3SHJLIWS S31dVd8!l_LIBRARiES SMITHSONIAN _ INSTITUTION NOimillSNI NVINOSH1IINS S3 1
in
Q O
HSONIAN-1 INSTITUTION NOlinjLIlSNI^NVINOSHiltAIS S3 I d Vd 8 II B R AR I E$Z SMITHSONIAN*” IN SI r- 2: r- * z r* z “
m x^vosv^ ^ m
OSHIUNS^SS I HVaairilBRAR I ES SMITHSONlAN~INSTlTUTION%OlinillSNI ~NVIN0SH1IINS S3 I
CO 2 * S2 ___ Z ♦ CO Z CO
I 1 Jfe. 1 ^ s
lull
<l CO JPf- S£ O
|A | | 1 y |.. 5 >
IONIAN JNSTITUTI0NOTN0liniliSNI_NVIN0SHilWSOTS3 I a Va a Ilf LI B RAR I ES^SMITHSONIANJNSl
M's w ^ . S g /5C22£5x “ -A- 5
)shiiims S3 iava an libraries Smithsonian institution Nou.ruu.SNi nvinoshiiws S3i
Z r- z r- z r- z
&<v ° m 2 2 /«\ ~ jSSf* O
m xg l?v>^ " rn ^ x^oius^x m
HSON IAN INSTITUTION^ NOIinillSM~NVINOSHllWS S3 I dVd8 IUI B RAR I ES ^SMITHSON IAN INS
Z ,yy. 00 Z
:VASV/H>. <
^ | 0^^ 1 I | 8
OSHilWS S3 I ava a nZ LI B RAR I ES^SMITHSONIAN^NSTITUTION^NOlinillSNI^NVINOSHilWS^Sa c n = co 5 __ 4/5 ^ . oo
OJ m W /»T\ ,&CX> IAJ
a
o X^M P.C^ — O X?. ov/ „ ^11^ Q
HSONIAN^NSTJTUTION^NOlinillSM^NVINOSHlIINS^SS I d Vd 8 11 "^Ll B R AR I ES^ SMITHSON IAN”* INS “ z r- * z r* Z r
2 m ^ 2 m °
c/5
»AK»wjy w m sg m
0SH1IWS S3 I ava a n~LI B RAR I ES SMITHSONIAN_INSTITUTIONWNOIinillSNI~NVINOSHilWS S3 I __ £ z 1 W , ^ z , w z “
-* z m. A h z mv4 h 2
o 4ft?1 ¥ \ S2 (|fe 8 ^ 8
§ x Xw § g ^ w i
CO
S .W > 2 ‘-'X^ >'y 2 >
S A z % 2 ^ 2
HSONIAN INSTITUTION NOlinillSNI^NVINOSHlIWS S3 I d Vd 8 I1_LI B R AR I ES SMITHSONIAN INS'
m%. 5 l /0%\ I 1 1 ? 4r
RARIES SMITHSONIAN INSTITUTION N011IUI1SNI NVINOSH1IWS SSIdVdSH LIBRARIES SMITHSONIAN INSTITUTION NQIilUtiSNI
Z (J) ^ VS c n Z
LIBRARIES SMITHSONIAN INSTITUTION NOIlfUliSNI NVINOSH1IWS S3!BVdail LIBRARIES <n ^ ~ co
" '<guu^ 5
N0UfUUSNI~NVIN0SHiIlAlS_S3 IdVHOII^LIBRARI ES^SMITHSONIAN^INSTITUTION NOIlfUliSNI
2 2 _ ......... O M&h, . _ O
CO
33 >
m '■ \^v 2 XjvAs^x m x^vosv^x ^ m
LI B R AR I ESC/,SMjTHSONIAN~INSTITUTIONC/>NOIinillSNl"‘NVlNOSHlllMS S3IBVaail LIBRARIES 03 z: 00 2 CO 2 \
< A S <
X & § I
vm Q til 2 ^
o
> 5 >*' 2
NOIlfUliSNI NVINOSHlMS^SB I d VB 8 ll\ I B R AR I ES^SMITHSONIAN^INSTITUTION ° NOllfUllSN
to ~ tn = co
LIBRARIES SMITHSONIAN INSTITUTION NOIlfUliSNI NVIN0SH1MS S3ldVdail LI B R AR I El
2 r* __ 2 r9 z r*
m ^ xguiissx m
NOIlfUliSNI NVIN0SH1IWS S3 I d Vd 8 n~ LI B R AR I ES^SMITHSONIAN- INSTITUTION NOlinillSh
— co Z CO 2 Co
2 o
CO
t
2 5 ^ > N^Jjiis^x ^ ^ >
LIBRARIES SMITHSONIAN INSTITUTION NOIlfUliSNI NVINOSHlIWS^Sa I d Va a II LI B RAR I E
to ~z co — CO
O __ o _ X<piiis£X O
NOllfUllSNI^NVINOSHlIlNS^Sa I BVB 8 ll^LI B RAR I ES^SMITHSONIAN^INSTSTUTION NOllfUllSN r; 2 r* z . [- 2
<»ASTiT(7>v _ ASk. O /W?V O __ /<O0:Vo\ O XtlVnT//>s
_ m
LIBRAR I ES SMITHSONIAN ^INSTITUTION^ NOllfUllSN i"-NVINOSHilWS</) S3 I HVH a 11 LIBRAR I E‘
2 » CO 2 CO 2 ’•
X o
> > '
NOIlfUliSNI _NVIN0SH1IWSW S3 IdVdS I T^L t B RAR I ES^SMITHSONIAN^INSTITUTION “ NOllfUllSN
co = m — _ co
CO
Vol. 89, No. April 1992
a
BOARD OF EDITORS
Executive Editor J. C. DANIEL
M. Rt ALMEIDA P. V. BOLE B. F. CHHAPGAR
Assistant Editor A. VARADACHARY
m
B. V. DAVID A. J. T. JOHNSINGH R. WHITAKER
INSTRUCTIONS TO CONTRIBUTORS
Papers which have been published or have been offered for publication elsewhere should not be submitted.
All words to be printed in italics should be underlined.
Trinomials referring to subspecies should only be used where identification has been authentically established by comparison of specimens actually col- lected.
Photographs for reproduction must be clear, with good contrast. Prints should be at least 8.20 x 5.60 cm (No. 2 Brownie) and on glossy glazed paper.
Text-figures, line drawings and maps should be in Indian ink, preferably on Bristol board.
References to literature should be placed at the end of the paper, alphabeti- cally arranged under author’s name, with the abridged titles of journals or periodicals underlined (italics) and titles of books not underlined (roman type), thus:
Banerji, M. L. (1958): Botanical Exploration in East Nepal /. Bombay nat , Hist. Soc. 55(2): 243-268.
Prater, S. H. (1948): The Book of Indian Animals, Bombay.
Titles of papers should not be underlined.
Each paper should be accompanied by a concise, clearly written synopsis, normally not exceeding 200 words.
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.
The editors reserve the right, other things being equal, to publish a member’s contribution earlier than a non-member’s.
Hornbill House,
Shaheed Bhagat Singh Road, Bombay 400 023.
Editors,
Journal of the Bombay Natural History Society
.
smTR
Date of Publication: 30-6-1992
VOLUME 89 (1): APRIL
CONTENTS
FLIGHT IDENTIFICATION OF INDIAN RAPTORS WITH PALE BARS ON UPPER WINGS (With two plates)
By William S. Clark and N. John Schmitt 1
SYSTEMATIC POSITION OF MOLOSSIDAE - AN EMBRYOLOGICAL ANALYSIS (With two text-figures)
By A. Gopalakrishna and N. Badwaik . 4
A STUDY OF THE FOOD HABITS OF SIX ANURAN TADPOLES (With two text-figures)
By A.G. Sekar 9
I. CLADOCERA OF KEOLADEO NATIONAL PARK, BHARAITUR, AND ITS ENVIRONS (With forty-nine text-figures)
By K. Venkataraman 17
FOOD AND FEEDING BEHAVIOUR OF THE GREAT INDIAN BUSTARD Ardeotis nigriceps (VIGORS)
(With two plates and four text-figures)
By Bharat Bhushan and Asad R. Rahmani 27
ON THE IDENTITY AND NOMENCLATURE OF CERTAIN INDIAN Ixora (RUBIACEAE)
By D.B. Deb and R.C. Rout 41
THE LAND TORTOISE IN NEPAL : A REVIEW (With a plate and a text-figure)
By J. Frazier 45
A CATALOGUE OF THE BIRDS IN THE COLLECTION OF BOMBAY NATURAL HISTORY SOCIETY - 35: TROGLODYTIDAE, CINCUDAE, PRUNELUDAE, PARIDAE, SITTIDAE AND CERTHIIDAE By Humayun Abdulali and Saraswathy Unnithan 55
BIOECOLOGICAL STUDIES ON THE BURROWING MAYFLY Ephemera (Aethephemera) nadinae
MCCAFFERTY AND EDMUNDS 1973 (EPHEMEROPTERA : EPHEMERIDAE) IN KURANGANI STREAM, WESTERN GHATS (With three text-figures)
By C. Balasubramanian, K. Venkataraman and K.G. Sivaramakrishnan 72
RELATIONSHIP BETWEEN CANOPY DENSITY AND BREEDING BEHAVIOUR OF Ploceus philippinus (LINN.) AND Ploceus benghalensis (LINN.) (With two text- figures)
By Satish Kumar Sharma 78
REPRODUCTIVE BIOLOGY OF THE HANUMAN LANGUR Presbytis entellus IN JODHPUR, WESTERN INDIA (With four text- figures)
By G. Agoramoorthy 84
NEW DESCRIPTIONS
FIRST RECORD OF THE GENUS Laurentina MALAISE (HYMENOPTERA : TENTH REDINIDAE) FROM INDIA, WITH DESCRIPTION OF A NEW SPECIES (With two text-figures)
By Malkiat S. Saini and Devinder Singh 94
Osteobrama bhimensis , A NEW CYPRINID FISH FROM BFIIMA RIVER, PUNE DISTRICT, MAHARASHTRA (With two text-figures)
By D.F. Singh and GM. Yazdani 96
FIRST REPORT OF THE FAMILY VAEJOVIDAE (SCORPIONIDAE : ARACHNIDA) IN MADHYA
PRADESH, WITH THE DESCRIPTION OF A NEW SPECIES Scorpiops (Scorpiops) pachmarhicus (With eight text-figures)
By Deshabhushan Bastawade 99
ON A NEW SPECIES OF Singhius TAKAHASHI (ALEYRODIDAE : HOMOPTERA) WITH A KEY TO INDIAN SPECIES ( With a text- figure)
By R. Sundararaj and B.V. David 103
A NEW SPECIES OF Liparis RICHARD (ORCHIDACEAE) FROM SIKKIM
By S.Z. Lucksom 105
Copidognathus krantzi, A NEW SPECIES OF HALACARIDAE (ACARI) FROM NICOBAR ISLANDS (INDIAN OCEAN) ( With ten text-figures)
By Tapas Chatterjee 106
REVIEWS
Flora of the Indian Desert
Reviewed by M.R. Almeida 110
A Revised Handbook to the Flora of Ceylon, Vol. 6
Reviewed by M.R. Almeida 112
MISCELLANEOUS NOTES
MAMMALS
1. Interspecific play behaviour between hanuman langur Presbytis entellus and rhesus macaque Macaca mulatta
By B. Ram Manohar and Reena Mathur .... 114
2. Notes on the food habits of nilgai Boselaphus tragocamelus
By K. Sankar and V.S. Vijayan 115
BIRDS
3. Marbled teal Marmaronetta angustirostris
(Menetries) in western India
By S.A. Akhtar, J.K. Tiwari and
N.N. Bapat 116
4. Balloons as a device for scaring birds
By H.S.A. Yahya 117
5. Unusual nesting site of brahminy kite Haliastur indus
By William Morrison, Lima Rosalind
and S. Balachandran 117
6. Great stone plover Esacus magnirostris (Vieillot) in Kerala By P.O. Nameer
7. Possible occurrence of four subspecies of lesser sand plover Charadrius mongolus at Pt. Calimere Wildlife Sanctuary, Tamil Nadu
By S. Balachandran and V. Natarajan 118
8. Occurrence of Larus minutus Pallas in Kutch
By N.N. Bapat and M.K. Himmatsinhji .... 119
A
9. On the black tern Chlidonias niger niger (Linn.)
By Vivek Menon 120
10. Sterna bergii thalassina Stresemann — an addition to the avifauna of Sri Lanka
By Thilo W. Hoffmann 120
11. Review of the status of the sandwich tern Sterna sandvicensis in Kerala
By D.K. Narayana Kurup 122
12. New nesting site of the Indian whitebreasted kingfisher Halcyon smyr- nensis fusca (Boddaert).
By P. Balasubramanian 124
118
13. Feeding by common nightjar Caprimulgus asiaticus and Indian roller Coracias ben- ghalensis in the light of mercury vapour lamps
By A.M.K. Bharos
14. Possible occurrence of the grey shrike Lanius excubitor Linn, in Assam
By Anwaruddin Choudhury
15. Breeding biology of the Malabar woodshrike Tephrodornis virgatus syl- vicola Jerdon at Thekkady, Kerala
By Lalitha Vijayan
16. Wintering range extension for the rubythroat Eritliacus calliope
By S. Balachandran, Lima Rosalind and S. Alagar Rajan
17. Plumages, female dimorphism and polymorphism of the endemic Indian species Par us xantliogenys
By S. Unnithan
18. Interesting feeding pattern of yel- lowthroated sparrow Petronia xanthocollis (Burton)
By A.M.K. Bharos
19. ‘Blind" nest of blackthroated weaver bird Ploceus benghalensis (Linn.)
By Satish Kumar Sharma
20. Spotted munia Lonchura punctulata (Linn.) from DachigamNational Park, Jammu and Kashmir
By S.A. Akhtar, Prakash Rao,
J.K. Tiwari and Salim Javed
21. An updated list of bird and bat species involved in collision with aircraft in India By S.M. Satheesan, Robert B. Grubh
and Rex J. Pimento
REPTILES
22. Gut contents of a mugger Crocodylus palustris
By R J. Rao and S.A. Hussain v
23 . Unusual nesting site of mugger Crocodylus palustris in Madhav National Park
By Rajiv Saxena 132
24. Swallowing of prey ‘leg first" by the cobra Naja naja
By Satish Kumar Sharma 133
AMPHIBIA
25. First record of Uperodon systoma from Rajasthan
By Satish Kumar Sharma 133
26. Additions to the list of amphibian fauna of Goa
By A.G. Sekar 134
FISHES
27. New record of a croaker, Johnius coitor (Hamilton-Buchanan) (Pisces : Sciaenidae) from Tripura, north-east India
By R.P. Barman 135
INSECTS
28. Dichocrosis festivalis Swinh. (Lepidoptera Pyralidae) — A new pest of litchi Litchi chinensis Sonn.
By Y.P. Singh and V. Kumar 137
OTHER INVERTEBRATES
29. Record of the arachnid order Schizomida from Arunachal Pradesh
By Deshabhusan Bastawade and
Tarun Kumar Pal 137
30. Redescription of Araneus fulvus Dyal (Araneae: Araneidae) from coastal Andhra Pradesh
By T.S. Reddy and B.H. Patel 138
31. Cladocera of Keoladeo National Park,
Bharatpur, IV. New records: Camptocercus cf. australis Sars, 1896 and Indialona globulosa (Daday, 1898)
By K. Venkataraman 140
32. Record of the cone shell Conus cumingii (Reeve, 1848) from Bombay seas ByDeepakApte 142
BOTANY
33. Corrections and additions to the flora of Gurdaspur district, Punjab By S.S. Bir, M. Sharma and C.P. Singh .... 143
124
124
125
126
126
128
128
129
129
132
34. Additions to the flora of Bihar
By S.K. Varma and R.R. Jha 146
35. Additional host species for Loranthus and their localities in Thanjavur district, Tamil Nadu
By S. Ragupathy and A. Mahadevan 149
36. Vetiveria lawsoni (Hook, f.) Blatter & Mc- Cann and Potamogeton crispus L. — Ad- ditions to the flora of Andhra Pradesh
By B. Ravi Prasad Rao and
T. Pullaiah ..f.. J.50
37. Indian doum palm in Khandesh — an un- usual occurrence
By A.S. Reddy and T.S. Patil 151
38. Another locality record for Cyathea spinulosa in Kumaon Himalaya
By H.C. Pande and P.C. Pande 152
JOURNAL
OF THE
BOMBAY NATURAL HISTORY
SOCIETY
April 1992 Vol. 89 No. 1
FLIGHT IDENTIFICATION OF INDIAN RAPTORS WITH PALE BARS ON UPPER WINGS1
William S. Clark2 and N. John Schmitt3 (With two colour plates)
INTRODUCTION
Diurnal raptors are notoriously difficult to identify in flight; raptors in India are even more so than in most other areas because of the greater number of species (68) and the lack of definitive information in bird field guides. In Europe raptor flight identification is easier not only because there are fewer species (38), but also because there is a very good specialized field guide (Porter et at. 1981). This guide is effective because it depicts correctly wing and tail shapes of the raptors, as well as pointing out definitive field marks , that is, noticeable fea- tures of each species that serve to distinguish it from other species.
No fewer than nine species of raptors that occur commonly over much of India share one field mark: a pale bar across each upperwing (Plate 1). Four of these — - black kite Milvus migrans , booted eagle Hieraaetus pennatus , short-toed eagle Circaetus gallicus, and white- eyed buzzard Butastur teesa — show this field mark in all plumages. The other five — brahminy kite Haliastur Indus , crested honey buzzard Per -
Accepted October 1991.
24554 Shetland Green Road, Alexandria, VA 22312, U.S.A.
311609 Alburtis Ave., Norwal\ CA 90650, U.S.A.
nis ptilorhyncus, crested hawk-eagle Spizaetus cirrhatus , crested serpent eagle Spilornis cheela , and Bonelli’s eagl e Hieraaetus fasciatus — show this mark only in juvenile plumage.
In spite of sharing this field mark, all nine are quite different, particularly when seen from below, and can be easily distinguished from each other by the use of other field marks, especially wing and tail shape.
We present herein, through simple text and illustrations, the field marks that can be used effectively to identify all nine species in flight.
Material and Methods
Field marks to distinguish these nine species were determined from our previous ex- periences, by reviewing the pertinent literature, including bird field guides and handbooks, by studying museum specimens, both in India and at major collections in the United Kingdom and United States of America, by studying many photographs of raptors in the field, and by ob- serving raptors in the field in many parts of India. Particularly helpful to us was Porter et al. (1981), as many species of European raptors (31) also occur in India. We field-tested the field marks presented here in the field in many parts of India during travel with the BNHS Birds of Prey Project surveys.
2
JOURNAL, BOMBAY NATURAL HIST SOCIETY, Vol. 89
Results
The field marks, including wing and tail shapes and overall proportions, that will positively identify each of the nine species with pale bars on upperw- ings are illustrated in Plate 1 (from above) and Plate 2 (from below) and are summarised under the head- ing ‘Captions FOR plates/ These are discussed below in more detail by species.
Blackkite: (This name for Milvus migrans applies to all races; pariah kite is unknown outside of India). This distinctive raptor is com- mon, widespread, and easily identified. The next two species are similar and could be con- fused with it, but its long forked tail is always definitive. However, caution is called for be- cause the tail appears somewhat more square when fanned and a few individuals will show a somewhat rounded tail. Also definitive is the barring on the pale primary panels. Subspecific differences between M. m. govinda and M. m. lineatus are not always seen in the field.
Booted eagle: This winter visitor is fairly common over most of India and can be confused with the black kite. Like that species it is aerial, hunting on the wing from morning to afternoon. It occurs in three colour morphs: pale, dark, and newly described rufous (see Clark 1989). Dark- and rufous-morph birds are similar to black kites, but the white uppertail coverts, more rounded tail, pale ‘head lights/ and dark line through the un- derwings of the rufous morph are diagnostic. All colour morphs appear alike from above.
Brahminy kite: Juvenile brahminy kites are similar in silhouette to black kites and booted eagles, but note the pale head and breast, rounded, unbanded tail, and larger, creamy primary panels on the underwings.
The next four species show pale wing bars only during their first year while in juvenile plumage. This plumage is quite different in all four from the respective adult plumages. All four juveniles are similar to each other in being rather pale buffy to creamy on the underparts and under- wing coverts, and have less distinct pale bars on the upperwings than do the previous three species.
Crested honey buzzard: This species has a distinctively long, slender neck and head that it moves constantly from side to side while in flight. Its comparatively narrow wings with darker secondaries on the underwing and its distinctive tail pattern are sufficient for identification.
Crested hawk-eagle: Compared to the other species considered here, it has a longer, distinc- tively banded tail and more strongly barred under- sides of primaries. The crest, if present, is visible only on birds seen flying near (Plate 1). The juvenile of the crestless changeable hawk-eagle Spizaetus c. limnaeetus is otherwise identical to the juvenile of the crested hawk-eagle.
Crested serpent eagle: This is the most easi- ly identified of these four juveniles, because of its black face patches, rufous underwing markings, strong tail pattern, and heavily streaked breast.
Bonelli’s eagle: Juveniles of this species have rufous underparts when they fledge. But this colour fades rather quickly, so that by winter they appear quite creamy on the underparts. The black line on the underwing may be prominent or indis- tinct or, in some cases, even absent. One field mark, darker secondaries on underwing, is shared with crested honey buzzard, but the indistinct banding on secondaries and tail and thicker head and shorter neck of this species separate it from the other.
Short-toed eagle: This species is the largest of the nine. Its pale upperwing bars are somewhat wider than all the others, except those of the white-eyed buzzard. The dark hood, lack of creamy tones on undersides, and strong banding on undeiwings are distinctive.
White-eyed buzzard: This is the smallest of the nine and is quite different from the rest. Note particularly the narrow black tips of the outer primaries, rufous tail, and unique wing shape.
Acknowledgements
The illustrations herein were prepared by Schmitt to help teach raptor identification to per- sonnel of the BNHS raptor ecology project, a collaborative project funded by the U.S. Fish & Wildlife Service. We thank D. Ferguson of the
J. Bombay nat. Hist. Soc. 89 Plate 1
Clark and Schmitt: Identification of raptors
Raptors with pale bars on upper wings — Identification from above. For explanation see captions (page 3).
J. Bombay nat. Hist. Soc. 89
Clark and Schmitt: Identification of raptors
Plate 2
Raptors with pale bars on upper wings - Identification from below. For explanation see captions (page 3).
FLIGHT IDENTIFICATION OF INDIAN RAPTORS
3
Office of International Affairs, U.S. Fish & Wildlife Service, for arranging our visits to India. We thank the curators and collection managers of the BNHS, India Wildlife Survey, British Museuifi (Natural History), and U.S. National Museum of Natural History for access to their specimen collections.
Special thanks go to our Indian field com- panions, Rishad Naoroji and Vibhu Prakash, for showing us Indian raptors. J.C. Daniel and R. Grubh of the BNHS are thanked for support while in India. C. Wilds made many helpful comments on earlier drafts.
References
Clark, W. (1989): Rufous morph of the Booted Eagle. Dutch Porter, RE, Christensen, S., Willis, I. & Neilsen, B.P. Birding 11: 57-60. (1981): Flight identification of European raptors. 3rd
Ed. Poyser, Carlton.
CAPTIONS FOR PLATES
Plate 1. Raptors with pale bars on upperwings — Iden- tification from above.
All nine species have a more or less distinct pale bar across each upperwing. Many also have pale heads. Wing shape and tail shape and pattern are usually the best field marks for identification, but others are helpful. Diagnostic field marks are listed below.
Raptors 1, 2 and 3 have similarly shaped wings.
1. Black kite Milvus migrans (adult shown). Long forked tail. 2. Booted eagle Hieraaetus pennatus. White ‘headlights’ at base of forewings; white ‘U’ on uppertail coverts; rounded, square-cornered tail. 3. Brahminy kite Haliastur indus (pale juvenile). Rufous on upperparts, especially primaries; short, unbarred, rounded tail.
Raptors 4, 5 and 6 have pale heads and similarly shaped wings. 4. Crested honey buzzard Pernis ptilorhyncus (juvenile). Long, slender neck and head; distinct tail pat- tern. 5. Crested hawk-eagle (includes juvenile changeable hawk-eagle) Spizaetus cirrhatus (juvenile). Rump, as well as uppertail coverts, are pale; distinct tail pattern. (Crest not visible). 6. Crested serpent eagle Spilomis cheela (juvenile). Dark cheek patch; bold black and white tail pattern.
7. Bonelli’s eagl e Hieraaetus fasciatus (juvenile). Tail bands indistinct. 8. Short-toed eagle Circaetus gallicus. Larger size; larger head; pale bars wider; distinct tail pattern. 9. White-eyed buzzard Butastur teesa (juvenile shown). Smaller size; white primary panels; rufous tail.
Plate 2. Raptors with pale bars on upperwings — Iden- tification from below.
All nine species appear quite different from below; under- wings; underbody, and undertail patterns and colouratio are the best field marks. Diagnostic field marks are listed below:
1. Black kit eMilvus migrans (juvenile shown). Dark body and underwing; coverts; pale, banded primary panels; tail squarish when spread. 2. Booted eagle Hieraaetus pen- natus (rufous morph shown). Rufous body; wide black bands across underwings; pale patch on inner primaries; dark central patch on pale tail. 3. Brahminy kit eHaliast- indus (pale juvenile). Creamy unbanded primary panel, lower body darker than upper body; rounded, unbanded tail.
4. Crested honey buzzard Pernis ptilorhyncus (juvenile). Banded dark secondaries; finely streaked underparts; dis- tinct tail pattern. 5. Crested hawk-eagle Spizaetus cir- rhatus (juvenile). Strongly banded primaries; distinct tail pattern. 6. Crested serpent eagle Spilomis cheela (juvenile). Black face patch; heavily streaked breast; rufous underwing coverts; distinct tail pattern.
7. Bonelli’s eagle Hieraaetus .fasciatus (juvenile). Dark wing tips; indistinctly banded dark secondaries; narrow black band across underwing (usually); tail banding indis- tinct 8. Short-toed eagle Circaetus gallicus (adult shown). Larger size; large head; dark hood (usually); white under- wings boldly banded black; distinct tail pattern. 9. White- eyed buzzard Butastur teesa (juvenile shown). Smaller size; rufous underwing coverts and lightly banded flight feathers; wide, black throat stripe; rufous tail.
