Tree and shrub mortality and debarking by sambar Cervus unicolor (kerr) in Gir after a drought in Gujarat, India

Biological Conservation 68 (1994) 149-154

TREE A N D SHRUB MORTALITY A N D DEBARKING BY SAMBAR Cervus unicolor (KERR) IN GIR AFTER A DROUGHT IN GUJARAT, INDIA Jamal A. Khan,* W. A. Rodgers,{ A. J. T. Johnsingh & P. K. Mathur Wildlife Institute of India, PO New Forest, Dehradun--248 001, Uttar Pradesh, India (Received 30 October 1992: revised version received 17 June 1993: accepted 14 July 1993)

protected area is now managed with the primary objective of long-term conservation of the lion. The frequent recurrence of severe droughts and a cyclone in 1983 must also have had a significant impact on the vegetation and on animal populations. The cyclone uprooted nearly 1.4 million trees in Gir (Sanat Chavan, pers. comm.). There is some information on the effects of livestock grazing on grass production (Hod& 1969), but little information on the impacts of droughts, cyclones, fires and wood cutting on the Gir ecosystem. In 1986, the Wildlife Institute of India started a research project in Gir to collect base-line data on the fauna and flora. The start of the project was followed by one of the worst droughts of this century in 1987. Extensive mortality in some woody species was subsequently noticed throughout Gir. Sambar Cervus unicolor (Kerr) also began stripping the bark of several tree species to supplement their diet during the drought. This paper summarises these observations during 1987 and 1988.

Abstract

The death and debarking of trees and shrubs, following a severe drought in 1987, were investigated in Gir Lion Sanctuary and National Park, Gujarat, India. Eight line transects totalling 48 km were used to sample the vegetation in three districts. The percentage mortality was highest in Sanctuary West (22.6%) compared to National Park (18.3%) and Sanctuary East (16.7%). Mortality differed significantly among different plant species and in some cases between districts. The extent of debarking pattern also difJered significantly between Sanctuary West (13.2%), National Park (7.7%) and Sanctuary East (none observed). Thirteen plant species were affected. The debarking pattern for different girth classes differed significantly in Sanctuary West and National Park. High plant mortality and debarking have several management implications for Gir and require careful monitoring of vegetation and ungulate populations. Keywords: debarking, Gujarat, India, plant mortality, management implications, sambar.

STUDY AREA INTRODUCTION

Gir Lion Sanctuary and National Park (hereafter called Gir, Fig. 1) in the Kathiawar peninsula of Gujarat, covers an area of 1412 km 2. The vegetation is tropical dry deciduous forest interspersed with tropical thorn forest (Champion & Seth, 1968). Nearly 70°/,, of the total area of Gir is covered with teak Tectona grandis (Linn.) and several associates. Three different management districts exist in Gir, i.e. Sanctuary West (690 km2), National Park (258.7 km 2) and Sanctuary East (463.4 kin2), differing in terms of vegetation, rainfall, topography, human settlement density and hence degradation. Sanctuary West is quite thickly wooded and water is available throughout the year in streams and springs between undulating hills. The National Park is more densely wooded and has more hilly terrain while Sanctuary East has more open wooded grassland vegetation with undulating topography. Grazing by livestock of the Maldhari pastoral community is heaviest in Sanctuary East and least in National Park. The winter, from December to March, in Gir is

Gir Lion Sanctuary and National Park form the last refuge of the endangered Asiatic lion Panthera leo persica (Meyer). It is also the largest (1412 km 2) compact tract of natural vegetation in the Saurashtra region of Gujarat. The dry deciduous forest ecosystem of Gir has a long history of interference and exploitation, which has resulted in the loss and modification of vegetation (Berwick, 1974). The main factors responsible for this alteration are extensive clearance of forest lor agricultural purposes by a growing human population, excessive grazing by local as well as migratory cattle, wood cutting and fires (Hodd, 1969). The * Present address: Centre of Wildlife & Ornithology, Faculty of Life Sciences, Aligarh Muslim University, Aligarh--202001, Uttar Pradesh, India. {Present address: c/o. FAO, PO Box 3, Dar-es-Salam, Tanzania. Biological Conservation 0006-3207/94/$07.00 © 1994 Elsevier Science Limited, England. Printed in Great Britain 149

