Andres, H.G. (1985): The Gammaridea (Crustacea: Ainphipoda) from the German Antarctic Expedition 1975/76 and 1977/ 78: Acanthonotozomatidae, Parampthithoidae and Stegocephalidae. Mitt. Hatnb. Zool. Mus. Inst. 8(20): 119- 154.
Barnard, J.L. (1969): The families and genera of marine gammaridean Amphipoda. Pmc. U.S. Nat. Mus. 271: 1-535. Bellan-Santini, D. & J.C. Dauvin (1981): Description of anew species of Ampelisca of the French coast (Amphipoda). Crustaceana 40(3): 242-252.
Dickinson, J.M. (1982): The systematies and distributional ecology of the family Ampeliscidae (Amphipoda: Gammaridea) in the north eastern Pacific region. Natl. Mus. Nat. Sci. ( Ottawa ) Publ. Biol. Oceanogr. 0(10): 1-40.
Goeke, G.D. (1985): Amphipoda of the family Ampeliscidae (Gainmariidea). V. H. hawaiiensis new species. Pac. Sci. 39(3): 261-265.
Goeke, G.D. (1987): Amphipods of the family Ampeliscidae (Gainmariidea) VI. Ampelisca macroclonta new species from