SYSTEMATIC POSITION OF MOLOSSIDAE - AN EMBRYOLOGICAL ANALYSIS1
A. Gopalakrishna and N. Badwaik2 (With two text-figures)
At present, morphological and anatomical characters constitute the main criteria for clas- sification of eutherian mammals, since other criteria are not available for most mammalian groups. But these systems of classification based on morphological characters, do not necessarily reflect the phylogenetic affinities of various sub- groups among mammals. This has been convinc- ingly argued by Mossman (1937, 1953) in his analysis of foetal membrane characters of various grades of eutherian groups. In the absence of adequate data from palaeontology, cytology, genetics, serology and such other disciplines, evidence from embryology assumes considerable significance for determining taxonomic position an ! phylogenetic affinities among lower grades of taxa, such as Super- families and Families.
So far all taxonomists have placed Pteropodidae at the begimiing and Molossidae along with Vespertilionidae within the Super- family Vespertilionidae, at the other end in the +axonomic hierarchy of the Order Chiroptera Jimpson 1945, Ellerman and Morrison-Scott 1951, Honacki et. al. 1982, Koopman 1984, Hill and Smith 1985). Jones (1917) examined the anatomy of the female genitalia of many species of bats and suggested that Chiroptera is a polyphyletic group, in which are included mem- bers derived from divergent ancestors.
Mossman (1937), basing his conclusions on foetal membrane characters, suggested that Megachiroptera share characters with Rodentia, whereas Microchiroptera are closer to Insec- tivora. It must, however, be conceded that very little information was available about the embryology of most families of Microchiroptera at that time. Moghe (1951), in his study of the embryology of Pteropus giganteus giganteus ,
1Accepted November 1989.
department of Zoology, Institute of Science,
Nagpur 440 001.
mentioned, "the two groups (Megachiroptera and Microchiroptera) are widely separated from each other in a large number of other characters and probably represent independent offshoots from some primitive insectivore". (Parentheses ours.) On the basis of embryological characters of four microchiroptera n families, Gopalakrishna (1958) mentioned, "the Megachiroptera and Microchiroptera are not as divergent as formerly believed. Many similarities and transitional char- acters are now apparent between the two sub-or- ders".
Luckett (1979), making an analysis of anatomical and embryological characters, sug- gested that the group Chiroptera is monophyletic, but he placed Molossidae as far removed from Pteropodidae. Gopalakrishna and co-workers (1981, 1983, 1987, 1988, 1989) examined the anatomy of the female genitalia, blastocyst-uterus relationship and development of foetal membranes of several families of bats, and postu- lated that not only is Chiroptera a monophyletic group but that the taxonomic hierarchy currently maintained by systematists needs some changes. One such suggestion was that the systematic posi- tion of Molossidae needs to be re-examined.
The basic premise for the present report is that in eutherian mammals embryological charac- ters are far more conservative than are mor- phological characters, since development takes place in a constant environment within the uterus, while morphological characters are directly in- fluenced by the environment and are therefore adaptive. Hence, similarities in embryological characters, according to Mossman (1937, 1953), indicate a closer phylogenetic affinity than similarities in morphological characters.
The present report is based on recent publi- cations and ongoing work in this laboratory on the embryology of four molossid species, namely Chaerephon plicata (Gopalakrishna et al. 1989),
SYSTEMATIC POSITION OF MOLOSSIDAE
5
I- c
Fig. 1. a-c. Uterus-blastocyst relationship at the time of implantation in (a) Pteropodidae, (b) Molossidae and (c) Vesper- tilionidae. The dark circle with a white central area represents the embryonic mass containing the primitive amniotic cavity, mes : mesometrium; ut 1 : uterine lumen.
Tadarida aegyptiaca (Sandhu 1986), Tadarida trageta and Molossus major aztecus (M. molos- sus ) (Gopalakrishna and Badwaik in press) and comparing the results with what is known of the embryology of other relevant families, namely Pteropodidae and Vespertilionidae.
Such a comparison reveals that the molos- sids share more embryological characters with pteropodids than with vespertilionids. Among pteropodids, implantation of the blastocyst is part- ly interstitial with the embryonic mass oriented towards the lateral side in Pteropus giganteus
giganteus (Moghe 1951). In Rousettus les- chenaulti (Karim 1976) and Cynopterus sphinx (pers. obs.) blastocyst implantation is superficial and the embryonic mass is oriented towards the tubo-uterine junction, which is sub-terminal and towards the lateral side of the uterus. The orienta- tion of the embryonic mass in the implanting blastocyst is lateral in all the molossid bats (San- som 1932, Pendharkar and Gopalakrishna 1983, Sandhu 1986).
Secondly, in Pteropodids and all molossids the blastocyst establishes contact with the uterine
6
JOURNAL, BOMBAY NATURAL HIST. SOCIETY, Vol. 89
Fig. 2a-c. Definitive arrangement of foetal membranes in (a) Pteropodidae, (b) Molossidae and (c) Vespertilionidae
all. pi : allantoic placenta; am: amnion; exo: exocoelom; tr. om: trilaminar omphalopleure; y-s: yolk sac; y-s.c.: yolk sac cavity;
y-s. spl: yolk sac splanchnopleure. Other legends as in Fig. 1.
wall on all sides, resulting in the obliteration of the uterine lumen at the level of implantation. This situation differs from what obtains in all vesper- tilionids, in which the blastocyst attaches itself to the antimesometrial side of the uterus by its embryonic pole, and the abembryonic region of the wall of the blastocyst lies freely hanging into the uterine lumen on the mesometrial side of the uterus (Fig. la-c).
In both Pteropodidae and Molossidae an ex- tensive yolk sac placenta is formed on all sides of the uterus except where the embryonic plate inter-
venes between the yolk sac and the uterine wall. This is at first non-vascular, but soon becomes vascularised and forms the chorio-vitelline placenta during early stages of pregnancy. In Vespertilionidae, on the other hand, only the lateral wall of the yolk sac forms the yolk sac placenta, while the abembryonic region remains non-vascular and free.
The unique modification of the yolk sac into a solid gland-like structure in both Pteropodidae (van der Sprenkel 1932, Moghe 1951, 1956; Wimsatt 1954, Gopalakrishna and Karim 1974,
SYSTEMATIC POSITION OF MOLOSSIDAE
7
Karim et. al. 1979, Gopalakrishna and Karim 1981) and Molossidae (Stephens 1962, Stephens and Easterbrook 1968, 1969, 1971; Sandhu 1986, Gopalakrishna et al. 1989) is unmatched in any other family of Chiroptera - and in fact in any other mammal. The yolk sac splanchnopleure be- comes free and undergoes progressive collapse until the yolk sac lumen is completely obliterated in Pteropodidae. In Molossidae the yolk sac lumen is reduced to a few isolated, very narrow streak-like spaces here and there within the solid yolk sac.
In both families the endodermal cells under- go enormous hypertrophy and form acinus-like groups; the mesodermal cells form the loose matrix and the outer covering to the gland-like yolk sac. In Vespertilionidae (Ramaswami 1933, Wimsatt 1945, Enders and Wimsatt 1968, Gopalakrishna 1950, Gopalakrishna and Sapkal 1974 Ramakrishna and Madhavan 1977, Gopalakrishna et al in press) the yolk sac lumen persists as a continuous space between the proximal invaginated, folded vascular splanchnopleure and the distal free trilaminar om- phalopleure (Fig. 2a-c). The uterine lumen per- sists on the mesometrial aspect of the uterus throughout gestation.
The definitive allantoic placental disc is mesometrial in both Pteropodidae and Molos- sidae, whereas it is squarely antimesometrial in
Refer
Ellerman, J.R. & Morrison -Scott, T.C.S. (1951): Checklist of Palaearctic and Indian mammals. British Museum of Natural History, London.
Enders, A.C. & Wimsatt, W.A. (1968): Formation and struc- ture of the haemodichorial chorio-allantoic placenta of the bat, Myotis lucifugus lucifugus. Amer. J. Anat. 122: 453- 489.
Gopalakrishna, A. (1950): Studies on the embryology of Microchiroptera, Part V - Placentation in the vesper- tilionid bat, Scotophilus wroughtoni (Thomas). Proc. Ind. Acad . Sci. 31: 235-251.
Gopalakrishna, A. (1958): Foetal membranes in some Indian Microchiroptera./. Morph. 102: 157-197. Gopalakrishna, A. & Badwaik, N. (1987): Is Rhinopoma a rhinolophoid bat? /. Bombay nat. Hist. Soc. 84(3): 664-670.
Vespertilionidae (Fig. 2a-c). With respect to the histological structure, the placenta is en- dotheliochorial in Pteropus and Cynopterus and haemochorial in Rousettus. In molossids a diffuse endotheliochorial chorio-allantoic placenta oc- curs concurrently with a small mesometrially lo- cated discoid placenta until about the third quarter of gestation. The discoid placenta is haemochorial. However, during the final quarter of gestation the diffuse endotheliochorial allan- toic placenta disappears, and only the mesometrially located discoid haemochorial placenta persists. Molossid bats, therefore, develop both endotheliochorial and haemochorial allantoic placentae. In all vespertilionids the placenta is haemochorial.
It is thus evident that embryological similarities between Molossidae and Pteropodidae and differences between Molos- sidae and Vespertilionidae suggest a closer relationship between Pteropodidae and Molos- sidae than between Molossidae and Vesper- tilionidae. It is, therefore, suggested on purely embryological grounds that Molossidae be separated from the Super-family Vespertilionidae and be placed somewhere between Pteropodidae and Emballonuridae.
We thank the U.G.C. and the C.S.I.R., New Delhi for financial assistance for carrying out this work.
ENCES
Gopalakrishna, A. & Badwaik, N. (in press): Foetal membranes and placentation in two species of molossid bats. Curr. Sci .
Gopalakrishna, A. & Chari, G.C. (1983): A review of the taxonomic position of Miniopterus based on embryological characters. Curr. Sci 52: 1176-1180. Gopalakrishna, A. & Karim, K.B. (1974): The yolk sac gland in the Indian fruit bat, Rousettus leschenaulti (Desm.). Curr. Sci 41: 639-641.
Gopalakrishna, A. & Karim, K.B. (1981): Female genital anatomy and the morphogenesis of the foetal membranes of Chiroptera and their bearing on the phylogenetic relationships of the group. Golden Jubilee Volume, Nat Acad. Sci India: 379-428. Gopalakrishna, A., Pendharkar, Y.D. & Badwaik, N. (1989): Morphogenesis of the foetal membranes and placentation in the Indian molossid bat,
8
JOURNAL, BOMBAY NATURAL HIST SOCIETY, Vol. 89
Chaerephon plicata (Buchanan). Proc. Ind. Acad. Sci. 98(3): 149-166.
Gopalakrishna, A., Phansalkar, R.B. Madhavan, A. & Bad- waik, N. (1988): Pre-implantation stages of develop- ment of Pipistrellus ceylonicus chrysothrix (Wroughton). Trends in Life Sci. 3: 47-50.
Gopalakrishna, A., Phasalkar, R.B. Madhavan, A. & Bad- waik, N. (in press): Development of the foetal membranes and placentation of the Indian vesper- tilionid bat, Pipistrellus ceylonicus chrysothrix (Wroughton). Trends in Life Sci.
Gopalakrishna, A. & Sapkal, V.M. (1974): The foetal membranes in the Indian pipistrelle, Pipistrellus dormeri.J. Zool. Soc. India. 26: 1-9.
Hill, J.E. & Smith, J.D. (1985): Bats - A natural history. Henry Ling Ltd., Dorchester, Dorset, U.K.
Honacki, J.H., Kinman, K.E. & Koeppl, J.W. (1982): Mam- mal species of the World. Allen Press Inc., Lawrence, Kansas, U.S.A.
Jones, F.W. (1917): The genitalia of Chiroptera. J. AnaL 51: 36-60.
Karim, K.B. (1976): Embryology of some Indian Chiroptera. Unpublished D.Sc. Thesis, Nagpur University.
Karim, K.B., Wimsatt, W.A., Enders, A.C. & Gopalakrish- na, A. (1979): Electron microscopic observations on the yolk sac of the Indian fruit bat, Rousettus les- chenaulti (Desmarest) (Pteropida €).Anat.Rec. 195 (3): 493-510.
Koopman, K.F. (1984): Asynopsis of the families of bats, Part VIII. Bat Res. News. 25: 25-27.
Luckett, W.P. (1979): The use of foetal membrane data in assessing chiropteran phylogeny. Proc. Fifth Int. Bat Res. Conf. 245-265.
Moghe, M.A. (1951): Development and placentation in the Indian fruit bat, Pteropus giganteus giganteus (Brun- nich). Proc. Zool. Soc. London 121: 703-721.
Moghe, M.A. (1956): On the development and placentation of the megachi ropteran bat, Cynopterus sphinx gan- geticus. Proc. Nat. Inst. Sci. India 22: 48-55.
Mossman, H.W. (1937): Comparative morphogenesis of foe- tal membranes and accessory uterine structures. Con- trib. Embryol. Carnegie Inst. Washington 26: 127-246.
Mossman, H.W. (1953): The genital system and the foetal membranes as criteria for mammalian phylogeny and taxonomy. Jour. Mammal. 34: 289-298.
Pendharkar, Y.D. & Gopalakrishna, A. (1983): Observa- tions on the early development and implantation of the blastocyst of Tadarida plicata plicata (Buchanan) (Molossidae). /. Shivaji Univ. (Sc.) 21: 179-188.
Ramakrishna, P.A. & Madhavan, A. (1977): Foetal membranes and Placentation in the vespertilionid bat, Scotophilus heathi (Horsefield). Proc. Ind. Acad. Sci. 86: 117-126.
Ramaswami, L.S. (1933): Some stages of the placentation in Vesperugo leisleri (Kuhl). Half-yrly. Jour. Mysore Univ. 7: 1-41.
Sandhu, S.K. (1986): Studies on the embryology of some Indian Chiroptera. Ph.D. thesis, Nagpur University (un- published).
Sansom, G.S. (1932): Notes on some early blastocysts of the South American Molossid bat, Molossus. Proc. Zool. Soc. London. Part 1: 113-118.
Simpson, G.G. (1945): The principles of classifications and a classification of mammals. Bull. Amer. Mus. Nat. Hist 85: 1-350.
Stephens, R.J. (1962): Histology and histochemistry of the placenta and foetal membranes in the bat, Tadarida brasiliensis cynocephala. Amer. J. AnaL 111: 259-286.
Stephens, R.J. & Easterbrook, N. (1968): Development of the cytoplasmic membranous organelle in the endoder- mal cells of the yolk sac of the bat, Tadarida brasiliensis cynocephala. J. Ultrastr. Res. 24: 239-248.
Stephens, R.J. & Easterbrook, N. (1969): Anew cytoplasmic organelle related to both lipid and glycogen storage material in the yolk sac of the bat, Tadarida brasiliensis cynocephala. Amer. J. Anat. 124: 47-58.
Stephens, R.J. & Easterbrook, N. (1971): Ultra-structural differentiation of the endodermal cells of the yolk sac of the bat, Tadarida brasiliensis cynocephala. AnaL Rec. 169: 207-242.
VanderSprenkel, H.B. (1932): Persistenz der Dottergefasse in den Embryo nen der Fledermause und ihre Ursache. Zeitschr.f. MikrA.nL Forsch. 28: 185-268.
Wimsatt, W.A. (1945): The placentation in the vespertilionid bat, Mytis lucifugus lucifugus. Amer. J. Anat. 77: 1-51.
Wimsatt, W.A. (1954): The fetal membranes and placentation of the Tropical American vampire bat, Desmodus rotundus murinus. Acta Anat. 21: 285-341.
A STUDY OF THE FOOD HABITS OF SIX ANURAN TADPOLES1
A.G. Sekar2 ( With two text-figures)
The intestinal contents of tadpoles of six anuran species, collected from different waterbodies, were studied to find out their food in natural habitat The gut contents revealed that all the tadpoles studied were largely herbivorous and ingested 36 genera of algae. The food is apparently determined by the nature of the habitat. It seems that in nature the tadpoles studied fed randomly, without any discrimination. The existing literature indicates that these tadpoles have the capacity to adj ust their feeding habits to available food in new environments.
Introduction
All adult amphibians are carnivorous and devour whatever they can overcome, such as Crus- tacea, small fishes, worms and insects. Tadpoles, on the other hand, are largely herbivorous (Cun- ningham 1912). They are fundamentally specialised for suspension feeding (Wassersug 1975) and depend mainly on algae for their food in natural habitats. The laboratory food for tad- poles as suggested by McCann (1932) was raw meat and also aquatic plants like Hydrilla, Ceratophyllum, Lemna and Vallisneria.
The dietary habits of tadpoles have been studied by Kamat (1962), Sabnis and Kolhatkar (1977), Sabnis and Kuthe (1980) and Wassersug et al (1981). Sabnis and Kuthe examined the natural food of tadpoles oiBufo melanostictus by gut analysis. Wassersug et al. described the tad- poles of Philautus sp. in Thailand as macro- phagus, feeding on frog eggs. Literature on natural food of tadpoles is meagre, whereas there is fairly adequate information on the diet of adult frogs (Andrews 1979, Davidson 1916, Isaac and Rege 1975, Joshee 1968, Mohanty-Hejmadi and Acharya 1982, Rangaswamy and Chari- nabasavanna 1972).
The present study examines the dietary com- ponents of tadpoles of Bufo melanostictus (Family Bufonidae), Ramanella montana (Microhylidae), Rana tigerina, Rana limnocharis, Tomopterna breviceps (Ranidae) and Polypedates macula tus (Rhacophoridae) in nature.
Accepted January 1992.
^Bombay Natural History Society, Ilornbill House, Shaheed Bhagat Singh Road, Bombay 400 023.
Material and Methods
Tadpoles were collected from various water- bodies like pools, river, cisterns and ponds in Sanjay Gandhi National Park in Borivli, Bombay (18° 55’N, 72° 54’E) during the monsoon of 1983. The tadpoles were collected with a net and preserved in 10% formalin. In the laboratory they were sorted out into pre-hindlimb and hindlimb stages.
To study the gut content, the intestine was removed, squashed in a small petri-dish and li- quefied by adding 5 ml of water. The fragments of empty intestine were removed and the liquid gut content was analysed under a 320 x micro- scope. The algal materials were identified and counted in viewing area of 0.732 sq. mm. Five similar squares were counted for algal com- ponents, and the average taken. Ten tadpoles per stage were examined.
Tadpoles of different species were collected from the following waterbodies (Figs. 1, 2).
(1) Bufo melanostictus : Dahisar river and a small stream near Film City. (2) Ramanella mon-4 tana : Dahisar river and roadside pool. (3) Rana tigerina : Pond near Jain temple and rectangular pond atKanheri Hill. (4) Rana limnocharis : Tem- porary puddle and water-logged grassland. (5) Tomopterna breviceps: Dahisar river and corridor of Kanheri cave No.l with 5 cm depth of still water. (6) Polypedates maculatus : Pool near Sanyasi’s hut and temporary puddle.
Results and Discussion
The data on food items in the gut of six tadpole species are shown in Tables 1-6. All six
10
JOURNAL, BOMBAY NATURAL HIST SOCIETY, Vol. 89
Figs. 1-2. Location of collection sites of tadpoles at Sanjay Gandhi National Park and Kanheri Caves.
FOOD HABITS OF SIX ANURAN TADPOLES
11
Table 1
PERCENTAGE OF FOOD ITEMS IN THE GUT OF TADPOLES OF Bufo melanostictus
|
Intestinal contents |
Dahisar river |
Stream near Film City |
||
|
Pre-hindlimb stage |
Hindlimb stage |
Pre -hindlimb stage |
Hindlimb stage |
|
|
Oscillatoria |
1.06 |
6.19 |
9.73 |
2.26 |
|
Scenedesmus |
- |
- |
- |
0.75 |
|
Phacus |
0.30 |
- |
0.54 |
- |
|
Oedogonium |
0.04 |
- |
- |
- |
|
Closterium |
0.23 |
- |
- |
- |
|
Cosmarium |
3.26 |
1.03 |
- |
- |
|
Pinnularia |
74.17 |
63.92 |
74.05 |
47.10 |
|
Navicula |
17.73 |
21.65 |
12.97 |
13.85 |
|
Cymbella |
2.35 |
2.06 |
1.08 |
- |
|
Synedra |
0.76 |
3.09 |
- |
32.49 |
|
Stauroneis |
- |
- |
- |
3.02 |
|
Euchalanis |
- |
- |
1.08 |
- |
|
Nematode worm |
- |
2.06 |
0.54 |
0.25 |
|
Table 2 |
||||
|
PERCENTAGE OF FOOD ITEMS IN THE GUT OF TADPOLES OF Ramanella montana |
||||
|
Intestinal contents |
Dahisar river |
Roadside pool |
||
|
Pre-hindlimb |
Hindlimb |
Pre-hindlimb |
Hindlimb |
|
|
stage |
stage |
stage |
stage |
|
|
Oscillatoria |
0.21 |
0.33 |
- |
- |
|
Spaerella |
- |
- |
- |
15.62 |
|
Oedogonium |
- |
- |
9.37 |
- |
|
Cosmarium |
0.85 |
1.76 |
- |
- |
|
Vaucheria |
- |
- |
3.13 |
6.25 |
|
Pinnularia |
98.50 |
97.23 |
56.25 |
43.75 |
|
Navicula |
0.43 |
0.66 |
12.50 |
12.50 |
|
Spores |
- |
- |
18.75 |
18.75 |
|
Daphnia |
- |
- |
- |
3.15 |
tadpoles were largely herbivorous; 36 genera of algae and four species of animalcules were recorded from the stomach contents. The intes- tines were long and spirally coiled like a watch- spring. Noble (1931) stated that the more car- nivorous tadpoles have a shorter digestive tract than herbivorous species.
Wassersug (1975) reported that the unique morphology of tadpoles is in some way as- sociated with herbivory, plankton feeding, filter feeding, suspension feeding etc. The elongated,
coiled intestines of most tadpoles contrast sharply with the shortened digestive tract of the few known carnivorous, non-feeding, or direct developing forms.
Food items of tadpoles collected from run- ning water differed from those collected from still water. For example the food of Bufo melanostictus tadpoles collected from Dahisar river as well as from the stream near Film City gate varied and certain food items like Eudorina, Opalina, Spirogyra, Ulothrix, Euglena , watermites and
12
JOURNAL, BOMBAY NATURAL HIST SOCIETY, Vol 89
Table 3
PERCENTAGE OF FOOD ITEMS IN THE GUT OF TADPOLES OF Rana tigerina
|
Intestinal contents |
Pond near Jain temple |
Rectangular pond at Kanheri Hills |
||
|
Pre-hindlimb stage |
Hindlimb stage |
Pre-hindlimb stage |
Hindlimb stage |
|
|
Oscillatoria |
7.48 |
1.51 |
0.04 |
0.14 |
|
Spirulina |
- |
- |
1.56 |
0.38 |
|
Scytonema |
- |
- |
0.09 |
0.13 |
|
Eudorina |
- |
7.09 |
1.95 |
1.78 |
|
Pediastrum |
2.04 |
3.92 |
- |
- |
|
Ankistrodesmus |
- |
5.13 |
- |
- |
|
Selenastrum |
- |
- |
11.64 |
12.94 |
|
Tetraedron |
- |
2.57 |
0.58 |
1.95 |
|
Scenedesmus |
34.69 |
12.08 |
77.83 |
81.09 |
|
Ulothrix |
- |
- |
0.24 |
0.23 |
|
Phacus |
2.72 |
28.70 |
0.44 |
0.51 |
|
Oedogonium |
0.68 |
0.90 |
0.14 |
0.10 |
|
Closterium |
- |
3.63 |
- |
- |
|
Euastrum |
- |
0.15 |
- |
- |
|
Cosmarium |
- |
- |
0.04 |
0.10 |
|
Pinnularia |
26.53 |
22.96 |
0.29 |
0.31 |
|
Navicula |
3.40 |
0.75 |
0.44 |
0.23 |
|
Spores |
21.76 |
8.00 |
4.45 |
0.38 |
|
Monostyla |
- |
- |
- |
0.07 |
|
Nematode worm |
0.68 |
2.26 |
- |
- |
|
Daphnia |
- |
0.15 |
0.19 |
- |
|
Tardigrada |
- |
0.15 |
- |
- |
Pleurococcus were absent. However, Sabnis and Kuthe (1980) have reported these food items in B. melanostictus collected from a pond.
Similarly, tadpoles of Tomopterna breviceps obtained from Dahisar river had eaten only a few varieties of food items, whereas tadpoles of the same species collected from still water in the corridor of Kanheri Cave 1 had consumed more food items. The food is apparently determined by the nature of the habitat. Running water, general- ly, contains less micro fauna than ponds and pud- dles. Tonapi (1980) also noted the conspicuous absence of many rooted plants and the relative absence of plankton in running water. But tad- poles otRamanella montana had fed on a limited number of food items though they were collected from stagnant water (rain pool). This might be due to the nature of the pool (fresh and muddy water, with meagre algal components).
The food items differed for different loca- tions, even for the same species of tadpole. Oedogonium, Closterium and Cosmarium were found in Bufo melanostictus tadpoles of Dahisar river but not in tadpoles from the stream near Film City. In Rana tigerina tadpoles, Scenedesmus was noticed both from the pond near Jain temple and the rectangular pond at Kanheri caves. However, there were several differences in food items from tigerina tadpoles from these two locations - Spirulina, Scytonema, Selenastrum, Ulothrix, Cosmarium and Monostyla were found in Kanheri caves tadpoles, but not in Jain temple tadpoles. There are similar location-related differences in the case of other species also.