J. A. Khan, I44. A. Rodgers, A. J. T. Johnsingh, P. K. Mathur

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Fig. 1. Map of the Gir Loin Sanctuary and National Park. - . . . . , Sanctuary West; . . . . , Sanctuary East; . . . . . , roads; e, ness; Q, forest settlement; - - , line transect. cool and dry (average minimum 9°C), followed by a hot dry summer (average maximum 42°C), lasting until mid June. The monsoon breaks in June and continues until September, followed by a dry post-monsoon season until mid December. Average rainfall over the past 28 years from the west end of the park and 10 years from the east is approximately 1000 and 800mm respectively. In 1987, however, Sanctuary West and East received only 199 mm and 456 mm of rainfall respectively. METHODS An initial vegetation survey was carried out while laying out a system of eight line transects during February-March 1987. Each transect was 6 k m in length. The transects were located in the different management districts (three in Sanctuary West, three in National Park and two in Sanctuary East) and were placed in a systematic random design with prior stratification according to management district and permanently marked in the field. As these line transects were used for regular monitoring of ungulate populations, it was also possible to monitor mortality and debarking until the next monsoon in 1988. Vegetation was sampled along the transects on 240 permanent sampling points at regular intervals of 200m. Mortality and debarking data were only collected for shrubs 2-6 m high and trees > 6 m. At every sampling point, the 10 nearest trees and 10 nearest shrubs were examined (all hereafter generally referred to as trees) and the following data were recorded: (a) Girth at breast height

, National Park;

(b) Mortality--only trees and shrubs which died during the drought period were recorded. These included standing as well as recumbent dead trees. (c) Debarking (assessed in four categories): 0 = none; 1 = slight (< 25% of bark circumference removed); 2 -- medium (25-50% of bark circumference removed); 3 -- heavy (> 50% of bark circumference removed). Data on plant mortality were summed over each management district for each species and girth class and the percentage of dead trees was calculated. The number of debarked trees was summed for each district, plant species, girth class and damage category. A series of chi-square contingency tables was used to detect significant differences in plant mortality and debarking between districts. To determine which species were preferred or avoided by sambar, and to decide whether there were significant differences in mortality between species, Bonferroni 95% confidence intervals were calculated following Neu et al. (1974) and Byers et al. (1984) for the proportions of dead and debarked trees. For each species, its proportion in the sample was calculated (P~o = n/]~n~). This was then compared with the proportion of dead individuals among all dead trees (Pie = m/~,m~). Mortality was significantly greater or less than expected when P~o lay outside the 95% confidence limits for Pie" For these calculations, 13 species were excluded which had less than 15 individuals in the whole sample. Similar tests were made for the proportion of debarked trees. Nomenclature of trees and shrubs follows Patel (1984).

151

Plant mortality and debarking in Gir

Table 1. Mortality in different plant species in different districts of Gift Sanctuary West

Plant species

Acacia catechu ,4. chundra A. leucophloea ,4. nilotica A. senegal Aegle marmelos Bauhinia racernosa Boswellia serrata Butea monosperma Cassia fistula Dalbergia latifolia Dichrostachys cinerea Dio~pyros melanoxylon Embliea ojficinalis Grewia tiliaefolia ltelicteres isora ttolarrhena antidysenterica Kydia calycina Lannea coromandelica Millusa tomentosa Sterculia urens Tectona grandis Terminalia crenulata Wrightia tinctoria A'eromphis spinosa Zizyphus mauritiana Z. xylopyrus Total h

National Park

Overall mortality %

Sanctuary East

n

m

%

n

m

%

6.4 20.0 0 13.9 91-1 0 16.6 0 5.1 14.2 33.3 36.8 4.5 32.1 68-1 55.2 81.7 0 7.6 20 20 9-1 52.5 5-3 62.9 4.6 41.8

139 26 18 10 0 0 13 44 8 19 21 2 18 40 28 334 114 17 26 19 9 478 74 139 27 33 1

9 3 1 0 0 0 0 1 0 4 2 0 0 16 1 95 83 14 0 2 0 29 23 2 6 3 1

6.4 11.5 5.5 0 0 0 0 2.2 0 21.1 9.5 0 0 40.0 3.5 28.4 72.8 82-3 0 10.5 0 6.0 31 1.4 22-2 9.09 1O0

0 5 15 19 6 133 11 18 0 0 0 11 17 14 3 4 1 0 6 0 2 0 103 3 2 27 1

0 0 3 2 1 23 0 1 0 0 0 1 0 5 1 0 1 0 1 0 0 0 36 0 0 1 0

0 0 20.0 10.5 16.6 17-2 0 5.5 0 0 0 9.1 0 35-7 33.3 0 100 0 16.6 0 0 0 34.9 0 0 3-7 0

6.4 14-2 9,3 12-1 82.3 17-2 5.5 2.8 4.2 19-2 12-5 25 1.7 365 32.0 34.1 76.9 77,7 4~4 12,5 6,2 7.6 36.8 3-4 52.5 5.2 42.2

22.6

1 828

335

18.3

514

86

16,7

20.2

n

m

%

139 25 10 86 45 0 12 9 39 7 3 19 22 28 22 96 93 1 13 5 5 538 40 150 89 130 43

9 5 0 12 41 0 2 0 2 1 1 7 1 9 15 53 76 0 1 1 1 49 21 8 56 6 18

1 809

410

" n, number of individuals sampled; m, number of individuals dead. J Total includes 13 species which had less than 15 individuals and are not listed in the table.