From this data it seems that tadpoles feed randomly without any discrimination, on whatever is available in the particular waterbody where they grow up. Costa and Balasubramanium
FOOD HABITS OF SIX ANURAN TADPOLES
13
Table 4
PERCENTAGE OF FOOD ITEMS IN THE GUT OF TADPOLES OF Rana limnocharis
|
Intestinal contents |
Temporary puddle |
Waterlogged grassland |
||
|
Pre hindlimb stage |
Hindlimb stage |
Pre hindlimb stage |
Hindlimb stage |
|
|
Oscillatoria |
0.93 |
2.38 |
0.26 |
1.00 |
|
Spirulina |
- |
- |
0.52 |
0.25 |
|
Lyngbya |
1.85 |
1.70 |
3.65 |
3.72 |
|
Anabaena |
0.93 |
0.68 |
- |
- |
|
Scytonema |
- |
- |
0.52 |
0.50 |
|
Tolypothrix |
1.39 |
1.36 |
- |
- |
|
Sphaerella |
2.78 |
3.40 |
1.30 |
1.49 |
|
Oocystis |
- |
- |
3.65 |
3.22 |
|
Ankistrodesmus |
- |
- |
0.52 |
0.25 |
|
Scenedesmus |
4.17 |
0.34 |
- |
- |
|
Oedogonium |
5.55 |
4.08 |
1.04 |
1.00 |
|
Phacus |
2.31 |
1.20 |
- |
- |
|
Zygnema |
- |
- |
0.26 |
0.25 |
|
Closterium |
11.11 |
8.84 |
3.65 |
4.96 |
|
Pleurotaneum |
- |
- |
2.08 |
1.00 |
|
Euastrum |
1.38 |
1.70 |
5.73 |
5.46 |
|
Microsterias |
- |
- |
1.30 |
0.25 |
|
Cosmarium |
26.85 |
15.99 |
13.54 |
18.11 |
|
Staurastrum |
3.70 |
2.38 |
2.60 |
3.47 |
|
Onychonema |
- |
0.34 |
9.11 |
7.20 |
|
Desmidium |
- |
0.34 |
3.13 |
6.20 |
|
Pinnularia |
24.07 |
41.84 |
33.59 |
30.52 |
|
Navicula |
12.04 |
10.20 |
7.55 |
6.70 |
|
Cymbella |
0.93 |
2.72 |
5.20 |
3.72 |
|
Synedra |
- |
0.68 |
- |
- |
|
Monostyla |
- |
- |
0.52 |
0.25 |
|
Daphnia |
- |
- |
0.26 |
0.25 |
(1965) showed from stomach content analyses that Rhacophorus cruciger larvae are qualitative- ly non-discriminant in the food that they ingest. Similar analyses for Rana clamitans tadpoles showed that these larvae are qualitatively and quantitatively non-discriminant in their suspen- sion feeding (Farlowe 1928). But, controversially, Kamat (1962) reported that tadpoles did not feed on all available algae. He found in the laboratory that tadpoles did not feed on certain algae like Chara, Cladophora, Pithophora. However, more work is required to prove that tadpoles show food preferences in natural environment.
Some diatoms {Pinnularia, Navicula , Scenedesmus, Closterium , Cosmarium) which are suspended in the water, were found in the present study to have been fed on in good percentage. Presumably, these were abundant in that par- ticular period and were therefore taken by the tadpoles. Wassersug (op. cit.) stated that tadpoles are highly specialised suspension feeders, adapted for utilizing rapid increases in primary production of a food source. Such sources are probably coupled to environmental fluctuations and available for only a limited amount of time during any year.
2
14
JOURNAL, BOMBAY NATURAL HIST. SOCIETY, Vol. 89
Table 5
PERCENTAGE OF FOOD ITEMS IN THE GUT OF TADPOLES OF Tomopterna breviceps
|
Intestinal contents |
Dahisar river |
Corridor of Cave 1 at Kanheri |
||
|
Pre-hindlimb s'tage |
Hindlimb stage |
Pre-hindlimb stage |
Hindlimb stage |
|
|
Gloeocapsa |
0.63 |
- |
1.70 |
5.65 |
|
Oscillcitoria |
21.65 |
19.59 |
19.57 |
17.39 |
|
Spirulina |
- |
- |
6.80 |
1.74 |
|
Lyngbya |
- |
- |
2.55 |
0.87 |
|
Scytonema |
- |
- |
25.95 |
15.21 |
|
Oedogonium |
1.91 |
0.50 |
5.95 |
1.74 |
|
Phacus |
- |
- |
0.85 |
1.30 |
|
Closterium |
- |
- |
- |
0.43 |
|
Cosmarium |
- |
- |
4.25 |
1.74 |
|
Spirogyra |
2.54 |
2.51 |
- |
- |
|
Pinnularia |
22.29 |
32.16 |
8.08 |
8.69 |
|
Navicula |
37.57 |
32.66 |
3.40 |
8.69 |
|
Cymbella |
- |
- |
1.70 |
1.30 |
|
Synedra |
12.10 |
11.05 |
- |
1.74 |
|
Spores |
- |
- |
17.87 |
32.17 |
|
Nematode worms |
1.70 |
1.50 |
0.43 |
1.30 |
|
Tardigrada |
- |
-- |
0.85 |
- |
McCann (1932) suggested raw meat as a food for tadpoles reared in the laboratory. Sekar (1990) fed tadpoles of the Malabar gliding frog Rhacophorus malabciricus with earthworms, meat and snail flesh in the laboratory to rear them. These non-algal food items were readily accepted.
Wassersug et al. (1981) reported that the larvae of Tlieloderma stellatum (Rhacophoridae) of Thailand, which developed in tree holes con- taining decomposing leaves, fed on amoeba tests, fungal spores, lepidopteran scales etc.
In contrast, tadpoles of Philautus sp., which developed in a tree hole containing less than 75 ml of water without any indirect source of food, appear to rely on introduced frog eggs for food. It seems that tadpoles might have the capacity to adapt to a new environment and adjust to feeding on the food available in that environment.
There was notably no difference between the pre-hindlimb and hindlimb stages; food items were similar in both stages.
Conclusions
Gut analysis of six species of tadpoles led to the following conclusions.
(1) All the tadpoles studied were largely her- bivorous in food habits; a variety of algal com- ponents constituted the major food items. (2) Tad- poles from still water fed on more food items than those obtained from running water. (3) In nature, the tadpoles studied fed randomly, without any qualita- tive discrimination. (4) Tadpoles which are fun- damentally specialised for suspension feeding fed more on diatoms like Pinnulario, Navicula, Cos- marium etc. (5) Food items were almost similar in both pre-hind limb and hindlimb stages.
Acknowledgements
I thank Mr J.C. Daniel, former Curator of the BNHS, for his constant encouragement during the' study; Dr Robert B. Grubh, Research Coor- dinator, BNHS for allowing me to use the project microscope, and Mr Ajay Varadachary for read- ing through the manuscript.
FOOD HABITS OF SIX ANURAN TADPOLES
15
Table 6
PERCENTAGE OF FOOD ITEMS IN THE GUTS OF TADPOLES OF Polypedates maculatus
|
Intestinal contents |
Pool near Sanyasi’s hut |
Temporary puddle |
||
|
Pre-hindlimb stage |
Hindlimb stage |
Pre-hindlimb stage |
Hindlimb stage |
|
|
Gloeocapsa |
- |
- |
1.01 |
- |
|
Oscillatoria |
4.79 |
6.61 |
2.53 |
0.39 |
|
Lyngbya |
- |
- |
1.01 |
0.39 |
|
Sphaerella |
2.17 |
3.78 |
4.04 |
1.76 |
|
A nJcistrodesmus |
0.09 |
- |
- |
- |
|
Scenedesmus |
3.44 |
3.78 |
7.07 |
5.69 |
|
Ulothrix |
- |
- |
0.25 |
0.19 |
|
Oedogonium |
16.00 |
17.12 |
2.77 |
0.76 |
|
Phacus |
7.78 |
6.85 |
0.76 |
0.19 |
|
Spirogyra |
- |
- |
2.70 |
1.96 |
|
Closterium |
2.44 |
1.18 |
- |
- |
|
Pleurotaenium |
- |
- |
0.25 |
- |
|
Euastrum |
- |
- |
1.26 |
0.98 |
|
Microsterias |
- |
- |
0.76 |
- |
|
Cosmarium |
23.32 |
12.87 |
20.96 |
26.93 |
|
Staurastrum |
2.99 |
2.72 |
1.76 |
2.95 |
|
Pinnularia |
8.41 |
6.14 |
28.03 |
18.86 |
|
Navicula |
14.92 |
10.86 |
6.31 |
18.23 |
|
Cymbella |
- |
- |
3.03 |
2.16 |
|
Synedra |
- |
- |
8.33 |
2.16 |
|
Colonies |
- |
- |
4.55 |
13.75 |
|
Spores |
13.38 |
27.74 |
- |
- |
|
Monostyla |
0.27 |
- |
0.76 |
0.98 |
|
Nematode worms |
- |
- |
0.25 |
0.19 |
|
Daphnia |
- |
0.35 |
- |
- |
|
Unidentified |
- |
- |
1.76 |
0.58 |
References
Andrews, M.I. (1979): Food of Ranci hexadactyla Lesson. J. Bombay nat. Hist Soc. 76: 175- 179.
Costa, H.H. & Balasubramanium, S. (1965): The food of the tadpoles of Rhacophorus cruciger cruciger (Blyth) in Ceylon .J. Sci. 5: 105-109 (quoted in Wassersug 1975).
Cunningham, J.T. (1912): Animal life: An evolutionary natural history. Reptiles, Amphibia, Fishes and Lower Chordata. Methuen and Co. Ltd., London.
Davidson, N. (1916): Food of the Bull Frog. J. Bombay nat Hist. Soc. 25: 152-153.
Farlowe, V. (1928): Algae ponds as determined by an ex- amination of the intestinal contents of tadpoles. Biol. Bull. 55: 443-448 (quoted in Wassersug 1975).
Isaac, S. & Rege, M.S. (1975): Food of Ram tigerina (Daud.). J. Bombay nat. Hist. Soc. 72: 143-157.
Joshee, A.K. (1968): Food habits of the Bull frog Rana tigerina (Daud). J. Bombay nat. Hist. Soc. 65: 498-501.
Kamat, N.D. (1962): On the intestinal contents of tadpoles and algae of small ponds. Curr. Sci. 31: 300.
McCann, C. (1932): Notes on Indian Batrachians.7. Bombay nat Hist. Soc. 36: 152-180.
Mohanty-Hejmadi, P. & Acharya, B.K. (1982): Observations on food habits of six species of Indian frogs. J. Bombay nat. Hist. Soc. 79: 120-124.
Noble, G.K. (1954): The biology of the Amphibia. Dover Publications Inc., New York.
16
JOURNAL, BOMBAY NATURAL HIST. SOCIETY, Vol. 89
Rangaswamy, H.R. & Channabasavanna, G.R (1972): Food and feeding habits of the Toad Bufo melanostictus Schneider. J. Bombay nat. Hist. Soc. 70: 558-563.
Sabnis, J.H. & Kolhatkar, B.L. (1977): Observations on the food preference oiRana cyanophlyctis tadpoles. Comp. Physiol. Ecol. 2(4): 232-233.
Sabnis, J. H. & Kutiie, K.S. (1980): Observations on food and growth otBufo melanostictus tadpoles./. Bombay nat Hist Soc. 77: 21-25.
Sekar, A.G. (1990): Observations on the developmental stages of ladpoles of the Malabar gliding frog Rhaeophorus malabaricus Jerdon, 1890
(Anura:Rhacophoridae)J. Bombay nat. Hist. Soc. 87: 223-226.
Tonapi, G.T. (1980): Fresh water animals of India (An Ecological approach). Oxford & IBH Publishing Co.
Wassersug, RJ. (1975): The adaptive significance of the tadpole stage with comments on the maintenance of complex life cycles in Anurans. Am$r. Zool. 15: 405-417.
Wassersug, R.J., Frogner K.J., & Inger, R.F. (1981): Adaptations for life in tree holes by Rhacophorid tadpoles from Thailand. Jour, of Herpetology 15: 41-52.
I. CLADOCERA OF KEOLADEO NATIONAL PARK, BHARATPUR,
AND ITS ENVIRONS1
K. Venkataraman2 (With forty-nine text-figures)
A study made on collections of zooplankton from shallow waters and ponds in and around Keoladeo National Park, Bharatpur, yielded 39 species of Cladocera, of which 25 are recorded for the first time from Rajasthan. Some selected species recorded in the present study are illustrated and described.
Introduction
Very little is known regarding the occurrence of different species of Cladocera in Rajasthan, particularly the Keoladeo National Park, which has a wide range of freshwater habitats and at- tracts various migratory birds from different parts of the world. The important earlier works on Cladocera of Rajasthan are those of Biswas (1964), Nayar (1971) and Venkataraman (1988, 1990). Ali and Vijayan (1983) studied the general limnology, primary productivity and secondary productivity in Keoladeo National Park. There are also some records of the protozoan species (Mahajan et al. 1980a), benthic fauna (Mahajan et al. 1980b) and dynamics of zooplankton (Mahajan et al. 1980c) made in freshwater habitats of the Keoladeo National Park (KNP).
The material for the present study was col- lected periodically from eight different places in the Park and 30 ponds and ditches of Rajasthan. The collections revealed 39 species of Cladocera, of which 25 are new records to Keoladeo National Park and Rajasthan. Short description including illustration of diagnostic features of a few inter- esting species is given in this paper. The impact of introduced fauna is also discussed.
Material and Methods
591 zooplankton samples were collected during the years 1984-85 from littoral and lim- netic regions of freshwater habitats of Keoladeo National Park (27° 7.6' to 27° 12.2' N, 77° 29.5'
Accepted May 1990.
2Zoological Survey of India, Andaman and Nicobar Regional Station, Port Blair 744 101. Present address: Zoological Survey of India, M-Block, New Alipore, Calcutta 700 053.
to 77° 39.9' E) and roadside ponds and ditches in and around Bharatpur, Rajasthan. The collections were made with 80 pm mesh size plankton nets of 30 cm upper diameter by taking both vertical and horizontal hauls. All samples were examined with a binocular microscope and the species were separated. Temporary slides were made in glycerine for confirming diagnosis. Drawings were made with a camera lucida and the measure- ments taken using a calibrated ocular micrometer. A list of species recorded is given in Table 1.
KNP has a wide range of freshwater habitats. About 10% of the land is covered with water that comes from a reservoir during the rainy season. This reservoir receives and retains faunal ele- ments from flowing and standing waters used for irrigation via Gambir and Banganga rivers. The climate is subtropical and temperate, with 1200 mm annual rainfall. There were 120 rainy days in the year June 1984 to May 1985. The mean annual temperature was 22° C; the lowest temperature (3°C) was recorded on 14 January and the highest (42°C) on 27 July 1984.
Description of Females of Selected Species Pseudosida bidentata Herrick, 1884 (Figs. 1-2)
Size: 1.20 mm. Body elongated oval; head short; eye relatively small and situated near antero-ventral comer. Antennules unsegmented, long and attached to postero-ventral part of head. Antenna not extending beyond posterior margin of valves. Ventral margin with a series of long setae followed by a series of spinules on postero- ventral corner. Postabdomen short and broad. Lateral side with 10 groups of spines. Claw long, curved dorsally; convex surface serrated; concave surface with series of short setules and three basal
18
JOURNAL, BOMBAY NATURAL HIST. SOCIETY, Vol. 89
Table 1
LIST OF CLADOCER A RECORDED FROM KEOLADEO NATIONAL PARK AND ITS ENVIRONS
Family Sididae
*1. Pseudosida bidentata Richard
*2. Latonopsis australis Sars
3. Diaphanosoma excisum Sars
*4. Diaphanosoma sarsi Richard
*5. Diaphanosoma senegalensis (Gauthier)
Family Daphnidae
6. Daphnia similis Claus *1. Daphnia longispina Muller
8. Daphnia lumholtzi Sars
9. Simocephalus vetulus elizabethae (King) * 10. Simocephalus acutirostratus Sars
11. Ceriodaphnia cornu ta Sars
12. Ceriodaphnia reticulata (Jurine)
13. Scapholeberis kingi Sars Family Macrothricidae
14. llyocryptus spinifer Herrick
15. Macrothrix spinosa King
16. Macrothrix triserialis (Brady)
*17. Grimaldina brazzai Richard *18. Guernella raphalis Richard Family Moinidae
19. Moina micrura Kurz
*20. Moinodaphnia macleayii Richard
Family Bosminidae
*21. Bosminopsis deitersi Richard
Family Chydoridae
*22. Alona costata Sars *23. Alona davidi Richard *24. Alona monacantha Sars *25. Alona karua King 26. Alona verrucosa Sars *27. Camptocercus australis Sars 28. Chydorus eurynotus Sars *29. Chydorus pan’us Daday *30. Chydorus ventricosus Daday *31. Dadaya macrops (Daday)
*32. Dunhevedia crassa King *33. Euryalona orientalis (Daday)
*34. Kurzia longirostris (Daday)
*35. Leydigia australis Sars 36. Leydigia acahthocercoides (Fischer) *37. Indialona ganapati Petkovski *38. Oxyurella sinhalensis (Daday)
*39. Pluroxus similis Vavra
spines. Occurs in all types of habitat except in turbid ponds.
Latonopsis australis Sars, 1888 (Figs. 3-4)
Size: 1.15 mm. Body oblong. Head short and indistinctly separated from the body. Eye small, situated near antero-dorsal end of head. Ocellus small and situated near base of labrum. Anten- nules long and segmented, attached to antero- ventral comer of head. Valves slightly convex dorsally and broadly rounded ventrally. Ventral margin with a series of long setae. Postabdomen short without anal denticles, lateral surface with a series of 8-10 denticles. Claw curved dorsally with two long basal spines. Occurs in all types of habitat except in turbid ponds.
Diaphanosoma excisum Sars, 1885 (Figs. 5-6)
Size: 1.05 mm. Head large and rounded anteriorly. Eye small. Postero-ventral corner broadly rounded with 5-9 marginal denticles fol- lowed by a series of fine setules. Claw serrated on the distal convex surface; concave surface with three long basal spines.
Very common. Occurs in all types of habitat except in marshes. This species has already been recorded in Rajasthan (Biswas 1971, Nayar 1971).
Daphnia similis Claus, 1876 (Figs. 7-9).
Size: 2.25 mm. Body slightly compressed, elliptical in shape. Head rounded anteriorly, rostrum pointed ventrally; antennules short, at- tached to posterior margin of rostrum. Eye large, situated slightly closer to the a ntero -ventral mar- gin of the head. Valves with a series of spinules on dorsal and ventral margins. Postabdomen nar- row distally with about nine pointed denticles. Rare. Occurs only in turbid ponds of Bharatpur.
Daphnia lumholtzi Sars, 1885 (Figs. 10-11)
Size: 1.52 mm. Head with pointed helmet. Rostrum small, fornix well developed. Body rounded. Dorsal and ventral margins with a series of spines. Postabdomen long and narrow with 9-11 denticles on the dorsal side.
* New records
Figs. 1-17. Cladocera of Keoladeo National Park and environs (females). P- postabdomen; PVC- postero-ventral corner. 1-2. Pseudosida bidentata ; 3-4. Latonopsis australis; 5-6. Diaphanosoma excisum; 7-9. Daphnia similis; 10-11. D. lumholtzi; 12-13. Ceriodaphnia cornuta; 14-15. Scapholeberis kingi ; 16-17. llyocryptus spinifer.
20
JOURNAL, BOMBAY NATURAL HIST. SOCIETY, Vol. 89
This is the commonest Daphnia sp. in the limnetic region of aquatic habitat of KNP. It has been already reported in Rajasthan (Biswas 1971, Nayar 1971).
Ceriodaplmia cornuta Sars, 1885 (Figs. 12-13)
Size: 0.39 mm. Head depressed and separated from the carapace by a dorsal impres- sion. Antennules short and broad, with a long seta and a group of sensory setae on the apex. Eye large, ocellus absent. Postero-dorsal corner of the carapace with two acute and diverging points. Postabdomen with four or five curved denticles. Claw long, gently curved with a series of setules along the concave surface.
Very common. Occurs in all types of habitat, especially shallow ponds of KNP. Both homed and hornless individuals are found together with Diciphanosoma excisum and Moina micrura. This species has already been recorded in Rajasthan (Nayar 1971).
Scapholeberis kingi Sars, 1903 (Figs. 14-15)
Size: 0.59 mm. Body rounded dorsally. Head small and slightly depressed, rostrum rounded and projecting ventrally. Eye large, ocellus small, situated closer to the rostrum than to the eye. Valves with lines and reticulations; posterior ventral margin has a long denticle. Postabdomen broad, dorsal margin with five or six denticles. Claw curved dorsally, with spinules along the concave surface. Common in all types of habitat but never occurs in large numbers.
Ilyocryptus spinifer Herrick, 1882 (Figs. 16-17)
Size: 0.75 mm. Body oval. Head small. Eye large, ocellus small, situated about halfway between eye and base of antennules. Antennules long with a group of sensory setae on distal end. Valves with a series of long feather-likc setae on ventral side. Postabdomen with slight depression in the middle. Preanal margin with eight marginal denticles, postanal margin with 12 denticles up to anal groove and with five long and stout spines on the lateral surface.
Claw with two basal spines. Three spinules present in between the base of claw. Common. Occurs in small numbers in marshes of KNP and in the other areas in and around Bharatpur.
Macrotkrix spinosa King, 1852 (Figs. 18-19)
Size: 0.41 mm. Body round-oval; dorsal margin serrated. Head rounded, ventral mar- gin slightly concave with slightly pointed antero-ventral corner. Antennules short, with a long seta near its base. Eye large, ocellus small and situated much nearer to the base of antennules than to the eye. Postabdomen broadly rounded. Claw short and serrated on the concave surface.
Very common. Occurs in all types of habitat in KNP and in the Bharatpur ponds.
Macrotkrix triserialis Brady, 1886 (Figs. 20-21).
Size: 0.56 mm. Body oval, dorsal margin with a slight cervical depression. Head with a round projection on anterior margin above eye. Eye laige, ocellus small and situated nearer apex of rostrum than eye. Antennules long with a long lateral seta. Antennae short with the longest seta having two or three larger spines in the middle. Ventral margin of the valve with a series of long setae in groups of three. Postabdomen bilobed with rows of spines increasing in size proximally. Claw short and serrated without basal spine.
Very common. Occurs in all the marshes of KNP and in the ponds of Bharatpur.
Grimaldina brazzai Richard, 1892
Size: 0.87 mm. Body quadrangular-oval. Head small, eye large. Ocellus small and situated closer to apex of rostrum than to eye. Antennules long and slightly segmented. Postabdomen bilobed with broadly rounded preanal margin. Postanal margin with two groups of long spines. Preanal comer with 2 large spines followed by a series of short spinules proximally. Claw long with basal spines.
Rare. Only a few specimens occurred in marshes of KNP. It has already been recorded by Venkataraman (1990).
CLADOCERA OF KEOLADEO NATIONAL PARK
21
0.1mm 0,03mm 0,05mm 005mm 02mm
18,20,24,26,28,30.32 19.21.25.33 23,27 29,31 22
Figs. 18-33. Cladocera of Keoladeo National Park and its environs, (females). P - postabdomen.
18-19. Macrothrix spinosa ; 20-21. M. triserialis ; 22-23. Moina micrura; 24-25. Alona davidi; 26-27. A. monacantha; 28-29. A. karua; 30-31. >1. verrucosa. 32-33. Chydorus eurynotus.
22
JOURNAL , BOMBAY NATURAL HIST. SOCIETY, Vol. 89
Guernella raphalis Richard, 1892
Size: 0.38 mm. Body slightly oval. Head concave ventrally, eye large, ocellus situated closer to apex of rostrum. Antennules short and broad with a group of sensory setae at apex. Valves with polygonal reticulations and serrated without setae. Postabdomen slightly bilobed with transverse rows of spinules and without anal den- ticles. Claw short without basal spines.
Not common, but occurs in decaying marshy regions of KNP. It has already been recorded by Venktaraman (1990).
Moina micmra Kurz, 1874 (Figs. 22-23)
Size: 0.73 mm. Head large, rounded with a deep cervical depression posteriorly. Eye large, ocellus absent. Antennules long and movable with a group of sensory setae on apex. Postabdomen with 6-8 ciliated lateral spines. Claw long, slightly curved dorsally, with pecten at base.
Very common. This is the most widely dis- tributed species of Moina. It occurs both in ponds of Bharatpur and shallow regions of KNP.
Moinodaphnia macleayii (King, 1853)
Size: 0.81 mm. Head with distinct cervical depression. Eye large, ocellus small, situated closer to antennules than to eye. Antennules slender with long lateral seta and a group of sensory setae on the apex. Ventral margin of valve rounded with series of short marginal spines. Pos- tabdomen without anal denticles, lateral surface with 8-10 ciliated spines. Claw long with a series of short setules along concave surface.
Not very common. Occurs only in the mar- shes of KNP. It has already been recorded by' Venkataraman (1988).
Bosminopsis deitersi Richard, 1895
Size: 0.38 mm. Body oval. Head rounded, rostrum long with two lateral branches, Eye large. Valves with polygonal reticulations, dorsal mar- gin with cervical depression, ventral margin with a long and pointed marginal spine on the postero- ventral corner. Postabdomen small with 4-6
groups of short spinules. Claw serrated with a large basal spine.
Not common. Occurs in open waters of Ajanbund reservoir, Bharatpur. It has already been recorded by Venkataraman (1988).
Alona davidi Richard, 1895 (Figs. 24-25)
Size: 0.35 mm. Maximum height slightly before middle. Postero-dorsal and postero-ventral corners rounded. Ventral margin projecting in the middle. Rostrum blunt. Antennules not reaching apex of rostrum. Ocellus smaller than eye, situated slightly nearer to the eye than apex of rostrum. Postabdomen with prominent preanal and postanal comers. Preanal corner projecting, anal margin with 8-10 groups of denticles. Claw with short basal spines.
Common. Occurs mainly in marshes with Hydrilla sp. in KNP and roadside ponds of Ban- baretha, Bharatpur.
Alona monacantha Sars, 1901 (Figs. 26-27)
Size: 0.27 mm. Valves with longitudinal lines. Postero-dorsal and postero-ventral comers rounded, postero-ventral comer with 1-3 denticles. Ocellus smaller than eye. Antennules not reaching apex of rostrum. Labrum with small denticle on anterior margin. Postabdomen with distinct preanal corner. Lateral margin with 6 -8 anal denticles followed by 3-4 groups of spines along the anal groove. Claw with long basal spines. Not common. Occurs mainly in marshes of KNP.
Alona karna King, 1853 (Figs. 28-29).
Size: 0.29 mm. Body with distinct lines and polygonal patterns. Postero-ventral corner rounded with 2-3 denticles followed by a row of setules. Antennules not reaching apex of rostrum. Ocellus small, situated closer to eye. Postab- domen broadly rounded. Claw with a short basal spine. Very common. Occurs in marshy habitats of KNP.
Alona verrucosa Sars, 1901 (Figs. 30-31).
Size: 0.28 mm. Body oval. Postero-ventral and postero-dorsal comers rounded. Antennules
CLADOCERA OF KEOLADEO NATIONAL PARK
23
almost reaching apex of rostrum. Ocellus small, situated slightly closer to eye than to apex of rostrum. Postabdomen with 5-6 denticles. Claw with short basal spine.
Common. Occurs in all marshy areas of KNP and in the Bharatpur ponds.
Caniptocercus australis Sars, 1896
Size: 0.82 mm. Head smoothly curved. Postero-ventral margin slightly convex with 3-4 small denticles. Ocellus smaller than eye, situated closer to the eye than to the tip of rostrum. Labrum wedge-shaped and slightly rounded at apex. Postabdomen long with 16- 18 anal denticles. Claws long, slightly curved dorsally and pointed.
Very rare. Occurs in Ghana canal of Keoladeo National Park. This is the first record of its occurrence in the Oriental region. However, the same species has been reported as a new species by Battish (1989) fromRenuka lake. Only the comparison of type-specimens will give a clear picture about the validity of this species.
Chydorus eurynotus Sars, 1901 (Figs. 32-33)
Size: 0.24 mm. Body shape slightly oval. Postero-dorsal and postero-ventral corners dis- tinct. Valves with faint reticulation. Rostrum slightly curved posteriorly. Ocellus smaller than eye, situated closer to eye than to apex of rostrum. Postabdomen with 10-12 short denticles. Claw with two basal denticles.
Common. Occurs in all marshy areas of Keoladeo National Park and in the Bharatpur ponds.
Chydorus parvus Daday, 1898 (Figs. 34-35)
Size: 0.28 mm. Body rounded. Postero- dorsal corner distinct, postero-ventral corner rounded without denticle. Ventral margin with 2-3 chitinous tubercles. Surface of carapace without reticulation. Ocellus smaller than eye and situated closer to eye than to tip of rostrum. Postabdomen with distinct preanal corner. Dorsal margin with 6-8 anal denticles. Claw with two basal spines.
Not common. Occurs in marshy regions of KNP and in the ponds of Bharatpur.
Chydorus ventricosus Daday, 1898 (Figs. 36-37).
Size: 0.76 mm. Body oval. Postero-ventral comer rounded, without denticle. Valves with hexagonal markings. Rostrum long and pointed. Labrum long, curved anteriorly and slightly pointed. Postabdomen long with distinct preanal corner. Dorsal margin with 9-10 anal denticles. Claw setulated along concave surface with two basal spines.
Not common. Occurs in marshes of KNP and in the ponds of Bharatpur.
Dadaya macrops (Daday, 1898) (Figs. 38-39).
Size: 0.48 mm. Body oval with polygonal reticulations. Postero-ventral corner with a dis- tinct denticle. Rostrum short, antennules long. Ocellus and eye large, ocellus situated slightly closer to eye than to apex of rostrum. Postab- domen with 10 groups of irregular sized denticles. Claw with setules on concave surface and a long basal spine.
Common. Occurs in marshes of Keoladeo National Park and roadside ponds of Banbaretha, Bharatpur.
Dunhevedia crassa King, 1853 (Figs. 40-41)
Size: 0.53 mm. Body curved dorsally. Postero-ventral corner of valves with a bifur- cated denticle. Rostrum blunt, labrum rounded with pointed apex. Ocellus small and situated slightly closer to eye than to apex of rostrum. Postabdomen with groups of scattered spinules, claw setulated with one basal spine.
Not common. Occurs in marshes of Keoladeo National Park and in the Ajan bund reservoir.
Euryalona orientalis (Daday, 1898). (Figs. 42-43)
Size: 0.99 mm. Valves with rectangular reticulations. Postero-ventral margin of valves with series of setae. Rostrum blunt, antennules almost reaching apex of rostrum. Labrum rounded with a nipple-like structure on apex. Pos- tabdomen long with 20 pointed denticles decreas-
24
JOURNAL, BOMBAY NATURAL HISTORY SOCIETY Vol. 89
0.01mm 0.1mm 005mm 0.05mm 0.05mm 0.2mm 0.2 mm
36.38,40,45.46,48 43,49 34.39 35,47 41 42 ' 44
0.03mm 37
Figs. 34-49. Cladocera of Keoladeo National Park and its environs (females). P-postabdomen.
34-35. Chydorus parvus ; 36-37. Chydorus ventricosus ; 38-39. Dadaya ma crops', 40-41. Dunhevedia crassa; 42-43. Euryalona orientalis; 44-45. Kurzia longirostris; 46-47. Leydigia acanthocercoides, 48-49. Oxyurella sinhalensis.
CLADOCERA OF KEOLADEO NATIONAL PARK
25
ing in size proximally. Claw long with a very short basal spine. Common among weeds in KNP.
Kurzia longirostris (Daday, 1898) (Figs. 44-45)
Size: 0.45 mm. Valves without longitudinal lines. Rostrum long, antennules short, reaching half of the rostrum. Ocellus smaller than eye, situated nearer to the eye than to the apex of rostrum. Labrum with slightly pointed apex. Pos- tabdomen long with 12 groups of denticles present dorsally; lateral side with 11-12 groups of short setules. Claw long with a short basal spine. Rare. Occurs among the weeds in KNP.
Leydigia acanthocercoides (Fischer, 1854)
(Figs. 46-47)
Size: 0.89 mm. Valves with longitudinal lines. Rostrum blunt, antennules not reaching apex of rostrum. Ocellus smaller than eye, situated closer to the eye than to the apex of rostrum. Labrum rounded with fine setae. Postab- domen with about 18 groups of small denticles, each group consists of 3 or 4 denticles with the distal-most denticle being the longest of each group. Claw with a short basal spine.
Rare, occurs in reddish-brown algal covered ponds of Bharatpur.
Indialona globulosa (Daday, 1898)
Size: 0.36 mm. Valves with striations. Rostrum short and blunt, antennules not reaching the apex. Ocellus smaller than the eye, situated closer to the eye than to the apex of rostrum. Labrum serrated on antero-ventral margin. Pos- tabdomen long with 12 or 13 anal denticles, claw long with 12 or 13 anal denticles, claw long with short basal spine.
Rare. Occurs in small numbers in marshes of KNP and Banbaretha ponds.
Oxyurella sinkalensis (Daday, 1898) (Figs. 48-49)
Size: 0.82 mm. Valves evenly rounded. Rostrum blunt, antennules not reaching the apex. Ocellus smaller than the eye and situated closer to the eye than to the apex of rostrum. Labrum round. Postabdomen long with 10-12 anal denticles
which decrease in size proximally. Claw long with a long basal spine and three short spines proximal to the basal spine. Not common. Occurs in the marshes of KNP.
Discussion
The cladoceran fauna of Keoladeo National Park has some features unique to this region. A total of 39 species of Cladocera belonging to six families is recorded in the present study, of which 25 are recorded for the first time from Rajasthan. An analysis of the published records shows that in a tropical region arounds60 species have been normally recorded, while a temperate region sup- ported around 95 species of Cladocera (Fernando 1980). The six species of limnetic Cladocera are limited in this region compared to temperate regions. As an example, in Ontario at Canada, Brandlova et al. (1972) recorded 18 species of limnetic Cladocera.
Similar figures can be quoted for all the northern temperate regions (Flossner 1972, Manuilova 1964, Scourfield and Harding 1966). In the southern temperate zone, Hebert (1977) found 10 limnetic species of Daphnia in south- eastern Australia. On the other hand, the number of species in Sri Lanka (Fernando 1980), south India (Venka tarn man 1983) as well as other tropi- cal areas of south-east Asia (Fernando 1980) is less than that of temperate regions. However, in Keoladeo National Park lying within the temperate zone, only three species of Daphnia , namely D. similis, D. lum/ioltzi and D. lon- gispina, occurred.
Venkataraman (1983) recorded and com- mented upon six species of non-Indian cladoceran species in south India. Ghetti (1970) also recorded one non-European ostracod species in Italy. More recently Fernando (1980) recorded two unex- pected species from Sri Lanka. Mukhamediev (1951) considered seed (agricultural) as a means of transporting tropical fauna and flora into sub- tropical regions of the USSR.
There is thus enough evidence to show the presence of temperate Cladocera in tropical India (Venkataraman 1983) and Sri Lanka (Fernando
26
JOURNAL, BOMBAY NATURAL HIST SOCIETY, Vol. 89
1980). The present study also reveals the cladoceran species Daphnia similis , Diaphanosoma senegalensis, Camptocercus australis, Leydigia australis and Pluroxus similis as introduced ones. Birds have been considered an important agency for the dissemination of microcrustaceans in freshwaters (Thienemann 1950, Loffler 1963, Smirnov 1974). Keoladeo National Park attracts several aquatic bird migrants (Ali and Vijayan 1983). Perhaps bird
sanctuaries, besides attracting aquatic migratory birds, also provide congenial conditions for the alien fauna and flora to colonise these aquatic ecosystems.
Acknowledgements
I am grateful to J. C. Daniel of BNHS, Bom- bay, Dr T. M. Haridasan of School of Energy Scien- ces, M.K. University, Madurai and to Dr G.C. Rao of Z.S.I., Port Blair for their encouragement.
References
Ali, S. & Vuayan, V.S. (1983): Hydrobiological Research at Keoladeo National Park, Bharatpur. First interim report. Bombay Natural History Society, Bombay.
Battish, S.K. (1989): Entamostracan fauna of Renuka lake, Himachal Pradesh. Geobios New Reports 8: 55-58. Biswas, S. (1964): A new species of cladoceran genus Latona Straus (1820) from Rajasthan, India. Proc. Zool. Soc . Calcutta 17: 149.
Biswas, S. (1971): Fauna of Rajasthan, India. Part II (Crustacea: Cladocera). Rec. Zool. Survey India 63: 95-141. Brandlova, J. Brandl, Z. & Fernando, C.H. (1972): The Cladocera of Ontario with remarks on some species and distribution. Can. J. Zool. 50: 1373-1403.
Fernando, C.H. (1980): The freshwater zooplankton of South- East Asia. Proc. V Symp. Intemat. Soc. Trop. Ecol. Flossner, D. (1972): Krebstiere, Crustacea. Kiemen-und Blat- tfusser, Branchiopoda. Fischlause, Branchiura. Tierwelt. Deutschl. 60: 1-501.
Ghetti, P.F. (1970): The taxonomic significance of Ostrocod larvae stages with examples from Burundi rice fields. i?u//. Zool. 37: 103-119.
Hebert, P.D.N. (1977): A revision of the taxonomy of the genus Daphnia in south-eastern Australia. Aust. J. Zool. 25: 371-398.
Loffler, H. (1963): Bird migration and the spread of Crustacea.
Verh. dt Zool Ges. 27: 311-316.
Mahajan, C.L., Arora, N.K., Sharma, S.D. & Sharma, S.P. (1980a): Drought and drought management in relation to Protozoan fauna in a wetland ecosystems with special reference to Ghana Bird Sanctuary, Bharatpur. Interna- tional Wetland Conference, New Delhi.
Mahajan, C.L., Arora, N.K., Sharma, S.D. & Sharma, S.P. (1980b): The role of benthic fauna in wetland ecosystem
with particular reference to the environmental stress due to drought in Ghana Bird Sanctuary, Bharatpur. Interna- tional Wetland Conference, New Delhi.
Mahajan, C.L., Arora, N.K., Sharma, S.D. & Sharma, S.P. (1980c): Dynamics of zooplankton in wetland ecosystems due to environmental stress with special reference to drought in Ghana Bird Sanctuary, Bharatpur. International Wetland Conference, New Delhi.
Manuilova, E.F. (1964): Cladoceran Fauna of the USSR-Nauka. Moscow and Leningrad.
Mukhamediev, A.M. (1951): Comparison and origin of the fauna of Crustacea in paddy fields in the Fergana Valley. Zool. Zn. 30: 376-378.
Nayar, C.K.G. (1971): Cladocera of Rajasthan. Hydrobiol. 37: 509-519.
Scourfield, D.J. & Harding, J.P. (1966): A key to the British species of freshwater Cladocera with notes on their ecol- ogy. Freshw. Biol. Ass. Sci. Pub. 5: 55.
Smirnov, N.N. (1974): Chydoridae Fauna of the World. Fauna of USSR. Crustacea 1(2): 644 .
Thienemann, A. (1950): The transport of aquatic animals by birds. Binnengewasser 18: 156-159.
Venkataraman, K. (1983): Taxonomy and Ecology of Cladocera of southern Tamil Nadu. Ph.D thesis, Madurai Kamaraj University.
Venkataraman, K. (1988): Cladocera of Keoladeo National Park, Bharatpur, Rajasthan II. New Records 1. Moinodaphnia macleayii (King, 1853) and Bosminopsis ddtersi (Richard, 1895). J. Bombay nat Hist Soc. 85(1): 229-233.
Venkataraman, K. (1990): New records of cladocera of Keoladeo National Park, Bharatpur-III. J. Bombay nat. Hist. Soc. 87 (1): 166-168.
FOOD AND FEEDING BEHAVIOUR OF THE GREAT INDIAN BUSTARD ARDEOTIS NIGRICEPS (VIGORS) 1
Bharat Bhushan2 and Asad R. Rahmani3 (With two plates and four text-figures)
Introduction
The great Indian bustard Ardeotis nigriceps , endemic to parts of the Indian subcontinent, is an endangered avian species. It is included in Schedule I of the Wildlife (Protection) Act, 1972, and a number of sanctuaries have been established for its protection (Rahmani 1987, 1989). The present study on the food and feeding behaviour of the species forms part of a wider investigation on its ecology. Studies on food and feeding be- haviour were mainly done at Karera Bustard Sanctuary, Madhya Pradesh, but wherever neces- sary, data from other sites such as Nanaj bustard area in Solapur district, Maharashtra, and Rol- lapadu Bustard Sanctuary in Kumool district, Andhra Pradesh, are also included.
Owing to its status as a game bird, the natural history of the great Indian bustard, including its food, has been described by many workers; but there is no detailed study on its feeding behaviour and seasonal dependence on different food items. This paper deals with these aspects. The study is based largely on faecal analysis. While previous reviews of food habits analyses (Hartley 1948, Van Tyne and Berger 1959, Korschgen 1969, Lorin 1970) maintain that analysis of faecal mat- ter should be the last alternative for study, they also accept that it is the only solution where endangered species are concerned.
Study Area
The 202.21 sq. km Karera Bustard Sanctuary (25°30’ to 24°40’N, 78°5’ to 78°12’ E) (hence- forth Karera) is located in Shivpuri district, Mad-
1 Accepted February 1991.
2,3Bombay Natural History Society, Hombill House, Shaheed Bhagat Singh Road, Bombay 400 023.
3 Present address : Centre for Wildlife and Ornithology, Aligarh Muslim University, Aligarh 202 001.
hya Pradesh. The average annual rainfall recorded by us from 1983 to 1985 was 966 mm. Summers (March to June) are very hot (maximum recorded 48°C), but temperatures as low as 4° C have been recorded in winter (November to February). The terrain is gently undulating, with scattered stones and boulders.
The original vegetation of the area was clas- sified as Tropical Dry Deciduous Forest (Cham- pion and Seth 1968). There are a few hillocks, which some decades ago would have been covered with Anogeissus pendula trees, but indis- criminate cutting and lopping have eroded the hills, leaving stunted specimens. The plains have degraded into open scrub. Wherever possible, the land has been cultivated. Today the shrubs Zizyphus rotundifolia and Acacia leucophloea are the dominant natural plants, the latter scattered and the former in clumps in all the uncultivated parts of the plains.
In the scrub area, along with Zizyphus, various grasses were found but severely exploited by livestock. Except for private fields, the entire area was totally grazed. Among grasses Heteropogon contortus, Cynodon dactylon, Andropogon pumilus, Pennisetum pedicellatum, Aristida spp. and Eragrostris spp. were common.
A number of ephemeral streams run across the sanctuary and flow into the village tanks or rivers. These streams dry up by the end of Sep- tember or October. With constant soil erosion in the hills, the dried stream beds show presence of soil, gravel and rocks all along their course.
There are 33 villages within the boundary of the Sanctuary, and both human density (127 per- sons/sq. km) and the livestock population (179.5/sq. km) are high.
The bustards are found in open scrubland. During our study period, there were about 25 bustards in the Sanctuary. The breeding season at
JOURNAL, BOMBAY NATURAL HIST. SOCIETY, Vol 89
Fig. 1. Study area in the Great Indian Bustard Sanctuary, Karera.
J. Bombay nat. Hist. Soc. 89 Plate 1
Bhushan and Rahmani: Great Indian bustard
Ghana Cicer arietinum plant matter recovered from bustard faeces. Left: whole shredded and fibrous pod material. Right above: whole seeds. Right below: shredded and fibrous stalk material.
Mantid cacoons (right) after recovery from faecal matter, as compared to their shape before being eaten (left).
Plate 2
J. Bombay nat. Hist. Soc. 89
Bhushan and Rahmani: Great Indian bustard
• • *
w ^
*)•*««*«
* • « f i 4c
tMI 1 9
- A"'
ill
Coleopteran body parts recovered from faecal matter.
Orthopteran body parts recovered from faecal matter.
FOOD AND FEEDING BEHAVIOUR OF THE GREAT INDIAN BUSTARD
29
Karera is from mid March to end June. Although occasional nests were found in July and August as well, courtship display of adult males was not seen after June.
The other areas were Nanaj (for details see Ali and Rahmani 1982-84, Rahmani and Manakadan 1989) and Rollapadu (see Manakadan and Rahmani 1989).
Methods
Study plots: Seven plots where bustards were seen foraging were selected (Fig. 1). The names of these plots are based on local names. Detailed descriptions of the plots are given by Bhushan (1985) and Rahmani (1989).
Analysis of faecal samples: More than 300 faecal samples were collected at Karera for analysis to determine the bustard's varying de- pendence on seasonal food. Most of the samples were collected from roosting sites. Wet samples were preserved in 10% formalin, and dry samples preserved in separate packets for later analysis. Analysis was done on the basis of Korschgen (1969) and Lorin (1970). Faecal material was studied under a stereoscopic binocular micro- scope. Various items in the diet were identified by comparison with reference material or directly in the case of such undigested parts as chelicera (arachnida) , elytra and mandible (insects), seed (Triticum, Zizyphus ) and pod (Cicer, Arachis).
Figs. 2 and 3 show the percentage weight of various food items in the faecal material collected fortnightly. Although insects appear to be preferred by the bustards (visual observations), they are under-represented in the figures, mainly because they were digested and thus did not show relative proportions in the faecal material. On the other hand, plant material is over-represented mainly because many plant parts (i.e. seed) remain undigested and are thus easily identified and weighed. Small reptiles which are opportunis- tically eaten (visual observations) were not recorded in the faecal material. These limitations should be kept in mind while interpreting figures.
Feeding rates: The bustards were observed while foraging by the scanning method, wherein
the bird was watched continuously for a five minute unit, termed here as a ‘scan'. During a single scan, each peck by the bustard away from its lateral position was counted. The total number of pecks in a single scan was considered to be the feeding rate (or peck rate) for that particular scan (after McKee 1982). The scan count was discon- tinued when the bird went out of view.
Peck rates were studied from May 1982 to August 1984 and 1271 readings were taken, spread over different seasons and months. For various reasons, equal number of readings could not be taken every month or every season. Most readings were taken during the breeding season (summer) and in winter, when the birds were watched more regularly.
Identification: To study the peck rate of different individuals, bustards were identified as (i) Alpha male = adult territorial male; (ii) Beta male = subadult non-territorial male; (iii) Juvenile male = less than one year old, generally seen with the mother; (iv) Mother = hen with juvenile; (v) Breeding female = hen with an egg or a chick; and (vi) Non-breeding female = hen without an egg or chick.
Entomological studies: Sweeping method was used to count the insect number. Weekly sweeps were carried out in different plots. 100 sweeps each were done over a fixed stretch of an area in each plot, between 0700 and 0900 hrs. The main emphasis was on the study of grasshoppers, which constitute the major animal food of the bustard, and are comparatively large and easy to identify and count. Moreover, sweeping method is much more effective to study the population of Orthoptera and Lepidoptera than Coleoptera. The insects caught or flushed during sweeping were counted.
The following were the common Orthop- terans and Lepidopterans found in the study area: Acrida exaltata, Chloeobora sp., Chrotogonus spp., Dnopherula sp., Gastrimargus africanus , Locusta migrator ia, Pyrgomorpha sp.,
Spathosternum sp .fAcorypha spp., Catantops sp., Chorthippus sp., Eyprepoenemis sp., Hieroglyphus sp., Patanga sp., Poicilorhynchus
3
30
JOURNAL, BOMBAY NATURAL HIST SOCIETY, Vol 89
Insect V/l Zizyphus
Other crop* 1 I Miscell.
'Arachls/Trl tlcum/Sesamum
Eruca l\\l Cicer
Unident. PlantLLU Pebbles
1983
100
75 - =
50 -
25 - ± == ==
Fig. 2. Percentage of various food items in faecal samples of great Indian bustard collected in 1983.
Insect
Other crops*
V/l Zizyphus 1 1 Miscell.
144-1 Eruca N\1 Cicer
I 1 Unident. PlantDZD Pebbles
1984
lOOnnnnfflPP
Fig. 3. Percentage of various food items in faecal samples of great Indian bustard collected in 1984.
FOOD AND FEEDING BEHAVIOUR OF THE GREAT INDIAN BUSTARD
31
No. of Orthopterans
Fig. 4. Monthwise and inter-annual variation in numbers of Orthoptera in the study area.
pictus and Sphingonotus sp.; and Tarucus indie a, I nara , I theoplirastus, Eurema brigitta , Precis lemonias , Precis orithyia , Papilio demoleus , Acraea violae , Hypolimnas misippus and Belenois mesentina .
Though Coleopterans are commonly eaten by bustards (Ali and Ripley 1969, Bhushan 1985), they are mainly crepuscular or nocturnal, and are rarely caught in the sweep net. Pitfall traps (using petri- dishes filled with formalin to trap beetles) were not used in order to avoid accidental drinking of for- malin by bustards and wild mammals. Moreover, there were always chances of petri-dishes being broken by grazing livestock or by blackbuck.
Five habitat types were selected for insect studies: (i) Open scrub; dominated by Zizyphus rotundifolia (average height 50 cm) and very low tree density (<1 tree/10 ha), (ii) Usar area; bare, alkaline area, dominated by Chloris stricta grass.
(iii) Wooded scrub: dominated by Z. rotundifolia and Acacia leucophloea (tree density > 5/10 ha).
(iv) Inside plantation; totally protected 20 ha
forest plantation, dominated by various grasses (height >100 cm) and some trees (density >1 tree/10 ha), (v) Outside plantation: very similar to Open scrub, i.e. dominated by Zizyphus rotun- difolia', this category was selected to compare the counts with the adjoining Plantation plot.
Results
Population fluctuation of Orthoptera and Lepidoptera: AtKarera, the orthopteran popula- tion was very low during winter (October to February). From the middle of March, it starts rising and reaches its peak during the late summer or monsoon. Inter-annual variation was seen in population as well as in the timing of peaks and troughs (Fig. 4). These were also dependent on local factors such as precipitation and temperature.
The lepidopteran number, on the other hand, is generally low in late winter and sum- mer, but soon after the monsoon breaks, it starts increasing and reaches its peak during August and September.
32
JOURNAL, BOMBAY NATURAL HIST. SOCIETY, Vol. 89
Table 1
LITERATURE REVIEW OF THE FOOD OF THE GREAT INDIAN BUSTARD
|
Food recorded |
References |
|
Insects |
|
|
''Grasshoppers, Mylabris, Buprestris , Scarabaei..." |
Hume and Marshall 1879 |
|
"Grylli, beetles of all kinds (Cetonidae, Elateridae, Buprestidae, Carabidae), frequently the spongy nidus of the mantis, caterpillars, Julidae, Scolopendridae...." |
Elliot 1880 |
|
"Locusts, grasshoppers, beetles, crickets, mole-crickets, ants" |
Dharmakumarsinhji 1957 |
|
Green blister beetle Cantharis tenuicollis |
Ali and Ripley 1969 |
|
Beetles {Gymnopleurusl , Atactogaster ) |
Gupta 1975 |
|
Curculionidae: Platynotus, Mylabris , Stemocera nitidicollis |
Manakadan 1985 |
|
Scorpions, spiders etc. |
|
|
"Centipedes, spiders, scorpions, worms" Reptiles |
Hume and Marshall 1879, Elliot 1880, Dharmakumarsinhji 1957, Gupta 1975 |
|
"Lizards, snakes..." |
Hume and Marshall 1879, Dharmakumarsinhji 1957, Ali and Ripley 1969 |
|
Echis carinatus |
Carter 1912 |
|
Uromastix hardwiddi |
Gupta 1975 |
|
Bird eggs |
|
|
Egg of Cotumix |
Hume and Marshall 1879 |
|
Mammals |
|
|
Rats |
Hume and Marshall 1879 |
|
Mice |
Dharmakumarsinhji 1957 |
|
Plants* |
|
|
"green shoots of lemon grass Citronella" |
Dharmakumarsinhji 1957 |
|
Carissa |
Hume and Marshall 1879, Gupta 1975 |
|
Salvadora |
Dharmakumarsinhji 1957 |
|
Inanimate matter |
|
|
"stones & gravel", "pebbles" |
Hume and Marshall 1879, Elliot 1880, Dharmakumarsinhji 1957 |
|
"brass ornament" |
Hume and Marshall 1879 |
♦Plants included here are not listed in Table 2.
Food Recorded Through Faecal Analysis Seasonal Variation
Winter (October to February): Winter in Karera starts in October; Insect numbers start declining, and the monsoon crop such as groundnut A rachis hypogea , as well as Zizyphus drupes, start ripening. In the beginning of Oc- tober, bustards were mainly seen feeding on in- sects and drupes of Zizyphus. By mid-October, the insect number declines rapidly and bustards feed more and more on the ripe drupes of Zizyphus. While insects constituted 13% of the diet (by
weight) in the first fortnight of October 1983, in the second fortnight they declined to 7% (Fig. 2). On the other hand, the percentage of Zizyphus increased from 65 to 68%.
In November also, drupes of Zizyphus constitute the major food (Fig. 2). Soeha Eruca sativa , which is a dryland winter crop at Karera, is relished by bustards. So is groundnut; bustards are frequently seen in groundnut fields, foraging on exposed nuts. They do not dig up the plants. How- ever, we did not obtain groundnut in faecal samples.
FOOD AND FEEDING BEHAVIOUR OF THE GREAT INDIAN BUSTARD
33
Table 2
PLANT SPECIES IDENTIFIED IN THE BUSTARD DIET
|
Family |
Species |
Remarks |
|
CRUCIFERAE |
Eruca sativa |
Crop |
|
Brassica campestris |
Crop |
|
|
MALVACEAE |
Hibiscus abelmoschus |
Wild plant/weed |
|
TTLIACEAE |
Triumfetta rhomboides |
Wild plant |
|
LEMACEAE |
Linum usitatissimum |
Crop |
|
RHAMNACEAE |
Zizyphus rotundifolia |
Wild plant |
|
LEGUMINOSAE |
Cicer arietinum |
Crop |
|
Arachis hypogea |
Crop |
|
|
Vigna sinensis |
Crop |
|
|
CUCURBITACEAE |
Cucumis melo |
Wild plant/weed |
|
PEDALIACEAE |
Sesamum indicum |
Crop |
|
GRAMINEAE |
Sorghum vulgare |
Crop |
|
Triticum vulgare |
Crop |
In December, the percentage of Zizyphus in the diet of bustards declined to less than 50% (Fig. 2), while the percentage of Eruca sativa did not change significantly. Similarly, there was not much change in the insect constituents. Vegetal matter remained the major part of the diet as indicated by the faecal samples.