RESULTS Mortality in plant populations No tree mortality was noticed during the initial survey, the trees beginning to die after the failure of the monsoon in 1987. Similarly, debarking by sambar also started after the monsoon failure. The plant-specific and overall mortality (percentage dead) for different districts of Gir are summarised in Table 1, which shows that 831 out of 4151 trees and shrubs (>2 m) were dead. The plant mortality differed significantly between the districts and was high in Sanctuary West 122.6%) compared to National Park (18.3%) and Sanctuary East (16.7%) 0( 2 = 14.4, d.f. = 2, p < 0.001). Out of 40 species sampled, 39 had suffered some mortality. Overall % mortality differed significantly among different plant species and was higher than 'expected' ;in relation to the available proportion of trees) in .4cacia senegal (82.3%), H. antidysenterica (76.9°/,) Helicteres isora (34.1%), Terminalia crenulata (36.8%) and Xeromphis spinosa (52.5%) while the remaining species suffered mortality either in proportion to their availability or significantly lower mortality than expected ITable 2). Plant species which were c o m m o n to all three districts, and had large sample sizes in at least two

districts, were tested for differences in mortality between the districts. The species which showed differential mortality between three units included Grewia tiliaefolia, Helicteres isora, Holarrhena antidysenteriea, Terminalia crenulata and Xeromphis spinosa. The mortality rates for different girth classes showed no significant differences in Acacia senegal, Emblica officinalis, Grewia tiliaefolia, Holarrhena antidysenterica, Terminalia crenulata and Xeromphis spinosa (t9 > 0-05). In Tectona grandis, the 0-20 cm girth class had significantly higher mortality whereas the 41--60 cm class showed significantly lower mortality than expected. In the case of Zizyphus xylopyrus, the 0-20 cm girth class showed significantly lower mortality while the 61-80 cm class had suffered higher mortality than expected.

Debarking pattern Debarking was greater in Sanctuary West as compared to National Park and was not recorded at all in Sanctuary East (Table 3). The proportion available, proportion debarked and 95% Bonferroni confidence intervals for different plant species in Sanctuary West and National Park are provided in Table 4. Thirteen plant species were found to suffer from debarking, of which five were preferentially selected by sambar and three were rarely selected. A similar analysis showed that the debarking pattern

152

J. `4. Khan, W. .4. Rodgers, A. J. T. Johnsingh, P. K. Mathur Table 2. Proportion available (P/o), proportion dead (P/~) and 95% Bonferroni confidence limits for different plant species in Gir

Plant species

Acacia catechu A. chundra A. leucophloea A. nilotica A. senegal Aegle marmelos Bauhinia racemosa Boswellia serrata Butea monosperma Cassia fistula Dalbergia latifolia Dichrostachys cinerea Diospyros melanoxylon Emblica officinalis Grewia tiliaefolia Helicteres isora Holarrhena antidysenteriea Kydia calycina Lannea eoromandelica Millusa tomentosa Stereulia urens Tectona grandis Terminalia crenulata Wrightia tinctoria Xeromphis spinosa Zizyphus mauritiana Z. xylopyrus

Pio

Pie

Confidence limits for Pie

Mortality rating~

0-075 0.015 0.011 0.030 0.013 0.035 0-009 6 0-019 0.012 0.006 9 0-006 4 0.008 5 0-015 0.022 0.014 0.116 0-055 0-004 8 0.012 0-006 4 0-004 2 0.27 0.058 0.078 0.031 0.050 0-012

0.023 5 0.010 4 0.005 2 0.018 2 0.054 8 0.030 0 0.002 6 0-002 6 0.002 6 0.006 5 0.003 9 0.010 4 0.001 3 0-039 1 0-022 1 0.193 2 0.208 8 0.018 2 0.002 6 0.003 9 0.001 3 0.101 8 0.104 4 0.013 0 0.080 9 0.013 0 0-024 8

0.006 532 < P1 < 0.040 467 0-000 963 < P2 < 0.021 763 0.002 855 < P3 < 0.013 255 0.003 227 < P4_< 0.033 172 0-029 308 < P5-< 0.080 291 0.010 892 _
-0 0 0 + 0 ---0 0 0 -0 0 + + 0 -0 0 -+ -+ -0