Although Zizyphus drupes are more or less exhausted by January, bustards were still able to pick up sufficient numbers to constitute up to 51% by weight in the first fortnight of 1984 (Fig. 3). In both 1983 and 1984, in the second fortnight of January the percentage of Eruca sativa increased (Figs. 2, 3).
This is further corroborated by our visual observations in the field. The insect number was very low (Fig. 4) and hence bustards were more frequently seen in the crop fields of soeha and Bengal gram Cicer arietinum . Sometimes com- plete pods of gram were found in the droppings but we rarely saw them eating the pods of soeha. Occasionally, they were seen feeding on mustard Brassica campestris and til Sesamum indie um.
In the first fortnight of February, Eruca sativa constituted 14.7 and 23.4% in 1983 and 1984 respectively, while in the second fortnight it in- creased to 33.6 and 45.7% (Figs. 2, 3). Similarly, the percentage of Bengal gram also increased in the second fortnight. In 1983, we found very little Zizyphus in the faecal samples, unlike 1984 when in both the fortnights, Zizyphus constituted 28% (Fig.
3). This could be due to a sampling error. In 1983 only five samples were analysed in each fortnight while in 1984, 12 and 16 samples were studied, which showed the diet variation more clearly. In February, as in other winter months, insects formed less than 10% of the diet of the bustard (as shown by faecal sample analysis).
Summer (March to June): In early March, the Bengal gram and soeha start ripening but the bustards are still seen in the crop fields, picking up the late flowers of soeha and unripe pods of Bengal gram. Insects start increasing in number (Fig. 4) but still formed less than 10% in the faecal samples analysed by us (Figs. 2, 3).
By the third week of March, when harvesting of early-grown wheat begins, bustards can be seen in newly harvested wheat fields, picking up the fallen grains. Soeha is also harvested by the fourth week of March, after which bustards are rarely seen in soeha fields. Breeding of bustards at Karera starts from the middle of March (Rahmani 1989). During the first fortnight of April Bengal gram is the main standing crop eaten by bustards at Karera, and constituted nearly 50% of the diet. Its percentage declined during the second fortnight (Figs. 2, 3). At the same time the percent- age of wheat increased in faecal samples as more fields are harvested. By the third week of April all wheat fields are harvested and livestock grazed on crop residue.
34
JOURNAL , BOMBAY NATURAL HIST. SOCIETY, Vol. 89
With the approach of summer and rise in temperature, small reptiles come out of hiberna- tion and the population of Sitana ponticeriana , Agama minor , Mabuya spp., Ophisops jerdonii and Calotes versicolor increases. Sitana, Ophisops and Agama are easily caught by forag- ing bustards because these small lizards generally depend on camouflage, remaining immobile when danger threatens. We often saw bustards with small lizards, but rarely saw them catching Calotes versicolor which is mainly arboreal. An injured Calotes offered to our captive -cum-free ranging juvenile bustard at Rollapadu was not eaten by it (Manakadan and Rahmani 1990). However, we suspect that an adult bustard can easily eat a Calotes because we have seen them eating snakes, sometimes larger than an adult Calotes.
During May and June there is hardly any crop at Karera and bustards depend totally on natural food (Figs. 2, 3). Insects, especially Orthopterans, reach maximum numbers, and form the main food, though this is not reflected in faecal analysis for reasons stated earlier (Fig. 4).
Monsoon (July to September): With the onset of the monsoon in end June or early July, agricultural activities begin and millet, pen- nisetum, sesamum and groundnut are grown. In- sect numbers remain high and constitute nearly 10% of the diet of the bustard (as indicated by faecal samples, though it must be higher) (Fig. 2). In August 1984 (we have no data for August 1983), insects constituted 47.31% in the first fortnight and 18.52% in the second fortnight (Fig. 3).
Bustards also feed on Cucumis melo var. momordicum. Locally known as ‘gila’, the cucumber-like Cucumis melo is locally con- sidered as a weed and grows naturally in groundnut fields. Bustards feed on the fruit (3-5 cm long and 2-3 cm broad), and presumably digest the entire fleshy matter and possibly the unripe seeds also, because only the hard seeds were found in faecal samples.
During September, insects constituted about 20% of the diet (Fig. 3). Groundnut is a new crop
which is added to the food of the bustard from this month onward till it is harvested in Novem- ber/early December. Bustards are first seen in the groundnut fields when it is sown, and later when it is ready for harvesting. As the groundnut plant is very short (20-30 cm), it is suitable for bustards and the birds are regularly seen there.
By the end of September, monsoon crops are harvested and f ields are prepared for winter crops such as wheat, soeha and Bengal gram. The insect numbers start declining. At the same time, drupes of Zizyphus start ripening and more bustards are seen picking the drupes from bushes. The cycle repeats itself.
Crop Plants Eaten by Bustards
Soeha: Soeha or taramira is a dry-land winter crop extensively grown in Karera and the whole of north-west India. We have seen it grown inSorsan, Sonkhaliya (Rajasthan), Ghatigaonand Pohri (Madhya Pradesh) bustard areas/ sanctuaries. The whole plant - inflorescence, leaves and shoots - is eaten by bustards. As long as there are flowers and fresh green leaves, bus- tards are seen in soeha fields. Some of the highest peck rates were found in the soeha fields (see Table 3).
Soeha is a marginal crop, grown only in those areas where irrigation facilities are scanty. If a field can be irrigated, farmers prefer to grow more commercially profitable crops.
Bengal gram: After soeha, Bengal gram Cicer arietinum is the most preferred crop of bustards at Karera. Cool dry climate and light well-drained soil are essential for successful cul- tivation. It can be grown with or without irriga- tion. The plant is 30-40 cm tall and is grown in narrow rows. At Karera, it is mainly grown in non-irrigated or marginally irrigated areas. Some- times it is grown with wheat.
Bustards eat young shoots, flowers and un- ripe pods of Bengal gram. The highest peck rates were observed in Bengal gram fields (Table 3). If undisturbed, they visit the same gram fields over consecutive days. We have recorded stems, leaves, seeds and pods in the faecal samples.
FOOD AND FEEDING BEHAVIOUR OF THE GREAT INDIAN BUSTARD
35
Occasionally, an entire pod with seeds within was recovered in the droppings.
Groundnut: This is one of the most impor- tant commercial crops of Karera and in almost all the bustard areas (except the Thar desert). It is grown mainly as a monsoonal crop, sown at the start of the monsoon and harvested in Oc- tober/November. As groundnut is favoured by bustards, their presence can be predicted in a particular field. Soon after the water has dried from a freshly irrigated groundnut field, bustards visit the field to pick up nuts that have been exposed by the flow of water. Similarly, they are seen for many days in a newly harvested field where some nuts are left behind. The birds move from one harvested field to another, consuming the scattered nuts. We were able to bait bustards during summer by spreading wheat and groundnuts in areas frequented by bustards. An interesting behaviour seen from the hide was that the bustards shake the nuts before eating; spoiled nuts, which possibly do not produce the charac- teristic rattling sound, were discarded.
Wheat: We never saw any bustard eating wheat Triticum vulgare from the spike in a stand- ing wheat field. They were mainly seen as soon as the wheat was harvested, picking up the fallen grains. On a few occasions we also saw them picking up fallen grains from cart-tracks after wheat-laden carts had passed by. Wheat was another food item by which we were able to attract bustards at Karera.
Incidentally, bustards at Nanaj were not at- tracted to wheat when we tried to catch them for colour banding. Similarly, at Karera, we were unable to' attract bustards to wheat bait after the onset of the monsoon. This could be due to two major reasons: (i) during monsoon bustards get sufficient food in their natural habitat, and so are not particularly attracted to wheat, and (ii) damp soil quickly spoils the grain which bustards do not eat.
Millet: Unlike wheat, bustards were found to eat millet (jowar) Sorghum vulgare from the standing crop plants. At Nanaj, many bustard droppings had millet seeds, and on a few oc-
casions we saw bustards pecking at spikes of millet.
Mustard: Although closely related to soeha, mustard Brassica campestris is not a preferred food of the bustard. Only once was a male found eating green leaves from standing plants. It is commonly grown at Karera, Pohri, Ghatigaon, Sonkhaliya and the whole of north-west India, either alone or with wheat or sesame.
Cowpea: Twice we saw a male bustard eating the beans of cowpea Vigna sinensis. How- ever, cowpea was never found in faecal samples.
Linseed Linum usitatissimum: This crop is commonly grown along with soeha and Bengal gram. Although we did not see the bustard feeding on this plant, seeds were recorded in the faeces..
Peck Rate
Feeding or peck rate (PR) per five minutes was analysed with a combination of various other parameters.
Sex-wise peck rate: The average peck rates (PR) of male and female bustards in the Open scrub area were significantly different (One-way ANOVA, F 1,1019 = 4.33, P < 0.05). Males had a higher PR than females.
Habitat- wise peck rate: (Table 3) The max- imum number of observations were taken in the Open scrub area (n = 1028, 80%). More than 80% of the natural area of the Sanctuary consists of Zizyphus rotundifolia- dominated scrubland.
Fallow fields are quickly invaded by Zizyphus and within two or three years become almost like the Open scrub area. In both Open scrub area and fallow fields, livestock grazing is allowed.
Expectedly, there was not much difference in the PR between these two similar habitats (Table 3). Some fallow fields in which livestock grazing is not allowed, become dominated by grasses. We have considered them as grassland. The PR was slightly higher in Grassland than in Open scrub area or fallow fields (Table 3). This could be due to the higher number of insects in the grassland. Unfortunately, our sample size is very small (only 12 observations). This was mainly because
36
JOURNAL, BOMBAY NATURAL HIST. SOCIETY, Vol. 89
Table 3
AVERAGE PECK RATES OF BUSTARDS IN DIFFERENT HABITATS
|
Habitat |
Mean |
SD |
No. of samples |
|
Chana (Bengal gram) |
67.40 |
37.29 |
40 |
|
Soeha |
46.48 |
28.00 |
93 |
|
Miscellaneous crops |
41.22 |
36.36 |
28 |
|
Groundnut |
15.33 |
3.77 |
6* |
|
Grassland |
13.91 |
4.75 |
12* |
|
Open Scrub |
12.09 |
10.23 |
1028 |
|
Fallow field |
10.87 |
4.34 |
56 |
|
Usar area |
3.87 |
2.89 |
8* |
♦Sample too small for statistical analysis n = 1271.
grassland patches in Karera are small and tem- porary. As soon as the grass becomes tall, it is either cut for hay or grazed.
The Usar is more or less totally bare ground with negligible ground cover. In the Usar, the peck rate was also very low (3.87, n = 8). The Usar was rarely used for foraging, hence the low sample size. We saw bustards in the Usar when they were crossing from one Open scrub area to another.
Among the crop fields, Bengal gram had the highest peck rate (Table 3). This is probably be- cause the plants are only about 30 cm high, and when ripe, the green pods are easily picked by bustards. As the plant is grown closely in narrow rows, the bird does not have to move much and from one or two spots, it can pick up a large number of pods. Moreover, we have also seen bustards plucking the green leaves of chana (which does not require much effort); hence the very high PR in chana fields.
We recorded bustards in chana fields in Oc- tober and December, though faecal analyses do not show presence of chana in these fortnights. The bustards must have fed on tender shoots that were easily digested or were not identifiable in faecal analysis.
The second highest peck counts, after chana fields, were seen in the soeha field (Table 3). Here too, the bustard does not have to move much or search for food once it is in a soeha field. We have often seen bustards eating the whole plant, starting from inflores- cence, leaves, tender branches to the green
part of the stem.
In groundnut fields, the bustard has to search for exposed pods. This could explain the PR of 15, compared to 46 and 67 in other crop fields (Table 3). Another explanation could be that groundnut pods being much bigger (and perhaps more nutritious) than chana pods and leaves and flowers of soeha, the bustard has to peck less to get the same amount of food.
Plot-wise peck rate: The plot-wise PR is shown in Table 4. In order to study the availability of natural food in different parts of the Sanctuary, we analysed the PR in the Open scrub areas of different plots (Table 4) and found no significant difference (One-way ANOVA, F 6,975 = 0.017, NS).
Peck rates of different individuals: There was no significant difference in the PR of Alpha and Beta males or Alpha and Juvenile males (Tables 5, 6). Similarly, there was no significant difference between Mother and Juvenile which foraged together in the same areas. The only sig- nificant difference in the PR was between Breed- ing hens and other individuals such as Non- Breeding hens, Mother, Alpha and Beta.
Peck rates of solitary and non-solitary birds: We compared the PR of the solitary and non-solitary bustards in the Open scrub areas (Table 7), where the bustards spent most of their time. There was no difference in PR between solitary and non-solitary hens (t = -0.29, St. Err. = 2.11, Df = 437, NS) and very little difference between solitary and non-solitary males (t = - 1.76, St. Err. = 0.67, Df = 587, P <0.1). This shows
FOOD AND FEEDING BEHAVIOUR OF THE GREAT INDIAN BUSTARD 37
food, so they peck faster before conditions be- come difficult for foraging. Maximum peck rates were observed in the evening. An additional reason for the high peck rate in the late evening could be that in the evening most of the villagers and livestock would return to villages; therefore bustards are left relatively undisturbed to forage, and could devote more time to eating (pecking). Expectedly, the peck rates drop suddenly after 1800 hrs because the birds are not able to locate their prey easily after dusk. At this time, most of the bustards roam in search of roosting spots and eat desultorily. With the approach of nightfall, risk of predation increases, so the birds spend more time looking for danger and less on forag- ing.
Minimum peck rates were seen during mid-day (Fig. 5), especially during summer when the temperature can reach 48° C. Mid- day foraging was seen more frequently in winter or during cloudy days in the monsoon.
Frequency
of pecks/ 5 minutes
Karera : 1982-84
10 11 12 13 14 15 16 17 18 19
Hours of the day
Fig. 5. Feeding intensity in the great Indian bustard at Karera.
that even when the bustard is in a flock, it feeds more or less individually.
Peck rates at different hours: Peck rates were higher in the late morning, and in the evening before roosting (Fig. 5). This could be because before roosting the birds have to eat sufficient
Fig. 6. Frequency distribution of peck rate in the great Indian bustard at Karera.
38
JOURNAL, BOMBAY NATURAL HIST. SOCIETY, Vol. 89
Table 4
PECK RATES IN THE OPEN SCRUB AREAS OF DIFFERENT PLOTS
|
Name of the Plot |
Mean |
SD |
No. of samples |
|
Turkani |
11.14 |
7.35 |
416 |
|
Hanuman-jhoora |
12.38 |
8.20 |
31 |
|
Rasori |
10.95 |
6.22 |
252 |
|
Kundpatha |
10.54 |
6.62 |
133 |
|
Karawwa |
11.31 |
4.89 |
94 |
|
Outside Plantation |
8.18 |
2.79 |
11 |
|
Baigawan |
12.57 |
7.66 |
45 |
Table 5
PECK RATES OF DIFFERENT SEXES AND DIFFERENT INDIVIDUALS IN OPEN SCRUB AREAS ONLY
|
Sex/Individuals |
Mean |
SD |
No. of samples |
|
Sex |
|||
|
Male |
11.5 |
7.05 |
589 |
|
Female |
10.6 |
6.41 |
433 |
|
Individuals |
|||
|
Alpha male |
12.11 |
7.14 |
288 |
|
Beta male |
11.89 |
7.46 |
167 |
|
Juvenile male |
9.97 |
5.01 |
109 |
|
Mother (with juvenile) |
12.44 |
7.36 |
98 |
|
Breeding hen |
6.69 |
2.71 |
59 |
Table 6
SCHEFFE’S TEST TO FIND THE SIGNIFICANCE BETWEEN VARIATIONS IN PECK RATE IN DIFFERENT SEXES AND DIFFERENT INDIVIDUALS
|
NBF |
MOF |
BF |
ALM |
BEM |
JUM |
|
|
NBF |
0.00 |
4.41 |
18.84* |
5.51 |
2.77 |
1.25 |
|
MOF |
0.00 |
27.83* |
0.18 |
0.43 |
7.19 |
|
|
BF |
0.00 |
32.95* |
26.92* |
9.42 |
||
|
ALM |
0.00 |
0.13 |
8.31 |
|||
|
BEM |
0.00 |
5.53 |
||||
|
JUM |
0.00 |
*P < 0.01
NBF = Non-breeding female, MOF = Female with chick, BF = Breeding female, ALM = Alpha male, BEM = Beta male, JUM = Juvenile male.
Table 7
PECK RATE OF SOLITARY AND NON-SOLITARY BIRDS IN THE OPEN SCRUB
|
Sex |
Mean |
SD |
No. of samples |
|
|
Solitary female |
12.78 |
8.84 |
194 |
|
|
Non-solitary female |
12.17 |
6.12 |
245 |
|
|
Solitary male |
12.30 |
8.76 |
272 |
|
|
Non-solitary male |
11.12 |
7.15 |
317 |
FOOD AND FEEDING BEHAVIOUR OF THE GREAT INDIAN BUSTARD
39
Discussion
Although the great Indian bustard is an op- portunistic feeder, its preferred food is insects, chiefly Orthopterans and Coleopterans. Numerous studies such as Parker (1929), Andrewartha and Birch (1954), Uvarov (1931), Edwards (1960), Dempster (1963) and Ali (1978, 1980) have shown seasonal fluctuations in the population of insects (grasshoppers), depending on the weather conditions, chiefly precipitation and temperature. In our study areas also, seasonal fluctuation of the grasshopper was noticed (Fig. 4), with maximum numbers during summer and/or monsoon and the minimum in winter.
The bustard shifts its food-preference seasonally depending on the more commonly available food items. The timing of the bustard’s nesting season at Karera was seen to synchronise with the increase in abundance of grasshoppers (Rahmani 1989).
Food items in the bustard diet identified by earlier workers are listed in Table 1. Insects, mainly grasshoppers and beetles, are perhaps the most important animal constituents in the diet. The habitats of the bustard (grasslands and open scrubland) are suitable for picking up in- sects during foraging. While walking the bus- tard flushes various types of insects, among which Hymenoptera, Orthoptera, Coleoptera and Lepidoptera are generally eaten. In addition to insects, other organisms such as lizards and small snakes are also opportunistically eaten. Hume and Marshall (1879) reported that a com- plete egg of quail (Coturnix) was found from the stomach of a bustard. Our tame bustard (age > 1 year) at Rollapadu in Andhra Pradesh easily pick- ed up two sandgrouse Pterocles exustus eggs from the nest (Manakadan and Rahmani 1990).
The two main advantages of foraging in a flock are a higher probability of detecting a predator by vigilance of companions, and im- proved location and chance of catching prey (Bertram 1978). Therefore, foraging rate of an individual in a flock should be more than in a solitary bird. We did not find any difference in the
peck rates between a solitary and a non-solitary female, and very little difference between a solitary and non-solitary male, indicating that even when the bustard is in a flock, it feeds independently; flushing of live prey by other members of a flock (‘beaters’) does not play a significant role in locating food. Similarly, predator detection by companions, thus leaving more time to search for food, also does not con- tribute significantly while foraging.
The peck rate of a hen that has laid an egg or is accompanied by a small chick was significantly lower than in hens which were not breeding (Tables 5,' 6). The main reason could be that a hen with an egg or a small chick is constantly looking for danger, and hence has less time for foraging. We found that adult bustards are not vulnerable to most ground predators (especially during daytime) such as wolf Cards lupus , jackal Canis aureus and fox Vulpes bengalensis. However, an egg or a chick is always in danger from these and many other smaller predators also; therefore a breeding hen has to be constantly alert .
The difference in peck rate of males and females could be due to two reasons: (1) Dif- ference in the behaviour of the two sexes - females were more timid and easily alerted than males, and therefore spend more time looking for danger than do males. (2) Sexual size dimorphism - as males are larger and almost twice the weight of females, they need more food, hence higher peck rates.
Acknowledgements
Studies on the great Indian bustard were started under the guidance of the late Dr. Salim Ali, to whom we are most grateful. The U. S. Fish and Wildlife Service funded the project, and the Ministry of Environment, Govt, of India, spon- sored the BNHS for conducting the research study. We are grateful to these organisations.
We express our sincere gratitude to J.C. Daniel, Co-investigator of the Project, for con- stant encouragement. We are grateful to the state governments of Madhya Pradesh, Maharashtra and Andhra Pradesh for cooperation during the
40
JOURNAL, BOMBAY NATURAL HIST SOCIETY, Vol. 89
study period. Dr P. Azeez of the BNHS helped us with the statistical analysis.
The first author (BB) would like to express
R EFER
Au, S. & Ripley, S.D. (1969): Handbook of the Birds of India and Pakistan, vol. 2. Oxford University Press, Bom- bay.
Ali, S. & Rahmani, A. R. (1982-84): Study of Ecology of certain Endangered Species of Wildlife and their Habitats: the Great Indian Bustard. Annual Report 2. Bombay Natural History Socipty, Bombay.
Ali, Shamshad (1978): Ecology and biology otAcrida exal- tata Walker. Ph.D. thesis, Aligarh Muslim University, Aligarh.
Au, Shamshad (1980): Seasonal variation in the population of Acrida exaltata , Walk, at Aligarh. J. Bombay nat. Hist. Soc. 81: 19-28.
Andrewartha, H.G. & Birch, L.C. (1954): The distribution and abundanceof animals. University of Chicago Press, Chicago.
Bertram, B. C. R. (1978): Living in groups: Predators and prey. In: Behavioural Ecology: An evolutionary ap- proach (eds. Krebs, J. R. and N. B. Davies). Blackwell, Oxford, pp 64-96.
Bhushan, B. (1985): The food and feeding behaviour of the Great Indian Bustard Choriotis nigriceps (Vigors). Class Aves: Otididae. M.Sc. dissertation, University of Bombay, Bombay.
Carter, J. R. (1912): Bustard in Kathiawar. J. Bombay nat Hist. Soc. 21: 1333-4.
Champion, H. G. & Seth, S. K. (1968): A revised survey of the Forest Types of India. Government of India Press, New Delhi.
Dempster, J. P. (1963): The population dynamics of grasshop- pers and locusts. Biol. Rev. 38: 490-529.
Dharmakumarsinhji, R.S. (1957): Ecological study of the Great Indian Bustard in Kathiawar peninsula. Jour. Zool. Soc. India 9: 140-152.
Edwards, R.L. (1960): Relationship between grasshopper abundance and weather conditions in Saskatchewan: 1930-58. Can. Ent. 92: 619-624.
Elliot, W. (1880): Notes on the Great Indian Bustard, with special reference to its gular pouch. Proc. Zool. Soc. Loud. Volume 486-489.
Gupta, P. D. (1975): Stomach contents of the Great Indian Bustard. J. Bombay nat. Hist. Soc. 71: 361-81.
his gratitude to his family for their patience during his absence during the study period.
ENCES
Hartley, P.H.T. (1948). The assessment of food of birds. Ibis 90 : 361-381.
Hume, A.O. & Marshall, C. H. T. (1879): Game Birds of India, Burmah and Ceylon, vol. 1. Calcutta.
Korschgen, L.J. (1969): Procedures for food-habits analysis. In: Wildlife Management Techniques, 3rd ed. (ed. R.H. Giles). The Wildlife Society, U.S.A.
Lorin, E. H. (1970): Nutrition research techniques for domes- tic and wild animals. Vol. I. An international record system and procedures for analysing samples. Utah State University, Utah, U.S.A.
Manakadan, R. (1985): The ecology of the Great Indian Bustard Choriotis nigriceps habitat. M.Sc. dissertation, Bombay University, Bombay.
Manakadan, R. & Rahmani, A. R. (1989): Rollapadu Bustard Sanctuary, with special reference to the Great Indian Bustard. J. Bombay nat. Hist. Soc. 86: 368-380.
Manakadan, R. & Rahmani, A.R (1990): Growth and development of a captive great Indian bustard Ardeotis nigriceps (Vigors) chick. Avicultural Magazine 96(3): 133-140.
McKee, J. (1982): The winter feeding of Turnstone and Purple Sandpipers in Strathclyde. Bird Study 29: 213-216.
Parker, J. P. (1929): Some effects of temperature and mois- ture upon the activities of grasshoppers and their rela- tion to grasshopper abundance and control. 4th Int. Congr. Ent. 1:322-332.
Rahmani, A.R. (1986): Status of the Great Indiau Bustard in Rajasthan. Technical Report No. 11. Bombay Natural History Society, Bombay.
Rahmani, A.R. (1987): Protection to the Great Indian Bustard. Oryx 21: 174-179.
Rahmani, A.R. (1988): The conservation of the Great Indian Bustar d Ardeotis nigriceps (Vigors) in the Karera Bus- tard Sanctuary. Biol. Conserv. 46: 135-144.
Rahmani, A.R. (1989): The Great Indian Bustard: Final Report. Bombay Natural History Society, Bombay.
Rahmani, A. R. & Manakadan, R. (1989): Return of the Great Indian Bustard in Maharashtra. Jour. Ecol. Soc. 2: 19-29.
Uvarov, B. P. (1931): Insects and Climate. Tr. Ent. Soc. London 79: 1-247.
Van Tyne, J. & Berger, A. J. (1959): Fundamentals of- Ornithology. John Wiley and Sons, New York.
ON THE IDENTITY AND NOMENCLATURE OF CERTAIN INDIAN IXORA (RUBIACEAE)1
D.B. Deband R.C. Rout2
The revision of the genus Ixora L. (Rubiaceae) by Bremekamp (1937a) has been criticised by Comer (1941) in his study of the Malayan Ixora. Comer observed "the size and hairiness of corolla and even the shape of the petals I find to be most variable and, for the majority of the Malayan species, by no means specific. From the most recent works on the genus (Craib 1934, Bremekamp loc.cit.), the impression is gained that many new species can be blocked out from the old and that minute differences in hairiness, leaf size or shape and length of corolla tube suffice to define them. But I am certain that this view is mistaken, and if followed, must lead to the making of so many species that the clas- sification of the genus will become impossible. Such splitting, based on relatively few herbarium specimens, merely obscures the issue which is to have names for the well defined groups of in- dividuals".
Ixora chinensis Lam., 7. coccinea L .,7. javanica (Bl.) DC., 7. grandifolia Zoll. & Mor., I. nigricans Wt. & Am. etc. treated therein by Corner (loc.cit.) are very common and widespread in India. On the other hand, field studies conducted by the senior author (D.B.D.) of this note during the last three decades and that of the other author (R.C.R.) for the last four years support the observations of Corner. Again, specimens available in herbaria after the study by Bremekamp (loc. cit.) and Comer (loc.cit.) tend to bridge the gap in knowledge on variability of some species. Husain and Paul (1989) did not examine many specimens in CAL and K as there is no indication of their study on the herbarium specimens to which the authors of this note had access.
Accepted January 1991.
“Botanical Survey of India, P.O. Botanic Garden, Howrah 711 103.
In the light of these facts we are in a position to comment on the status of several species there- by reducing them to synonyms. Those reduced to synonyms do not deserve infraspecific status.
1. Ixora goalparensis Bremek. (1938 : 336) was postulated on the basis of U.N. Kanjilal 5758, collected from Goalpara district \)f Assam. This was distinguished from I. subsessilis in much longer calyx teeth. 7. longibracteata Bremek. (1959: 371) was distinguished from I. goalparen- sis Bremek. in longer bracts, bracteoles and calyx teeth. A study of the protologue and type specimens along with other specimens of both the species suggests that I. goalparensis and 7. lon- gibracteata differ neither from each other nor from 7. subsessilis in any respect. The latter is variable in the length of bracts, bracteoles, calyx teeth and corolla tube, thereby covering up the distinctions noted by Bremekamp (loc.cit.). Hence 7. goalparensis and 7. longibracteata are treated as synonyms as follows. They do not even deserve any infraspecific status.
Ixora subsessilis Wall, ex G. Don, Gen. Syst. 3: 572. 1834 (Lectotype : Jaintiapore (Jowai), May 1826, F. De Siva s.ti. ex Wall. Cat. 6139 A CAL!). Hook, f., FI. Brit. Ind. 3: 139. 1880; Husain & Paul in Journ. Econ. Taxon. Bot. Addl.Ser. 6:119. 1989.
7. oxyphylla Wall, ex G. Don, Gen. Syst. 3: 572. 1834 (Type: Wall Cat. 6159 & 6159 ACAL!)
7. goalparensis Bremek. in Journ. Bot. 76: 336. 1938; Husain & Paul in Journ. Econ. Taxon. Bot. Addl. Ser. 6: 117. 1989 (Type : Assam, Goal- para dist., Guma Reserve, 22. 5. 1916, U.N. Kan- jilal 5758 holo. DD!, photo and iso. CAL!), Syn. Nov.
7. longibracteata Bremek. in Ind. For. 85(7): 371. 1959; Hussain & Paul in Journ. Econ. Taxon. Bot. Addl. Ser. 6: 119. 1989 (Type: Bangladesh, Chittagong. Hill Tracts, Tintilla, 16.5.1939, T. V. Dent 14, holo. DD!, photo CAL! Syn. Nov.
42
JOURNAL, BOMBAY NATURAL HIST SOCIETY, Vol. 89
Husain and Paul (1989 : 119) while select- ing Wall. Cat. 6139 A CAL as the lectotype misinterpreted Wall. Cat. entry and considered that this specimen was collected by De Silva & W. Gomez. This is not correct. The specimen con- cerned in CAL was collected by F. De Silva in May 1826 from Sylhet mountain which is in fact Jowai, now in Meghalaya.
2. Ixora andamanensis Bremek. (1937b: 260) was described on the basis of C.E. Parkinson 140. It was probably wrongly compared with I. lacei and I. ackr ingae. Rather it agrees with I. barbata Roxb. wherein corolla tube varies in length from 20 to 35 mm and the throat is bearded to glabrescent. In/, andamanensis , corolla tube is 15 to 17 mm and the throat is glabrous or glabres- cent. Thus it gets merged with I. barbata and does not deserve any infra-specific status.
I. katchalensis Husain & Paul (1984 : 153 - 156) was postulated on the basis of four gather- ings : P. Chakraborty 1134 & 5305, N. Bhargava 5031 and N. P. Balakrishnan 5325. It was distin- guished from I. barbata Roxb. in "distinctly longer petiole, more lateral nerves, different in- florescence pattern, number of flowers per head and structure of stigma". A study of protologue along with the types (holo. and iso.) suggests that, I. katchalensis does not differ from I. barbata in any respect and agrees with it. The stigma described as "slightly cleft in the middle by 1/3 from above" is found to be bifurcated to two linear stigmatic arms. Hence, I. katchalensis does not stand as a distinct species, or does not deserve any infraspecific status.
I. andamanensis and I. katchalensis are reduced here to synonyms as follows.
Ixora barbata Roxb. (Hort. Beng. 10. 1814 non. nud. &) ex Smith in Rees, Cycl. 19. no. 6. 1811; Roxb. FI. Ind. 1: 394. 1820 (Type: Andaman Islands. Cultivated at H.B.C. (CAL), Roxburgh s.n. holo. K!, photo CAL!); Hook.f., FI. Brit. Ind. 3: 148. 1880; Hussain & Paul in Journ. Econ. Taxon. Bot. Addl. Ser. 6: 146. 1989.
I. andamanensis Bremek. in Journ. Bot. 75: 260. 1937; Husain & Paul in Journ. Econ. Taxon. Bot. Addl. Ser. 6: 94. 1989 (Type: Andaman Is-
land, Havelock, 1914, C.E. Parkinson 140 holo. DD!, photo CAL!), Syn. Nov.
I. katchalensis Husain & Paul in Blumea 30: 153-156. 1984 & Journ. Econ. Taxon. Bot. Addl. Ser. 6: 153. 1989 (Type: India, Andaman & Nicobar Island, North Nicobars, Katchal Is., sea level, 22 Apr. 1974, P. Chakraborty 1134 PBL), Syn. Nov.
3. Husain & Paul (1986, 1989) proposed I. beddomei, I. manantoddi and I. mercaraica , on the basis of a single gathering each, collected from the same locality in Wynaad district of Kerala, and the adjacent locality Mercara of Coorg, now in Karnataka. They distinguished these species from I. lawsonii , which was also originally collected from these localities.
Husain and Paul do not appear to have ex- amined the type specimen of I. lawsonii located at K and MH as no sign of exclamation is added after the name of herbaria where they are extant nor they seem to have studied the original descrip- tion properly. In describing this species they say "habit unknown... stipules not seen... anthers and filaments not seen". In ‘distribution and ecology’ they say "populations are usually encountered in ghats of Wynaad, Manantodde and Coorg", but no specimens has been cited. If populations are en- countered then the habit and stipules cannot remain unknown. The notes on the species on p. 133 are ad verbatim copied from Gamble’s original publication.
Gamble stated inter alia "Arbor vel frutex
elatus ... Folia basi rotundata, juniore aliquan-
do attenuata; nervi lateralis 10-16; stipules
ovatae, apiculo dorsali longi subulata stamina
recurva, filamentis brevibus, antheris linearibus". This appears to be a rare species. In spite of the area having been thoroughly explored, only one collection has been made after the original collec- tion.
These species differ from/, lawsonii only in slight pubescence of the leaf, a character that may arise due to ecological variation. Moreover for this slight variation in hairiness only, these taxa cannot be distinguished even in infraspecific status.
IDENTITY AND NOMENCLATURE OF CERTAIN INDIAN IXORA
43
Ixora beddomei is distinguished from/, law- sonii in foliis basi acutis, calycum tubis pubescen- tibus et corollae tubis 17-17.5 mm longis. In I. lawsonii , leaf base varies from acute to obtuse or rounded. The calyx is pubescent in the same specimen (Wight s.n. K!) and corolla tube is 7-20 mm long, smaller size is evident in bud stage. Thus I. beddomei cannot stand as a distinct species.
Type specimens of I. mercaraica ( Hohenacker 439 a) is too poor to be the basis of a species. It is an incomplete specimen with only a pair of leaves at the base of peduncle, which is normally variable from those of the other leaves in form, length of the petiole and hairiness. The peduncle may be pubescent when the remaining part of the stem is glabrous. In/, lawsonii , petiole varies from 2 to 10 mm. -Calyx tube pubescent outside, calyx teeth 3-7.5 x 0.5-1 .0 mm, pubescent outside, glabrous within. Moreover, a species should not be distinguished on the basis of slight differences in quantitative characters like length of petiole, bracteoles and calyx lobes. Thus /. mercaraica also does not stand.
I. manantoddi agrees with I. lawsonii in all respects, except hairiness of young branches and leaves beneath. So it does not stand as a distinct species.
I. beddomei , I. mercaraica and I. manantod- di are reduced to synonyms as follows:
Ixora lawsonii Gamble in Kew Bull. 1920: 247. 1920; Husain & Paul in Joum. Econ. Taxon. Bot. Addl. Ser. 6: 131. 1989 (Type: Kerala, Wynaad dist., Manantodde, ± 1000 m, Jan. 1884, MA. Lawson 43 lecto. K!, photo CAL!, Isolecto. MH!).
I. beddomei Husain & Paul in Candollea 41 (1): 87. 1986 & Joum. Econ. Taxon. Bot. Addl. Ser. 6: 124. 1989 (Type: Kerala, Wynaad, 1885, R.H. Beddome 3909 holo. BM!, photo CAL!, iso. K), Syn. Nov.
I. mercaraica Husain & Paul in Candollea 41(1): 88. 1986 & Journ. Econ. Taxon. Bot. Addl. Ser. 6: 135. 1989 (Type: Karnataka, Mercara, 1&41, Hohenacker 439 holo. BM!, photo CAL!, iso. K), Syn. Nov.
I. manantoddi Husain & Paul in PI. Syst. Evol. (MSS) & in Journ. Econ. Taxon. Bot. Addl. Ser. 6: 133. 1989 (Type: Kerala, Wynaad, Manan- todde, R.H. Beddome 3908, holo. BM! photo CAL!, iso. K), Syn. Nov.
4. Ixora capituliflora Bremek. (1937 b) was based on C.E. Parkinson 1198 and/.//. Lace 2818 from Andaman Islands. The author distinguished this species from three distantly related species:/. merguensis Hook.f. in calyx lobes longer, glabrous, corolla not bearded; from I. korthal- siana Kurz in corolla not bearded and from /. kurziana (Teysm. & Binn.) Kurz in leaves smaller, inflorescence subcapitate, calyx lobes longer. On examination of types and other collections from Andaman Islands, it is observed that this species is not distinguishable from I. finlaysoniana Wall, ex G. Don. The leaves of I. finlaysoniana are narrowly obovate or oblanceolate, obtuse or mucronulate at apex, atenuate at base, whereas in Andaman specimens (/. capituliflora ), it is vari- able from elliptic to -oblong, -obovate or -lanceo- late. The cyme is also variable, subcapitate to distinctly branched corymbose one. So, /. capituliflora Bremek. does not stand as a distinct species and is reduced to a synonym as follows. It does not even deserve any infraspecific status.
Ixora finlaysoniana Wall, ex G. Don, Gen. Syst. 3: 572. 1834 (Type: East India, Finlay son s.n. ex Wall. Cat. 6166 K-WH microfiche CAL!).
I. capituliflora Bremek. in Joum. Bot. 75: 297. 1937; Husain & Paul in Joum. Econ. Taxon. Bot. Addl. Ser. 6: 124. 1989 (Type: Andaman Islands, 1916, C.E. Parkinson 1198, holo. & iso. DD!, photo & iso. CAL!), Syn. Nov.
5. Ixora roxburghii Balakr. (1981 : 232) was proposed as a substitute name for I. villosa Roxb. (1814 & 1820) as the latter was a later homonym of that of Poiret (1813), which represents a synonym of Pavetta villosa Vahl. The name I. roxburghii Balakr. was accepted by Husain and Paul (1989 : 102). However, they overlooked that /. roxburghii Balakr. is also a later homonym of that of O. Kuntze (1891), which represents a synonym of Pavetta tomentosa Roxb. ex Smith. So, this name is rejected as per Art. 64 of Interna-
44
JOURNAL, BOMBAY NATURAL HIST SOCIETY, Vol. 89
tional Code of Botanical Nomenclature. Conse- quently a substitute name is proposed as follows: Ixora balakrishnii Deb et Rout, nom. nov. Ixora villosa Roxb. (Hort. Beng. 10. 1814, nom. mid. &) FI. Ind. 1: 382. 1820, non Poir. 1813
Refer
Balakrisiinan, N.P. (1981): Flora of J&wai, Vol. I. Botanical Survey of India, Howrah.
Bremekamp, C.E.B. (1937a): The Malayan species of the genus Ixora (Rub.). Bull. lard. Bott. Bull Ser. 3, 14: 197-367.
Bremekamp, C.E.B. (1937b): Th & Ixora species of Burma and the Andaman Islands. Journ. Bot. 75: 108-111, 169- 175, 260- 266, 295-298, 318-326.
Bremekamp, C.E.B. (1938): Ixora species of Burma and the Andaman Islands - Additions and Emendations. Journ. Bot. 76: 330- 336.
Bremekamp, C.E.B. (1959): New Ixora species from Bengal, Burma and Nicobar Islands. Ind. For. 85 (7): 371-375.
Corner, E.J.H. (1941): Notes on the Systematy and Distribu- tion of Malayan Phanerogams IV: Ixora. Garden’s Bull. Str. settlements 11 (3): 177-235.
Craib, W.G. (1934): Florae Siamensis Enumeratio. Vol. 2.
Don, G. (1834): A. General System of Gardening and Botany,
(Type: Sylhet, Roxburgh s.n. ex Wall. Cat. 6137 A, CAL!, K); Hook.f, FI. Brit. Ind. 3: 144. 1880.
I. roxburghii Balakr. FI. Jowai 1: 232. 1981, non O. Kuntze 1891; Husain & Paul in Journ. Econ. Taxon. Bot. Addl. Ser. 6: 102. 1989.
ENCES
Vol. 3. London.
Gamble, J.S. (1920): In: Decades Kewenses (XXXVIII): Plantarum novarum Herbario Horti Regii Conser- vatarum. Decas XCIX. KewBull. 1920: 247.
Husain, T. & Paul, S.R. (1984): A new species of Ixora (Rubiaceae) from the Andaman and Nicobar Islands. Blumea 30: 153-156.
Husain, T. & Paul, S.R. ( 1986): Two new species of Ixora L. (Rubiaceae) from India. Candollea 41 (1): 87-93. Husain, T. & Paul, S.R. (1989): Taxonomic Studies on Indian species of genus Ixora L. (Rubiaceae). Journ. Econ. Taxon. Bot. Addl. Ser. 6: 1-205.
Kuntze, O. (1891): Revisio Generum Plantarum. Vol. I. Paris. Poiret, J.L.M. (1813): In: Lamarck, Encyclopedic Methodi- que suppl. 3: 208. Paris.
Roxburgh, W. (1814): Hortus Bengalensis.
Roxburgh, W. (1820): Flora India, vol. I. Serampur.
THE LAND TORTOISE IN NEPAL : A REVIEW 1 J. Frazier2
(With a plate and a text-figure)
Little is known of land tortoises (family Testudinidae) in the Indian subcontinent, and the information available from Nepal is remarkably muddled. Three very different species have been reported, but there are numerous unsupported statements and several claims of misidentification. Many authors have simply repeated (perhaps in a modified form) the statements of earlier publications without critically examining specimens or information. The present note reviews published and other information on land tortoises in Nepal, concluding that the only species definitely recorded in the country is Indotestudo elongata.
Introduction
Three species of land tortoise (Testudinidae) have been claimed to occur in Nepal: Geochelone elegans (Schoepf), Testudo horsjieldii Gray and Indotestudo elongata (Blyth). The first named is found from north-eastern Rajasthan (western India) south to Sri Lanka (Iverson 1986 : 140; Frazier in prep.); hence, a Nepalese record would represent a major range extension.
Testudo horsjieldii is known to occur from the Caspian and Aral areas of West Central Asia (Shammakov 1981 : pi. 3; Yatyayev 1985 : pi. 4), eastward to Baluchistan and Afghanistan (Smith 1931 : 146, Hora 1948 : 296, Iverson 1986 : 172) and even to Xinjiang province of western China (Zhao 1973). Auffenberg (1974 : 195) suggested that T. horsjieldii occurs in the environs of Dehra Dun, Uttar Pradesh, India, west of the western border of Nepal, but no evidence has ever been presented to support this claim. It is, therefore, remarkable that the Nepalese ‘record’ (see below) is the easternmost for this species.
The third species recorded from Nepal, In- dotestudo elongata , is known to occur from Indo- china westward to India (Smith 1931, Hoogmoed and Crumly 1984 : fig. 3, Iverson 1986 : 156), and as far west as Corbett National Park, Uttar Pradesh, India (Ross and Crumly 1983). This geographic range stretches across the east-west extent of the southern extreme of Nepal (Fig. 1).
1 Accepted October 1990.
2 Centro de Investigacion Y de Estudios Avanzados, Apartado Postal 73, ‘Cordemex’, Merida, Yucatan, Mexico, C.P. 97310.
Zoogeographically, G. elegans is typical of the central Indian and Deccan areas of the Indian subcontinent, I horsjieldii is part of the Palearctic fauna, and I. elongata is characteristic of the Indo-Chinese subregion of the Oriental Region (Smith 1931 : 16, Hora 1948 : 296, Jayaram 1949 : 397, 1974 : 545-546). Species from all of these zoogeographic regions, subregions or areas are known to occur in the herpetofauna of Nepal (Swan and Leviton 1962, Waltner 1973a, b, c, d); hence, a priori any, or all, of the three above- named species of land tortoise could occur in Nepal.
The confusion stems mainly from the (mis)identification of a drawing donated by B.H. Hodgson to the British Museum (Natural History) (BM[NHj) in the mid-19th century. There is a long history of problems regarding drawings of herpetological specimens from this region (see Webb 1980), and the Hodgson tortoise drawing is remarkable in this respect.
Discussion
Geochelone elegans (Schoepf)
There is only one record of this tortoise from Nepal. Laurie (1978 : 41) stated that "Reptiles and amphibians recorded in Chitawan include the marsh crocodile or mugger, the gharial, two species of monitor lizards, the starred tortoise and several species of lizards, snakes and frogs." The common name ‘starred tortoise’ is the most usual English name for G. elegans (e.g. Gunther 1864:4, Daniel 1983 : 30), and it is not regularly used for any other species in Asia.
4
46
JOURNAL , BOMBAY NATURAL HIST. SOCIETY, Vol. 89
Fig. 1. The northern area of the Indian subcontinent, showing locality records for Indotestudo elongata (circles 1-17) at the western limit of its range, and also locality records of Testudo horsfieldii (triangles A-H) at the eastern limit of its range. See Appendices 1, 2 for details of each locality record. Dotted lines show political boundaries; only major river systems are indicated.
In the absence of any other supporting evidence, this record must be rejected as an error (indeed, the appendix of reptiles in Chita wan in Laurie’s (1978) thesis lists Testudo elongata). No further mention of G. elegans will be made in the present discussion of Nepalese tortoises.
Testudo horsfieldii Gray
The first species of land tortoise reported from Nepal was T. horsfieldii. Gunther (1861 : 214) listed and described 41 species of reptiles and amphibians collected (either as actual
specimens or as drawings) by B.H. Hodgson. The first species in Gunther’s list was Testudo horsfieldii, for which he simply stated "one coloured drawing," indicating that he had not examined a biological specimen. In fact, this ‘specimen’ (Plate 1), commissioned by Hodgson, is the root of nearly all misunderstanding regard- ing land tortoises in Nepal; it is, therefore, dis- cussed in detail below.
It is noteworthy that three years later, Gun- ther (1864 : 8), evidently less confident about the identity of this drawing, wrote "A drawing made
THE LAND TORTOISE IN NEPAL : A REVIEW
47
from a Nepalese specimen, and presented by B.H. Hodgson to the British Museum, appears to rep- resent this tortoise ( T. horsfieldii ); if the deter- mination be correct, this species (T. horsfieldii ) would extend to Nepal" (italics added). For some reason, Gunther was no longer simply stating that T. horsfieldii occurred in Nepal, but instead he qualified his' listing of this species by suggesting that - on the basis of a drawing - it may extend into Nepal. He pointed out his unfamiliarity with T. horsfieldii , stating that he had seen only one specimen (1864 : 8); clearly, he was not certain of the identity of the species represented by the drawing.
Smith (1931 : 146), in his classic review of the chelonians of British India, made no mention of T. horsfieldii in Nepal, and he did not even cite Gunther’s papers (1861, 1864) under the discus- sion of this species. However, he stated (p. 143) that Hodgson’s drawing was of Testudo elongata , an earlier name for Indotestudo elongata. The fact that Smith (1931) included no mention of the previous (mis)identifications made by Gunther (1861: 218, 1864:8) left his re-identification as a source of confusion. One might sunnise that in deference to the grand old guru of herpetology, Albert Gunther, Smith chose not to mention his elder’s error.
Whatever his reasons, Smith’s (1931) omis- sion of any mention of a misidentification or his re-identification was seized upon as evidence that the identity of the land tortoise in Nepal was iii a state of confusion. Swan and Leviton (1962 : 110) flatly rejected Gunther’s identification by listing " Testudo horsfieldii Gunther, 1861, p. 214 (Nepal); 1864, p. 7 (Nepal)" as a synonym for " Testudo (= Indotestudo) elongata". They went on to claim that "It is uncertain whether Smith con- sidered Hodgson’s drawing (which Gunther described as T. horsfieldi , supra cit.) to be T. elongata or whether Smith’s reference is in error.
This treatment by Swan and Leviton (1962) is itself not without shortcomings. They failed to take into account that Gunther’s second listing (1864: page 8' which deals with Nepal, not page 7 as they indicated) was a tentative identification,
clearly including simple and obvious reserva- tions. Thus, to list ‘T horsfieldi Gunther 1864 : 8’ as a synonym of/, elongata is an oversimplifica- tion and misrepresentation of what Gunther had in fact written. In addition, Smith’s (1931 : 143) meaning could not be clearer in his identification of the Hodgson drawing, and there is no reason to claim that he was confused. It is remarkable that, in spite of these identifications, re-identifications and criticisms of earlier authors, there is no evidence given by Swan and Leviton (1962) that they ever examined either a specimen of a land tortoise from Nepal or the root cause of all the confusion and discussion - the drawing presented by Hodgson to the BM (NH)! (Levitson, pers. comm, stated that they had not seen the drawing.)
In his summary of the reptiles of the Himalaya, Waltner (1973a, b) made no mention of T. horsfieldii. He did list I. elongata , evidently following Swan and Leviton (1962). Auffenberg (1974 : 195) suggested that T. horsfieldii occurs at Dehra Dun, to the west of Nepal, and "will eventually be found to inhabit much of the foothill area of the Himalayan mountains." However, he now believes {in lift. Dec. 1987) that the species involved is Indotestudo elongata , not T. horsfiel- dii.
Jayaram (1974 : 548) stated that "Testudo horsfieldi is widely distributed from the Caspian and Aral Seas to the north-western comer of India". There is absolutely no evidence that T. horsfieldi has ever been recorded in the territory known as ‘India.’ This error was apparently caused by Jayaram having lifted Smith’s (1931 : 146) description of the range of this species, which specified ".... to the north-western corner of British India." Omitting the word ‘British’ completely changes the area from the British India of the days of the Empire (which included Baluchistan, known to harbour T. horsfieldi ) to modern India (which has never included Baluchistan).
The most recent mention of T. horsfieldii in Nepal is that of Majupuria (1981-82 : 152); he stated that "Testudo horsifieldi (sic) is represented in Hodgson’s collection." No reference or sup-
48
JOURNAL , BOMBAY NATURAL HIST. SOCIETY, Vol. 89
porting evidence was given, but this claim was evidently based on Gunther (1861 : 218). Majupuria (1981-82 : 174) stated that T. horsfiel- dii was "Reported by Prater, 1928'° in Nepal, but Prater’s book of Indian animals, first published in 1928 and now in its fourth edition, although an invaluable source book on mammals of the Indian subcontinent, does not deal with tortoises or other reptiles. Hence, Majupuria’s (1981-1982) state- ments about I horsfieldii occurring in Nepal are without support.
Indotestudo elongata (Blyth)
The first mention of this species in Nepal is that of Smith (1931 : 143): "Hodgson obtained a specimen in the Saul forests of Nepal, and has left a fine coloured sketch of it in his collections of draw- ings." The mention of Saul forests implies that Smith had examined the drawing in question, for the words ‘Land tortoise of Saul Forest’ are pencilled on the bottom left, and this information has not appeared in previous - or subsequent - publications. In addition, the evaluation of the drawing as "fine" is further support for his having examined it because it is a realistic depiction (Fig. 2).
The next mention of 7. elongata is that of Swan and Leviton (1962 : 110). As stated above, these authors rejected Gunther’s (1861 : 218, 1864 : 8) accounts of T. horsfieldii and doubted the certainty of Smith’s (1931 : 143) identification of T. elongata.
Yet, it was not explained how they ar- rived at their own identification, with no men- tion of having examined any relevant material. Nevertheless, they (1962 : 110) stated that 7. elongata is found in central Nepal, presumed (p. 107 footnote 2) on the premise that Hodgson’s material came mainly from the area around Kathmandu. More remarkable, they listed (Table 1, p. 138) this tortoise as known to occur in "Sikkim-Dar- jeeling"; no support for this claim was given, and none is known (see below). Swan and Leviton (1962 : 138) also predicted that 7. elongata occurs in eastern Nepal, and they concluded that the species could be charac-
terisedashavinganlndo-Chinesedistributionbut extendinginto the eastern Himalaya.
Waltner (1973b : 29), in his review of Himalayan reptiles, listed Testudo (= Indotestudo) elongata as recorded from "Sikkim, Darjeeling, Teak forests of Nepal ...." and from 0 to 9,000 feet (0 to 2,740 m) in altitude. The distributional data appear to be based on Swan and Leviton (1962), but the occurrence of this tortoise in teak forests or at any altitude in Nepal is unsupported by any published information. Waltner (in litt. 20 Oct. 1987) stated that he had no firsthand experience with chelonians in this region. Furthermore, teak Tectona grandis does not naturally occur as far north as Nepal (Champion and Seth 1968, Stain- ton 1972).
It is important to point out that there is no evidence that Indotestudo elongata occurs in Sik- kim and Darjeeling (c.f. Swan and Leviton 1962, Waltner 1973b : 29), an error which has evidently arisen from the inappropriate use by Swan and Leviton (1962 : Table 1) of ‘Sikkim-Darjeeling’ to refer to Sikkim and all of North Bengal (includ- ing Darjeeling, Jalpaiguri and Koch Bihar dis- tricts). There are several records of 7. elongata from Jalpaiguri dist. (Smith 1931 : 96, Das 1988 : 21, 22), but none are known from Darjeeling or Koch Bihar districts or from Sikkim.
The name Testudo elongata appears in an appendix of the reptiles of Chitawan Park, Nepal, in Laurie’s thesis (1978) on the rhinoceros Rhinoceros unicornis. However, this was ap- parently based on previous publications, not original data; and, as stated above, there is con- fusion in the text of the thesis as to which tortoise was being referred to.
Majupuria (1981-82 : 152, 174) stated that Testudo (= Indotestudo) elongata is recorded from Central Nepal, in Bagmati and Narayani zones. Although this is the first attempt to provide specific locality data for land tortoises in Nepal, it is not stated on what information these claims were based. A photograph published by Majupuria (1981-82) clearly shows three 7. elon- gata together, but no indication of where these tortoises were collected is given; the caption
THE LAND TORTOISE IN NEPAL : A REVIEW
49
simply says "Land tortoises are distributed in Central Nepal."
Clearly, it has been common for authors to reiterate the Nepalese ‘record’ of Smith (1931 : 143). After it was repeated by Biswas et al. (1978), Ross and Crumley (1983) referred to it. However, as the last named authors pointed out, there is no exact locality for this ‘record.’
Of the half dozen authors who have written about land tortoises in Nepal, none (with the pos- sible exception of Majupuria) appears to have examined a specimen from that country; and only Gunther and Smith had obviously examined the drawing donated by Hodgson to the BM (NH).
Hodgson’s drawing (Plate 1)
Smith (1931 : 5) gave a brief biographical account of Brian (or Bryan) Houghton Hodgson, and other details are presented by Archer (1962 : 11-12, 80) and Sawyer (1971 : 140). In short, Hodgson commissioned collections and drawings of local animals while based in Kathmandu from 1820 until 1844. A folio of 29 drawings bound into one volume is in the library of the BM (NH).
Only one Chelonian is depicted in this col- lection (Plate 9), and the coloured drawing of the land tortoise clearly illustrates several important features: nose and sides of the head are pink; head scales posterior to the frontal are relatively small in size; five claws are on each of the fore feet; the carapace is conspicuously more elongate than wide; there are conspicuous black, irregular blotches on each scute of the carapace; no spurs are evident in the area of the thighs (although they might not be seen from the angle shown); and a spur is conspicuous at the end of the tail. With the exception of black blotches on the carapace (which can occur in both species), all of these features are consistent with - if not diagnostic of -Indotestudo elongata , and inconsistent with Tes- tudo horsfieldii.
At the bottom of the drawing/ are several annotations. ‘Land Tortoise of Saul Forest, nat. size.’ is pencilled in at the far left (‘Saul,’ or more commonly ‘Sal,’ is the tree Shorea robusta). At the centre, in what appears to be different hand-
writing is ‘? Testudo indica.’ That name, as well as the question mark, has a line through it, and ‘Horsfieldii’ is pencilled at the right of ‘indica’; this appears to be in a third hand.
Directly under the pencilled species names are three lines of North Indian script: a pencilled line and below, in another hand, two inked lines. The pencilled line (with apparently four charac- ters) is indecipherable. The literal translation of the two inked, Hindi-like, lines is ‘R.V. Motiya Voli’ and ‘Kachhuwa.’ R.V. Motiya Voli appears to be the name of a person. The last line, Kach- huwa, means turtle.
At the far right of the base of the drawing, in pencil, are two more lines of Hindi-like script; these appear to be in two additional hands. The upper line, of three characters, may be an ab- breviated signature of R.V. Motiya Voli. However, it could also translate to ‘Sugali’ (or ‘Sungali’) the significance of which is unknown. In the bottom right comer is ‘A.V. Lapcha’ (or ‘Zapcha’); this is evidently a person’s name - probably a Nepali (although Archer (1962 : 11) stated that Hodgson hired Indian assistants). Which, if any, of these names apply to the artist or collector is unknown.
In summary, it appears as if at least five different people have annotated this drawing: three in English and between two (in the case that one person signed in full and then with initials) and four in Hindi, or another north Indian script. Some of the annotations are clearly attempts to identify the drawing, others may indicate who was involved in its execution. The comments relevant to habitat and size are likely to have been written soon after the drawing was given to Hodgson, and the north Indian script was almost certainly added while the drawing was in ‘British India’.
On the back side of the drawing, in pencil, is the list of measurements (apparently in inches and fractions of inches, except for one value in pounds), as shown in Table 1.
Records in the India Office Library (Archer 1962 : 12) and the BM (NH) library, archives and publications show that a number of well known herpetologists had corresponded with B.H. Hodgson and examined material donated by him,
50
JOURNAL, BOMBAY NATURAL HIST. SOCIETY, Vol. 89
Table 1
MEASUREMENTS INDICATED ON HOGDSON’S DRAWING
|
Length of shell |
0.11.0 |
|
Width |
7 1/2 |
|
Height |
-5- |
|
Length of head |
-2 3/4 |
|
Width of head |
-15/8 |
|
Length of leg, fore, as fully exeserted (sic) |
4 1/4 |
|
Length of leg, hind, as fully exeserted (sic) |
4 1/2 |
|
Weight |
6 lbs |
|
Tail to vent |
1 5/8 |
|
Tail to dorsal shell |
2 1/4 |
|
including T.E. Cantor, J.E. Gray, A.C.L.G. Gun- ther, Dr Hooker, Dr T. Horsfield, and M.A. Smith. |
However, neither Andrew Stimson (of the Her- petology Section) nor the Librarian of the BM (NH) could identify any of the handwriting on the tortoise drawing; it did not match handwriting samples of either Gunther or Smith.
Archer (1962 : 80) stated that light pencilled inscriptions on some of Hodgson’s drawings deposited in India House (India Office Library) were in Nagari. On the basis of this she suggested that the draughtsmen were Hindus from Bihar or United Provinces (now Uttar Pradesh).
It is probably of little significance that the Hodgson drawing was originally named ‘? Tes- tudo indica ,’ for very little was known of chelonians in the Indian region during the last century, and there was tremendous nomenclatural confusion. That Boulenger (1889 : 172) regarded T. indica as an extinct species from Mauritius, and considered the name as a synonym for one of the Galapagos tortoises (‘ Testudo nigrita ’ =
Geochelone elephantopus) (Boulenger 1889 : 169) shows how confused the species names were (see also Theobald 1870 : 674 for a criticism of the use of the species indica for Indian species of tortoises). There is no reason to further confuse the question of which tortoise is in Nepal by considering these island species.
Of more importance is: who re-identified the drawing as T. horsfieldii ? However, it may never be possible to determine who annotated this draw- ing.
Assuming that the values on the back of the
drawing are reliable measurements of the specimen depicted on the front, its identity is even more clear. When compared with measurements of specimens of the two species in question (Frazier, unpublished data), the dimensions of the shell — 11" (28 cm) long; 7.5 " (19 cm) wide; and 5 " (13 cm) high — are consistent with Indotes- tudo elongata and inconsistent with Testudo horsfieldii.
Recent specimens and records from the terai: In 1974 C.A. Ross (Ross and Crumly 1983) found a specimen of Geochelone (= Indotestudo ) elongata "in the vicinity of Gairal Forest Rest House, Corbett National Park, about 25-30 km north-west of Ramnagar." This locality (Fig. L) is about 75 km west of the Nepalese border, in Garhwal district, Uttar Pradesh.
On 16 April 1985, Dr J.C. Mitchell found the remains of a shell of Indotestudo elongata (USNM 267020) at Sauraha, Chitwan, Narayani district, Nepal (Fig. 1). Although incomplete and dog-chewed, this appears to be the only Nepalese specimen of this species deposited in a museum (although it could not be found ip September 1988).
Indotestudo elongata apparently also occurs in the vicinity of Dehra Dun, Uttar Pradesh, India, to the west of Nepal (Fig. 1). Dr R.K. Bhatnagar, formerly in charge of the Herpetology Section of the Zoological Survey of India (ZSI) station in Dehra Dun, wrote ( in litt. 4 December 1987) that "before 1970" he collected a gravid female tor- toise (unidentified) £rom Phandowala, Dehra Dun Siwaliks, now Rajaji National Park; the specimen was left at the ZSI station. Dr W. Auffenberg reported (in litt. December 1987) that he has "seen a slide of a specimen photographed near Dehra Dun and it is elongata ". Apparently, the slide came from Dr. R.K. Bhatnagar.
Dr R. Tilak, Officer-in-charge of the ZSI station, Dehra Dun, reported (in litt. 5 February 1988) that they had no trace of any specimen of T. horsfieldii , but did have a mounted specimen of I. elongata. Measurements and photographs of this specimen, provided by B.C. Choudhry (in litt. 13 March 1989), show that it is an adult female I.
THE LAND TORTOISE IN NEPAL : A REVIEW
51
elongata (curved carapace measurements: length 27.3, width 22.7 cm.; 17 to 18 growth rings), with very little black on the carapace.
These recent records show that Indotestudo elongata occurs along the Terai, or Himalayan foothills, as far west as Dehra Dun. This further supports the occurrence of this species in Nepal.
Conclusions
There is no evidence to support the conten- tion that Testudo horsfieldii occurs in Nepal. The occurrence of Indotestudo elongata , although confused by more than a century of misidentifica- tion, is traceable back to the first evidence of a species of Testudinidae in that country - viz. Hodgson’s drawing.
It must be appreciated that when Gunther (1864 : 8) wrote about Hodgson’s drawing, he had only seen one specimen of T. horsfieldii. Further- more, I. elongata was described in 1853 from Burma (Blyth 1853), and the first recorded acces- sions in the BM (NH) of this species, all from Indo-China, were in 1861 and 1862, and at the time of Boulenger’s Catalogue (1889: 174) there was still not a single specimen in the BM (NH) from a locality near to Nepal. In short, when Gunther was writing, both species were repre- sented by very few specimens, in the BM (NH) at least, and the known range of T. horsfieldii was much closer to Hodgson’s ‘locality’ than was the known range of I. elongata.
The resolution of which species of tortoise occurs in Nepal is of central importance to zoogeographic arguments. Well known for har- bouring not only endemics, but various faunal elements from diverse zoogeographic regions and subregions (e.g. Swan and Leviton 1962), Nepal provides unique insights into the biogeographical history of these animals. One interpretation is that the Indo-Chinese tortoise has been able to expand its range westward across the Brahmaputra, along the Himalayan foothills and past the Ganges. This is consistent with the Satpura Hypothesis of Hora (1948).
However, together with the extensive north Indian/Nepalese distribution of I. elongata , one
must consider the closely related Travancore tor- toise Indotestudo forstenii (Schlegel and Muller), isolated some 2,000 km to the south in the Western Ghats. This situation is strong support for Smith’s statement (1931 : 16): "The Indo-Chinese hill tortoises, Testudo elongata and Geoemyda tricarinata, did not extend their range into the peninsula of India (Chhota Nagpur) by crossing the Gangetic Plain" (and on p. 143: "That it [I. elongata ] ever crossed the Gangetic Plain as we know it today is, of course, highly improbable"). Instead, these represent relict distributions of an ancestor that was widely distributed during a period when environmental conditions were very different from what they are now.
It is important to point out that Ross and Crumly (1983 : 429) misrepresented Smith (1931 : 16, 143) in stating that "Smith also contended that it was unlikely that G. elongata ever extended across the Gangetic Plain." Smith clearly was concerned that there are species whose present range extends across the Gangetic Plain, and he seemed to favour the argument that this geographic distribution predated the Gangetic Plain.
Elements of both Smith’s (1931) and Hora’s (1948) explanations are compatible; the two hypotheses would need to be mutually exclusive only if the time period under consideration were the same. If I. elongata truly is of Indo-Chinese origin, then at some point it, or its ancestor, had to expand its range westward some thousands of kilometres, across what is today Nepal and as far as Dehra Dun. This could then have been followed by a change in environmental conditions (perhaps even the birth of major rivers such as the Ganga and Brahmaputra) and subsequent isolation of the population into southern and northern hill sites.
The Nepalese and Uttar Pradesh records show that despite changes which might have taken place in the environment and distribution of I. elongata , it has managed to survive across the length of the Gangetic plain, nearly to the south- eastern limit of the Palearctic region. Its occur- rence in both the terai and Chhota Nagpur show that it is on both sides of the Ganga, and the
52
JOURNAL, BOMBAY NATURAL HIST. SOCIETY, Vol. 89
species is unquestionably on both sides of the Brahmaputra.
Ironically, a complementary question which surfaces in the light of this evidence is: why has I horsfieldii not be able to expand into the western Himalayan area? The eastern limit of its geographic range (Fig. 1) extends to central Af- ghanistan and Baluchistan (although there are questionable records from the major cities of Kabul and Islamabad). The answer appears to be in the major mountain ranges of eastern Afghanis- tan, northern Pakistan and Kashmir. Although the species is said to live at altitudes between 1,000 and 2,000 m (J. Anderson, pers. comm.), there is no evidence that this palea retie species has ever been able to survive - much less cross - these
Refer
Anderson, J. (1878-9): Anatomical and zoological researches and zoological results of the Yunnan Expeditions. Reptilia and Amphibia. Calcutta, pp. 705-860.
Annandale, N. (1913): The tortoises of Chota Nagpur. Rec. Ind. Mus. 9 (2) : 63-78, pis. v & vi.
Archer, M. (1962): Natural history drawings in the India Office Library. Her Majesty’s Stationery Office, London. Auffenberg, W. (1974): Checklist of fossil land tortoises (Tes- tudinidae). Bull. Florida State Mus. 18(3): 121-251.
Baylis, H.S. & Daubney, R. (1922): Report on the parasitic nematodes in the collection of the Zoological Survey of India. Mem. Ind. Mus. 7(4): 263-347.
Biswas, S., Acharjyo, L.N. & Mohapatra, S. (1978): Notes on the distribution, sexual dimorphism and growth in cap- tivity of Geocheloneelongata (Blyth)./. Bombay nat. Hist. Soc. 75: 928-930.
Boulenger, G.A. (1889): Catalogue of the Chelonians, Rhynchocephalians, and crocodiles in the British Museum (Natural History). British Museum, London. Reprinted 1966, Wheldon & Wesley and Verlag Cramer; Leher, Ger- many.
Blyth, E. (1853): Notices and descriptions of various reptiles, new or little known./. Asiat. Soc. Bengal 22 (5): 639-655. Champion, H.G. & Seth, S.K. (1968): A revised survey of the forest types of India. Govt, of India Press, New Delhi. Daniel, J.C. (1983): The bookof Indian reptiles. Bombay Natural History Society, Bombay.
Das, I. (1988): A survey of land tortoises and freshwater turtles of north-eastern India: Final report. IUCNAVWF Project 6343.
Ghaub, S.A., Rahman, H., Iffat, F. & Hasnain, SA. (1976): A checklist of the reptiles of Pakistan. Rec. Zool. Surv. Pak.
high ranges (over 3,000 m) with rigorous climate and impoverished soil and vegetation.
Acknowledgements
Valuable information and comments were given by K. Adler, K. Asher, W. Auffenberg, R. K. Bhatnagar, B.C. Choudhry, I. Das, J.B. Iverson, J.C. Mitchell, A.E. Leviton, W.A. Rodgers, A.F. Stimson, R. Tilak, R.C. Waltner, R.G. Webb, and C. Wemmer. The library of the British Museum (Natural History) kindly allowed me to examine the Hodgson drawings and to photograph and reproduce plate 9 from that collection here. Infor- mation was compiled while supported by the Indo-American Fellowship Program and the U.S. Fish and Wildlife Service.
•NCES
8 (1 & 2): 37-59.
Gunther, A. [C.L.G.] (1861): List of cold-blooded vertebrata collected by B.H. Hodgson, Esq., in Nepal. Proc. Zool. Soc., Load. 1861: 213-227.
Gunther, A.C.L.G. (1864): The Reptiles of British India. Ray Society, London. Reprinted 1987 ? Oxford and IBH Publishing Co., New Delhi.
Hodgson, B.H. (n.d.): Drawings of reptiles, fishes, etc., presented by B.H. Hodgson. Original in the library of the British Museum (Natural History).
Hoogmoed, M.S. & Crumly, C.R. (1984): Land tortoise types in the Rijksmuseum van Natuurlijke Historie with comments on nomenclature and systematics (Reptilia : Testudines: Testudinidae). Zool. Meded. 58 (15): 241-259.
Hora, S.L. (1948): The distribution of crocodiles and chelonians in Ceylon, India, Burma and farther east. Proc. Nat. Inst. Sci. India. 14 (6): 285-310.
Iverson, J.B. (1986): A checklist with distribution maps of the turtles of the world. Privately printed; Richmond, Indiana. Jayaram, K.C. (1949): A note on the distribution of chelonians of peninsular India with Malayan affinities. Proc. Nat. Inst. Sci. 15 (8): 397-398.
Jayaram, K.C. (1974): Ecology and distribution of freshwater fishes, amphibians and reptiles.//i: Mani, M.S. (ed.). Ecol- ogy and biogeography in India. Junk, The Hague, pp. 517-584.
Laurie, W A. (1978): The ecology and behaviour of the greater one horned rhinoceros. Unpublished D. Phil, thesis, Cambridge University.
Majupuria, T.C. (1981-82): Wild is Beautiful: Introduction to the magnificent, rich and varied fauna and wildlife of Nepal. S. Devi, Lashrar (Gwalior).
J. Bombay nat. Hist Soc. 89 Plate 1
Frazier: Land tortoise in Nepal
mgKm m
Plate 9 of the BM (Nil) collection of B.H. Hodgson’s drawings. Photo courtesy British Museum (Natural History).
THE LAND TORTOISE IN NEPAL : A REVIEW
53
Prater, S. (1980): The Book of Indian Animals (4th ed.). Bom- bay Natural History Society, Bombay.
Ross, C.A. & Crumly, C.R. (1983): A range extension of Geochelone elongata. J. Bombay nat. Hist. Soc. 79(2): 429-430.
Sawyer, F.C. (1971): A short history of the libraries and list of mss. and original drawings in the British Museum (Natural History). Bull. br. Mus. nat. Hist. (Hist. Ser.) 4: 77-204.
Shammakov, S. (1981): Presmikayushcheyesya Ravnennovo Turkmyenestayanya. Academy of Sciences of Turkmenis- tan S.S.R., Ashkabad Ilim.
Smith, M A. (1931): The Fauna of British India, including Ceylon and Burma. Reptila and Amphibia. Vol. I. Loricata, Tes- tudines. Taylor and Francis, London. Reprinted 1973, Ralph Curtis Books, Miami; and 1974, Today & Tomorrow’s Printers & Publishers, Faridabad.
Stainton, J.D A. (1972): Forestsof Nepal. John Murray, London.
Swan, L.W. & Leviton, A.E. (1962): The herpetology of Nepal: A history, checklist and zoogeographic analysis of the herpetofauna. Proc. Cal. Acad. Sci., Sr. 4. 32 (6) : 103-147. Republished 1966, J. Bengal, nat. Hist. Soc. 34 (2): 88- 144.
Theobald, W. (1870): Observations on a paper by Dr J.E. Gray,
entitled ‘Notes on the families and genera of tortoises’ etc. Proc. Zool. Soc., Land. 1890: 674-677.
Waltner, R.C. (1973a): Geographical and altitudinal distribu- tion of amphibians and reptiles in the Himalayas (Part I). Cheetal 16(1): 17-25.
Waltner, R.C. (1973b): Geographical and altitudinal distribu- tion of amphibians and reptiles in the Himalayas (Part II). Cheetal 16 (2): 28-36.
Waltner, R.C. (1973c): Geographical and altitudinal distribu- tion of amphibians and reptiles in the Himalayas (part III). Cheetal 16 (3): 14-19.
Waltner, R.C. (1973d): Geographical and altitudinal distribu- tion of amphibians and reptiles in the Himalayas (Part IV). Cheetal 16 (4): 12-17.
Webb, R.G. (1980): Gray, Hardwicke, Buchanan-Hamilton, and drawings of Indian Softshell turtles (family Trionychidae). Amphibia-Reptilia. 1(1): 61-74.
Yatyayev, Ch. (1985): Presmikayushcheyesya gor Turkmyenes- tyanya. Academy of Sciences of Turkmenistan S.S.R., Ashkabad Ilim.
Zhao Ermi (1973): A new record of Chinese land tortoise from Sinkiang - Testudo horsfieldi Gray. Acta Zoologica Sinica 19 (2): 198. (in Chinese).
Appendix I
LOCALITY RECORDS OF Indotestudo elongata AT THE WESTERN LIMIT OF ITS RANGE
1. Phandowala, Rajaji National Park, Dehra Dun Siwaliks, Uttar Pradesh, India; before 1970; coll. R.K. Bhatnagar; gravid female; probably the unnumbered mounted female in the Zoological Survey of India (ZSI), Dehra Dun.
2. Gairal Forest Rest House, Corbett National Park, 25-30 km NW of Ramnagar and about 75 km W of Nepal, Garhwal District, Uttar Pradesh, India: April 1974; Ross and Crumly 1983; photo only.
3. Sauraha, Chitwan District, Narayan, Nepal; 16 April 1985; 200 m; coll. J.C. Mitchell; USNM 267020; dog- eaten shell.
4. Sarugara Forest, ‘4th mile’ on Siliguri - Kalimpong road, Darjeeling District, West Bengal, India; 1970’s; Kumirmari in Das 1988: sight records, kept as pets.
5. Baradighi Tea Estates, Jalpaiguri District, West Bengal, India; 1915-1916; coll. W.L. Traves; seven specimens/records in ZSI: 17992 entire in spirit, 18016 entire in spirit, 18046 given to Agra College, not traced;
18124 donated to Punjab government, not traced,
18125 entire in spirit, 18162 dry shell, 18171 not seen since 1984; see also Baylis and Daubney 1922 : 303, 312 and Das 1988 : 21.
6. Gorumara, Jalpaiguri District, West Bengal, India; Sub- imal Roy in Das 1988 : 22; sight record.
7. Damanpur, Buxa Division, Jalpaiguri District, West Bengal, India; Subimal Roy in Das 1988 : 22; sight record.
8. Western Assam, India; locality from Hoogmoed and Crumly 1984 : Fig. 3; it is not known on what this record was based; see Das 1988 : 4.
9. Mangaldai, Darrang District, Assam, India; 1957-58; S.K. Sharma in Das 1988 : 3; sight record.
10. Tura, Garo Hills, Meghalaya, India; referred to in parasitological study of Baylis and Daubney 1922 : 304, 312.
11. West Bhanugach Reserve Forest, Moulri Bazar Dis- trict, Bangladesh; 11 January 1989; Das in litt. 4 April 1989; shell.
12. Nainimukhi, Chittagong Hill Tracts, Bangladesh; 20 February 1922; ZSI 19239 entire in spirit.
13. Chaibasa, Chhota Nagpur, Bihar, India; Anderson 1878-9: specimen not traceable according to Annandale 1913 : 76.
14. Jasandra Singhbhum, Chaibasa, Singhbhum District, Chhota Nagpur, Bihar, India; 17 February 1901; ZSI 11379 - type of Testudo parallelus Annandale 1913 : 76ff.; shell etc. in spirit, dry skull.
54.
JOURNAL , BOMBAY NATURAL HIST. SOCIETY, Vol. 89
15. Kumdi Village and Kodlibad Village, Saranda Forest,
Si nghbhum District, SW of Chaibasa, Bihar; 1982;
Coll. E.O. Moll and J. Vijaya, EOM 2711 and 2712;
Moll in litt. 9 Sept. 1988 & 7 Nov. 1988; two dry shells from the villages.
16. Simuli Pahar, Mayurbhanj District, Orissa; 22 February 1971; Biswas et al. 1978; live male in Nandankanan Zoo.
Appendix II
LOCALITY RECORDS OF Testudo horsfieldii AT THE EASTERN LIMIT OF ITS RANGE
Florida 20643 and 21353, D.L. Auth in litt. 3 October 1988.
E. Khanai, Quetta District, Baluchistan^Pakistan: 28 April 1906; two ZSI records; 15552 dry shell and skull, 15553 sent to Munich Museum.
F. 12 miles east of Balkh, Afghanistan; BM(NH) 1940. 5.1.1; dry shell.
G?) Safed-i-Rak, Kabul, Afghanistan; ZSI 5591 to 5600 ten eggs in spirit. This locality is questionable since it is a major city, and it is likely that the specimen was a captive.
H?) Islamabad, Pakistan; MTKD (Staatliches Museum fur Tierkunde, Dresden, Germany (DDR)) N. N. This locality is questionable since it is a major city, and it is likely that the specimen was a captive.
A. Pap Hills, Lasbela district, Baluchistan, Pakistan; PZS 40 a; entire specimen in spirit; Ghalib et al. 1976 : 39.
B. Kalat (Kelat), Baluchistan, Pakistan; 1906; coll. G.H. Tipper; eight ZSI specimens/records; 15537 not trace- able. 15538 entire in spirit, 15539 sent to Vienna Museum, 15541 dry shell and skull, 15542 dry shell and skull, 15543 dry plastron and skull (‘sent to Munich Museum’), 15551 dry shell and skull and 3 eggs, 15725 eggs not traceable.
C. Kowas, Sibi District, Baluchistan, Pakistan; 5 March 1976; coll. M. Farooq Ahmed; Zoological Survey of Pakistan (PZS) 40; entire in spirit; Ghalib et al. 1976 : 39.
D. Quetta, Baluchistan, Pakistan; at least four specimens: BNHS 3, dry shell; ZSI 16480, skull; University of
17. Puri District, Orissa; 27 April 1969; ZSI 23550; Biswas
et al. 1978; dry shell. However, L.A.K. Singh in litt. 3 February 1988, states that ZSI 23550 was collected in the autumn of 1968 from near Barbara Rest House, Banapur village, Tamana or Rajin Forest Block, con- tiguous with Satkoshia Gorge Sanctuary (north of Mahanadi River), evidently in Cuttack disk
A CATALOGUE OF THE BIRDS IN THE COLLECTION OF BOMBAY NATURAL HISTORY SOCIETY - 35 : TROGLODYTIDAE, CINCLIDAE, PRUNELLIDAE, PARIDAE, SITTIDAE
AND CERTHIIDAE
Humayun Abdulali and Saraswathy Unnithan (Continued from Vol. 88 (1) : 80)
This part covers 866 specimens of 88 species and subspecies, nos. 1769-1851 in handbook of the birds ofindiaand Pakistan & synopsis and 14 extra limitals. Of the 88 from Indian limits we have no specimens of 18 forms (3 species and 15 subspecies). As in Part 34, the bulk of the work has been done by S.U. and H.A. has only read over and checked the final work.
1769. Troglodytes troglodytes magrathi (Whitehead) (Safed Koh) Magrath’s Wren. 1 : 446
nil.
1770. TVoglodytes troglodytes neglectus Brooks
(Kashmir) Kashmir Wren 1 : 446
16: 6 males (2 juv.) 4 females (2 juv.) 6 ? (2 juv.).
5 Liddar Valley, 1 Barnaibut, 2 Kashmir, 1 Tara Devi, 1 Patiala State, 1 Fagu, Keonthal State, 1 Koti State, 4 Simla, N.W. Himalayas.
There are six juveniles collected during June July and August among them whose measure- ments do not show much difference. Measurements on p. 64.
1771. TVoglodytes troglodytes nipalensis Blyth
(Nepal) Nepal Wren 1 : 445
Darker than 1770 though collected in the cold weather
11:3 males 7 females 1 ?
1 Mangalbara, E. Nepal, 4 Phalut, Darjeeling DL; 3 Lachung, N. Sikkim, 2 Chimakothi, 1 Chapcha, W. Bhutan.
There are no juveniles all being collected in winter, November to February.
Measurements on p. 64.
EL. Troglodytes troglodytes hyrcanus (Zarudny and Loudon) Northern Iran and Turkmenia
1 ? Dohuk, Kurdistan.
Chin to vent washed with white, upper parts paler than in both 1770 and 1771.
1772. Cinclus cinclus leucogaster Bonaparte (West Siberia) Whitebellied Dipper
2:3
2: 1 male 1 female
1 Tashkent, U.S.SJl., 1 Boston Terek.
Measurements on p. 64.
1773. Cinclus cinclus cashmeriensis Gould
(Kashmir) Whitebreasted Dipper 2 : 2
3: 1 male 1 female 1 ?
1 Chagre, 1 Kardong, Ladak; lAmuchhu; R. Phaxma, Tibet 11000'.
Measurements on p. 64.
1774. Cinclus cinclus przewalskii Bianchi (Den-
chu, basin of the Yangtse-kiang) Eastern whitebreasted Dipper 2 : 2
nil.
1775. Cinclus pallasii tenuirostris Bonaparte (Himalayas) West Himalayan Brown Dipper
2:4
23: 14 males (6 juv.) 7 females (2 juv.) 2 ? (both juv.)
6 Chitral, 1 Banihal village, 1 Kulotran, Badrawar, 1 Kashmir, 2 Patiala State, 1 Koti State, 1 Glow stream, 1 Narkanda, 5 Simla; 1 Guptakashi, 1 Yoshinath, 2 Badrinath, Garhwal.
There are 10 juveniles (6 m, 2 f, 2 ?). Entirely different in colouration from the adult. They are grey with numerous white spots, and are smaller in measurements, which are given separately. Measurements on p. 64.
1776. Cinclus pallasii dorjei Kinnear (Sakden, E. Bhutan) East Himalayan Brown Dipper
2:5
6 :1 male 4 females 1?
1 Tibet, 1 Kurseong, 1 Mangdechu, 1 Batase, C. Bhutan, 1 Gomchu. E. Bhutan, 1 Mishmi Hills, Burma.
One each from Batase (March) and Kur- seong (Dec.) are paler above than the others. Measurements on p. 64.
1777. Prunella collaris rufdata (Severtzov) (Turkestan) Turkestan Alpine Accentor 2 : 189
3 : 2 males 1 female
56
JOURNAL, BOMBAY NATURAL HIST. SOCIETY, Vol. 89
2 Kesun, 1 Drosh, Chitral.
Measurements on p. 64.
With the material available no. 1777 and 1778 are barely separable.
1778. Prunella collaris whymperi (Baker) (Garhwal) Garhwal Alpine Accentor 2 : 188
11:3 males 4 females 4 ?
2 Liddar Valley, Kashmir, 3 Kufri, Koti State, 4 Simla, N.W.H.; 2 Nila Valley, Garhwal.
Measurements on p. 64.
1779. Prunella collaris nipalensis (Blyth)
(Kachar region of Nepal) Eastern Alpine Accen- tor* # 2 : 188
5 : 3 males 1 female 1 ? (2 * missing)
1 Lachung, N. Sikkim, 1 Chapcha, W. Bhutan, P*Kama Valley*, 1 E. Everest, Tibet*, 1 Mishmi Hills.
The two marked are missing, but the remain- ing three have the darkest upperparts and grey of breast.
Measurements on p. 64.
1780. Prunella himalayana (Blyth) (Himalaya
range) Altai Accentor 2:191
14 : 4 males 9 females 1 ?
2 Duala-Dar range, Dharmsala, 3 Tara Devi, Patiala State, 3 Mashobra, 2 Simla N.W.H.; 4 Chungthang, Lachung, Sikkim.
Measurements onp. 65.
1781. Prunella rubeculoides (Moore) (Nepal)
Robin Accentor 2 :193
5 : 2 males 1 female 2 ?
1 Debring, 1 Rupshu, Kashmir, 1 Sukta, Ladak, 1 Gyangtse, Tibet, 1 no locality.
Measurements on p. 65.
1782. Prunella strophiata jerdoni (Brooks) (Kashmir) Western Rufousbreasted Accentor
2: 197
16 : 4 males 7 female 5 ?
1 Gulmarg, 1 Mornaula, 1 Pyas, Kishtwar, 3 Liddar Valley, 1 Kashmir; 1 Fagu, Keonthal St., 2 Koti State, 2 Patiala State, 3 Simla N.W.H.; 1 Pindari, Garhwal.
3 (sex?) immature from Liddar (2) and Pin- dari (Garhwal) differ in the absence of the broad band across the breast, the orange-rufous eyebrow and the faint trace of the double wing-bar. The last in white exists in the one from Pindari 10500', and the bird has a paler breast band and pale eyebrows.
Measurements on p. 65.
1783. Prunella strophiata strophiata (Blyth) (Nepal) Eastern Rufousbreasted Accentor 2 : 196
18 : 7 males 3 females 8 ?
2 Lachung, N. Sikkim, 1 Sikkim, 1 Native Sikkim; 1 Honka, 2 Chimakoti, 1 Chapcha, W. Bhutan, 1 Shamgong, 1 Batase, C. Bhutan, 5 Rongtong, 1 Wamrong, 1 Gomchu, 1 Narphong, E. Bhutan.
Measurements onp. 65.
1784. Prunella fulvescens fulvescens
(Severtzov) (Turkestan) Turkestan Brown Accen- tor 2 : 198
5: 1 male 2 females 2 ?
1 Chitral, Drosh, 1 Gilgit, 1 Kashgar, Vibulak, 1 Kar- dong, 1 Sasar Pass, Ladak.
Measurements on p. 65.
1785. Prunella fulvescens sushkini Collin &
Hartert (Khamba-jong, Tibet) Tibet Brown Ac- centor 2 : 198
1 male Kaungmar; Tibet.
Measurements on p. 65
1785a. Prunella fulvescens ocularis (Radde) (Kiz Yurdi Mt, Talych) Radde ’s Accentor 8 : 626 1 (sex?) Chaman, Baluchistan.
1786. Prunella atrogularis huttoni (Moore) (Simla) Turkestan Blackthroated Accentor
2: 194
11: 4 males 4 females 3 ?
1 Quetta, 1 Wana, Waziristan, 2 Drosh, 1 Ayun, 2 Chitral, 1 Gora Gali, Murree Hills, 1 Rawalpindi; 1 Patiala St., N.W.H.; 1 Chandigarh, Ambala Dt.
Measurements on p. 65.
1787. Prunella atrogularis atrogularis (Brandt) (Semipalatinsk) Ural Blackthroated Accentor
2:194
nil.
1787a. Prunella montanella montanella (Pal- las) (Dauria) Siberian Accentor nil.
1788. Prunella immaculata (Hodgson) (Central
and northern regions of the Hills, Nepal) Maroon- backed Accentor 2:193
6: 1 male 2 females 3 ?
1 Shamgong, C. Bhutan, 3 Gomchu, 1 Narphong, E. Bhutan, 1 North Shan States, Burma.
Measurements on p. 65.
CATALOGUE OF BIRDS IN THE BNHS COLLECTION
57
1789. Melanochlora sultanea sultanea
(Hodgson) (Central and northern regions of the Hills, Nepal) Sultan Tit. 1 : 101
28 : 17 males 11 females
1 Nepal, 2 Longview TE, 2 Savoke, Darjeeling Dt, 2 Pershoke, Sikkim, 1 Martham, Rongni Valley, 2 Tama, C. Bhutan, 2 Deothang E. Bhutan, 1 Singtam, Teesta Valley, 1 Hungrum, N. Cachar, 1 Bagh-o-bahar, Cachar, 2 Margherita,
2 Tezu, Lohit Valley, U. Assam; 1 Tarajuli, Arunachal Pradesh, 1 Memon, U. Burma , 1 SE of May my o, 1 Mansum, 1 Dabahka, 3 Tangti, North Shan States, Burma , 1 no locality.
Measurements on p. 65.
1790. Parus major ziaratensis Whistler (Ziarat, Baluchistan) Baluchis.tan Grey Tit
1 : 76
1 female, Harboi, Baluchistan.
Measurements on p. 66.
1791. Parus major decolorans Koelz (Jalabad, eastern Afghanistan) Afghanistan Grey Tit 1 :76
nil.
1792. Parus major caschmirensis Hartert (Gil-
git) Kashmir Grey Tit 1: 76
14 : 6 males 5 females 3 ?
3 Chitral, N.W.F.P.; 1 Dachigam, 1 Badat, Kishtwar, 1 Kashmir, 1 Jhelum, 1 Jhalor, nr. Campbellpur, 1 Lahore, 1 Ambala, Punjab, 1 Bhajji St., 3 Simla, N.W.H.
A high altitude bird, descends to lower eleva- tions in winter.
Measurements on p. 66.
1793. Parus major nipalensis Hodgson (Nepal)
Nepal Grey Tit 1 : 74, 77
10 : 6 males 3 females 1 ?
1 Bhajji State, 1 Gama-ki-Hatti, Dharmi State, 1 Kalka,
3 Simla N.W.H., 1 Kqruprayag, 1 Ranighet, 1 Almora, Garhwal, 1 Pilibhit Terai, U.P.
Averages smaller than caschmirensis. Measurements on p. 66.
1794. Parus major stupae Koelz (Sanchi,
Bhopal) Indian Grey Tit 1 : 77
41: 19 males 12 females 10 ?
3 Sunda Hill, Jaswantpura, Dt. Jodhpur, 1 Narwar Fort, Gwalior State, 1 Gangasagar, Palanpur St., Gujarat, 2 Jam- bughoda, 2 Dohad, Panch Mahals, 1 Dalkhania, Amreli Dt., Kathiawar, 1 Ghatwad,S. Kathiawar, 1 Malegaon, 1 Laochali, Surat Dangs, 2 Bijwar, Indore St, 1 Jabalpore, C.I., 1 Bhanupratappur, Kanker, 2 Konta, Bastar Dt, C.P., 1 Nasik, 1 Suriamal, Thane Dt, 1 Mehda, 1 Satara, 2 Dodballapur,
Bangalore, 1 Kumili, 1 Merchiston, Ponmudi, S. Travancore, 1 Kotagiri, Nilgiris, 1 Manalur, Palni Hills, 2 Nelipaka, 1 Kaulas, 1 Paloncha, Hyderabad, 1 Kutri, Daspalla, 3 Badrama, Bamra, 1 Chahala, Simlipal, Orissa, 3 Rajapati, Chapra, Saran Dt., Bihar.
Measurements on p. 66.
1795. Parus major mahrattarum Hartert
(Ceylon) Ceylon Grey Tit 1 : 77
nil.
1796. Parus major vauriei Ripley (Chabua, Northeastern Assam) Lakhimpur Grey Tit 1 : 74
4 : 3 males 1 female
3 Dibrugarh, 1 Sadiya, U. Assam.
Measurements on p. 66.
1797. Parus major tibetanus Hartert (Chaksam in Tsangpo Valley, Tibet) Tibet Grey Tit 1 : 346
nil.
EL. Parus major commixtus (Swinhoe) (South China) Burmese Great Tit 1 : 78
4 : 1 male 1 female 2 ?
1 U. Burma, 1 Maymyo, 1 Saw pakoku, 1 Shatzusth, North Shan States, Burma.
Measurements on p. 66.
EL. Parus major minor Temminck & Schlegel (Japan)
2 males
1 Forgyi, Shan States, Burma, 1 Peking, China.
Measurements on p. 66.
EL. Parus major major Linnaeus (Sweden)
1 male, Hungary ?
Measurements on p. 66.
EL. Parus major intermedius Zarudnyi (Kopet Dagh mountain system of Afghanistan & Baluchistan)
7 : 4 males, 3 females
1 Shiraz , 1 Turbat, nr Meshed , 5 Meshed, Persia.
Measurements on p. 66.
EL. Parus major blanfordi Prazak (Tehran)
6 : 2 males 2 females 2 ?
1 Dohuk, Kurdistan, 1 Shush, Karkheh River, 1 Jungle on Karkheh R., 1 Legation-Gulahek, Tehran, 1 Shiraz, 1 Engeli, Persia.
Measurements on p. 66.
1798. Parus nuchalis Jerdon (Eastern Ghats)
Whitewinged Black Tit 1 : 79
58
JOURNAL, BOMBAY NATURAL HIST. SOCIETY, Vol. 89
11: 7 males 3 females 1 ?
1 Gangasagar, Palanpur, 4 Bhuj, 2 Mata-no-madh, 2 Chaduva, 1 Godsar, Bhuj env., 1 Kutch.
Baker (Faun^t Vol. 1, p. 79) gives one description of the bird, so we presume he intended this for both the sexes. Indian Handbook (Vol. 9, p. 172) says the sexes are alike. Mr Simon Harrap who has been working on the Paridae at the British Museum drew our attention to the following in his mss. He has examined the material at the B.M. and says the sexes are distinct in so far as the male has the entire upper parts deep black, glossed blue, strongest on the crown and mantle. The female has the upper parts duller, sooty brown-black glossed blue on the crown only, flanks and sides of breast with a more distinct pale yellow wash than in the male, the median line on the underparts very sjightly paler. In general our observations agree with this.
In the three female specimens the crown is sooty brown-black without a blue gloss and flanks and sides of breast without any pale yellow wash, (HANDBOOK OF THE BIRDS OF INDIA AND PAKISTAN says this colour fades into pure white in museum specimens) and the median line is definitely paler than in the male. Of the two males from Kutch, one collected on 28 Sept. 1943 and the other going back to July 1896, both lack the glossy black of the other males; the former is either wrongly sexed or is in juvenile plumage while the latter may have faded. The unsexed specimen from Gangasagar is in male plumage.
Perhaps the publication of the Harrap notes may produce additional information from other sources.
Measurements on p. 66.
1799. Parus monticolus monticolus Vigors (Himalayan Mountains = Simla) Greenbacked Tit
1:80
43 : 23 males 13 females 7 ?
1 Dachigam, Kashmir, 2 Dalhousie, Punjab, 12 Simla, N.W.H., 1 Mussoorie, 1 Bora, Almora, 1 Ghat, 1 Pothibassa, 1 Yoshinath, Garhwal, 1 Himalayas, 1 Godavri, Nepal, 1 Kurseong, 4 Rinchinpong, W. Sjkkim, 1 Sikkim, 1 Chimakothi, W. Bhutan, 1 Khosela,! Narphong 1 Wamrong E. Bhutan, 5 Shillong, 2 Kohima, Naga Hills; 1 Dam, 1 Lac-Tango, S. Tibet, 1 Chin Hills, 1 Loi Wong N. Shan States,
Burma.
Measurements on p. 66.
1800. Parus cyanus flavipectus Severtzov (Turkestan) Yellowbreasted Blue Tit
2 : 1 male 1 female
2 Chitral. No yellow is visible on the breast Measurements on p. 67.
1800a. Parus cyanus tianschanicus (Menzbier) (Tien Shan) Tien Shan Blue Tit 1 : 81
nil.
1801. Parus hypermelas (Berezovski & Bianchi) (Shensi and the border of Kansu) Blackbibbed Tit
1:82
nil.
1802. Parus melanolophus Vigors (Himalayan
Mountains= Simla-Almora area) Crested Black Tit 1 : 83
28 : 17 males 6 females 5 ?
2 Chitral, 1 Nultar Valley, Gilgit, 2 Liddar Valley, 1 Kashmir, 1 Rawalpindi, 2 Dalhousie, Punjab, 3 Simla, 1 Fagu, Keonthal St., 1 Mt Hutto, Kumarsain, 2 Kufri, 1 Koti St, 4 Dakuri, Kumaon, 1 Dungari, 1 Mounkhal, 1 Talla, Garhwal,
1 Sirkunda 4 m fromDhanaulti, Mussoorie, 1 Bhawali, Naini- tal; 2 no locality.
Measurements on p. 67.
1803. Parus ater aemodius Hodgson (Nepal)
Himalayan Coal Tit 1: 84
6: 3 males 3 females
5 Lachung, N. Sikkim, 1 Bumthang C. Bhutan. Measurements on p. 67.
1804. Parus rubidiventris rufonuchalis Blyth (range beyond Simla) Simla Black Tit 1 : 85
20: 10 males 9 females 1?
2 Tashkent, U.S.S.R. , 4 Rawalpindi, 4 Chitral, 1 Srinagar, 1 Liddar Valley, 2 7m below Yus, 1 Danguil, Kishtwar, Kashmir, 1 Kaying Bashi, 2 Baghi, Bushahr St., N.W.H., 1 Harsi, 1 Gangotri, Garhwal.
Measurements on p. 67.
1805. Parus rubidiventris rubidiventris Blyth (Nepal and Sikkim, restricted to Kathmandu Val- ley by Ripley) Rufousbellied Crested Tit 1 : 84
1 male, Kalabagh, Jubbal St., Garhwal.
Rufous abdomen separates this subspecies from others.
CATALOGUE OF BIRDS IN THE BNHS COLLECTION
59
Measurements on p. 67.
1806. Parus rubidiventris beavani (Jerdon) (Mount Tonglo, Sikkim) Sikkim Black Tit 1 : 86
3 males
1 Tonglo, 2 Phalut, Darjeeling.
Measurements on p. 67.
1806a. Parus rubidiventris saramatii Ripley. (Mount Saramati, Naga Hills) Nagaland Black Tit.
nil.
1807. Parus dichrous kangrae (Whistler) Koti State, near Simla) Western Brown Crested Tit
1:87
4: 1 male 2 females 1 ?
2 Narkanda, 1 Baghi, Bushahr State, 1 Simla Hills, N.W.H.
Measurements on p. 67.
1808. Parus dichrous dichrous Hodgson
(Nepal) Eastern Brown Crested Tit 1 : 87
3: 2 males 1 female
1 Lachung, N. Sikkim, 2 Tongloo, near Darjeeling. Measurements on p. 67.
1809. Parus xanthogenys xanthogenys Vigors (Himalayan Mountains restricted to Murree by Baker) Northern Yellowcheeked Tit
18 : 11 males 3 females 4?
1 Ghora gali, Murree Hills, 2 Dalhousie, Punjab; 2 Keonthal State, 6 Simla, N.W.H., 1 Bhim Tal, 1 Ranibaug, 1 Mornaula, 1 Bhawali, 1 Naini Tal, 1 Kumaon, 1 Godavri, Nepal.
Measurements on p. 67-68.
1810. Parus xanthogenys aplonotus Blyth (Mountains of central India, restricted to Chaibasa, Singhbhum, Bihar by Whistler and Kinnear) Central India Yellowcheeked Tit
1 :92
33: 17 males 13 females 3 ? (1 male 2 females by plumage).
3 Juna, Rajpipla, 1 Pandwa, 1 Malegaon, Surat Dangs, 1 Lonawala, Western Ghats, 1 Poona, 2 guna, 1 Badarwas, Gwalior State, 1 Saugar, 1 Sehore, Bhopal State, 1 Paryat, Jabalpur, 1 Mandu, Dhar State, 1 Melghat, Raipur, Berar, 1 Lahottar Reserve Forest, 2 Bhanupratappur, Ranker, 1 Darba, Bastar DL 1 Upper Bharakhamba, 1 Chahala, Simlipal, 3 Badrama, Bamra, Orissa, 5 Anantgiri, 1 Valaspara, near Sileru, 1 Sankrametta, Vizagapatnam, 2 Wangasara, Andhra Pradesh.
Measurements on pp. 67-68.
1811. Parus xanthogenys travancoreensis
(Whistler & Kinnear) (Mynall, Travancore) Southern Yellowcheeked Tit 1 : 92
23: 17 males 5 females 1?
1 Colvalli, Goa, 1 South Konkan, 2 Sunkal, 2 Castle Rock, Kanara, 1 Horabail, Sagar, Shimoga, 1 Jagar Valley, Bababudan Hills, Kadur Dt., 1 Sakleshpur, Hassan Dt., Mysore, 2 Mercara, Coorg, 1 Runnimadi, 1 Kottamalai, Cherambadi, 1 Kodanad, beyond Kotagiri, Nilgiris, 1 Perumalmalai, 6 Shembagnur, Palni Hills, 2Santanpara, Car- damom Hills.
There is no polymorphism as stated by Whistler and Kinnear in the description (Unnithan infra p. 126).
Measurements on p. 68.
1812. Parus spilonotus spilonolus Bonaparte
(Himalaya, restricted to N. Cachar by Baker) Himalayan Blackspotted Yellow Tit 1 : 89
15: 8 males 6 females 1 ?
1 Sikkim, 1 Darjeeling; 1 Tama, C. Bhutan, 2 Wam- rong E. Bhutan, 2 Barha Pari, 7 Shillong, 1 Che/ra, Assam.
All these specimens were wrongly identified and registered as Parus xanthogenys. Measurements on p. 68.
1812a. Parus spilonotus subviridis Blyth (Tenasserim) Burmese Blackspotted Yellow Tit.
1: 91
1 (sex?) Pimpri Ban, S. Shan States, Burma.
This was also in the Parus xanthogenys.
EL. Parus palustris hellmayri Bianchi (Peking).
4: 2 males 2 females 4 Temple of Heaven, Peking.
These were in P. major.
Measurements on p. 68.
1813. Sylviparus modestus simlaensis Baker
(Simla) Simla Yellowb rowed Tit 1:88
7: 4 males 3 females 3 Dharmsala, 4 Simla, N.W.H.
The yellow has almost disappeared from the forehead.
Measurements on p. 68.
1814. Sylviparus modestus modestus Burton
(Nepal) Eastern Yellowbrowed Tit 1 : 88
60
JOURNAL, BOMBAY NATURAL HIST. SOCIETY, Vol 89
11: 5 males 4 females 2 ?
1 Mangalbare, E. Nepal, 1 Chungthang, N. Sikkim, 1 Chapcha, West, 2 Chimakothi, West, 1 Shamgong, Central, 3 Gomchu, 2 Rongtong, East Bhutan.
Measurements on p. 68.
1815. Cephalopyrus flammiceps flammiceps (Burton) (Mussoorie) Western Firecapped Tit
2 : 545
9: 5 males 4 females
1 Kalka, 4 Simla, N.W.H., 1 Ladwa, Karnal Dt., 2 Baretha, Bharatpur, 1 Surwaya, Gwalior.
Measurements on p. 68.
1816. Cephalopyrus flammiceps olivaceus
Rothschild (Vicinity of Tengyueh) Eastern Firecapped Tit. 1 :545
3 males
2 Gomchu, E. Bhutan, 1 Goalpara, Assam. Measurements on p. 68.
1817. Remiz pendulinus coronatus (Severtzov)
(Khodzhent, Thrkestan) Penduline Tit 1 : 100
10: 3 males 7 females
1 Karram Darra, Persia ; 2 Kohat, N.W.F.R, 2 Lahore, Punjab, 1 Bahawalnagar, 2 Harunabad, 2 Bahawalpur town env.
Measurements on p. 68.
1818. Aegithalos concinnus iredalei (Baker)
(Himalayas restricted to Simla) Western Red- headed Tit 1 : 93
32: 16 males (3 juv.) 7 females (juv.) 9 ? (5 juv.)
1 Mugalmaidan, 2 Bandarkoti, Kishtwar, 3 Dalhousie, 1 Dharmsala, Punjab, 2 Dakuri, 10 Simla, 3 Koti St., 2 Kufri, Patiala St, N.W.H., 1 Pindari Valley, 2 Loliba, 2 Mussooree, 1 Mornaula, U.P., 1 Godavri, 1 Sheopur Ridge, Nepal.
Juveniles of both sexes have pale forehead and underparts.
Measurements on p. 69.
1819. Aegithalos concinnus rubricapillus (Ticehurst) (Sikkim) Eastern Redheaded Tit 1 : 93
8: 5 males 2 females 1?
1 Chungthang, 1 Rinchinpong, W. Sikkim; 1 Gedu, West, 1 Batase, 1 Shamgong Central, 1 Wamrong, 1 Gomchu, 1 Rongtong, East Bhutan.
Upperparts darker and underparts more rufous than in 1818.
Measurements on p. 69.
1820. Aegithalos concinnus manipurensis (Hume) (Eastern hills, Manipur) Manipur Red-
headed Tit 1 : 94
1 ? Chin Hills, Burma.
EL. Aegithalos concinnus pulchellus (Rippon) (Nanoi, S. Shan States) Shan Redheaded Tit
1 :95
1 ? South Shan States, Burma.
1821. Aegithalos leucogenys (Moore) (Balu
Chughur, north east Afghanistan) Whitecheeked Tit 1 : 97
5: 3 males, 2 females
3 Chitral, 2 Jhalor near Campbellpur, W. Punjab.
All the five are correctly marked leucogenys
by earlier workers.
Measurements on p. 69.
1822. Aegithalos niveogularis (Gould) (North
India restricted to northern Punjab) Whitethroated Tit 1 : 98
nil.
1823. Aegithalos iouschistos iouschistos
(Hodgson) (Nepal) Rufousfronted Tit 1 : 99
4 males
1 Darjeeling, 2 Chapcha, West, 1 Rongtong, East Bhutan.
Measurements on p. 69.
EL. Aegithalos bonvaloti sharpei (Rippon) (Mt. Victoria, Chin Hills) Mt. Victoria Blackheaded Tit 1 : 97
1 ? ML Victoria, Chin Hills.
1824. Sitta europaea cashmirensis Brooks
(Kashmir) Kashmir Nuthatch 1 : 128
5 : 1 male 3 females 1 ?
1 Shinghar, North East Baluchistan, 2 Chitral, NWFP, 1 below Pahalgam, 1 Kashmir.
Measurements on p. 69.
1825. Sitta europaea montium La Touche (Atuntze, N.W. Yunnan) Chinese Nuthatch
nil.
1826. Sitta europaea nagaensis Godwin- Austen
(Naga Hills) Naga Nuthatch 1 : 127
2 ? 1 Mt. Victoria, 1 Keelpkai, South Shan States, Burma.
1827. Sitta castanea almorae Kinnear &
Whistler (Valley of Ramganga between Almora and Pethora) 1 : 125
7: 4 males 2 females 1 ?
3 Ranibaug, Kumaon, 1 Gwaldam, Garhwal, 1 Bans, 1
CATALOGUE OF BIRDS IN THE BNHS COLLECTION
61
Gurua, 1 Bageswar, Almora, U.P.
Measurements on p. 69.
1828. Sitta castanea cinnamoventris Blyth (Darjeeling) Eastern Chestnutbellied Nuthatch
1 : 125
17: 11 males