0, Mortality proportional to availability; --, significantly lower mortality; +, significantly higher mortality. differed significantly between different girth classes in Sanctuary West (g 2 = 94.1, d.f. : 4, p < 0.001) and in National Park (X2 = 152.06, d.f. = 3, p -- <0.001) (Table 5). Sambar clearly selected smaller trees and avoided those with >40 cm girth. The proportion of trees in different damage categories, and the overall percent damage for different plants species, is given in Table 6. Out of a total of 1678 trees, 9.8% were severely debarked while 8.7% and 4.1% had medium and low levels of debarking. Among plant species, the damage was high in Wrightia tinctoria (79.9%), Xeromphis spinosa (73.2%) and X. uliginosa (62.5%). These species also had a high proportion of severely debarked trees, followed by medium and low categories. DISCUSSION The preliminary vegetation survey in Gir, prior to the failure of the 1987 monsoon, did not show mortality in Table 3. Debarking in different districts of Gir (trees and shrubs > 2 m height)

District Sanctuary West National Park Sanctuary East

Trees sampled 1 809 1 828 514

Trees debarked 239 ~ 142 0

% Debarking 13.2 7.7 0-0

any plant species. Since mortality and debarking by sambar only continued until the next monsoon in 1988, this clearly suggests that the severe drought in 1987 was responsible. The significantly greater mortality in Sanctuary West and National Park as compared to Sanctuary East was also associated with the relatively low rainfall in the west, which was unusual. The differential mortality patterns a m o n g different plant species were generally as expected as the species are distributed along a wet-to-dry moisture gradient in Gir (unpublished data). Those favouring wet conditions suffered higher mortality, the only exception being the thorny species Acacia senegal, which is adapted to dry conditions and had a very high mortality (82.3%). Uprooted individuals showed a very shallow root system, making this species vulnerable to such exceptional drought. Available literature on food habits of sambar (e.g. Brander, 1923; Schaller, 1967) suggests that sambar occasionally do eat bark of several plant species, though its contribution to the overall diet remains negligible. N o previous record of debarking by sambar exists from Gir (Berwick, 1974; Sanat Chavan, pers. comm.) and it seems to have been induced by the extremely low grass production after failure of the monsoon, and grazing by approximately 0.2 million cattle during October 1987 (Khan et al., 1990). Chital

Plant mortality and debarking in Gir

153

Table 4. Proportion available (P~), proportion debarked (P~) and 95% Bonferroni confidence limits for different plant species in Sanctuary West and National Park" Plant species

n

Pio

D

Pie

Confidence limits for Pie

Preference rating

Sanctuary West

Cassia fistula Ehretia laevis Emblica officinalis Flaucortia indica Sapindus emarginatus Schrebera swietenioides Tectona grandis Wrightia tinctoria Xeromphis spinosa 3: uliginosa

7 8 28 5 3 9 538 150 89 4

0.008 0.009 0.033 0-005 0.003 0.010 0.639 0.178 0.105 0.004

3 5 2 9 5 7 7 3 8 7

1 1 10 1 1 1 34 121 67 2

0-004 0.004 0-041 0.004 0-004 0.004 0.142 0.506 0.280 0.008

1 1 8 1 1 1 2 2 3 3

0.007 0.007 0-005 0.007 0.007 0.007 0.078 0.415 0.198 0.008

514 < 514 < 423 < 514 < 514 < 514 < 718 < 325 < 661 < 190 <

44 28 19 478 139 27 4

0.059 0.037 0.025 0-646 0.188 0.036 0.005

5 8 7 8 0 5 4

1 1 1 8 110 18 3

0.007 0.007 0-007 0.056 0.774 0.126 0.021

0 0 0 3 6 7 1

0.011 0-011 0-011 0.004 0.680 0.051 0.011

820 820 820 266 275 610 342

P~ < 0.015 714 P2 < 0.015 714 P3 < 0-078 176 P4 < 0.015 714 P5 < 0.015 714 P6 < 0-015 714 P7 < 0.363 118 P8 <- 0.597 074 P9 < 0.361 938 P10 < 0.024 790

0 0 0 + + 0 -+ + +

National Park

Boswellia serrata Grewia tiliaefolia Millusa tomentosa Tectona grandis Wrightia tinctoria Xeromphis spinosa X. uliginosa

< Pi < 0.025 < Pz -< 0.025 < P3 < 0-025 < P4 <- 0-108 _
820 820 820 333 924 789 542

--0 -+ + +

"n, Number of individuals sampled; D, number of individuals debarked; --, avoided; +, preferred; 0, debarking proportional to availability.

Axis axis switched over completely from grazing to browsing during this d r o u g h t period ( K h a n et al.,

a consequent shift in the predation pattern o f lions towards wild prey (Sinha, 1987) f r o m the situation described by Berwick (1974), H o d d (1969) and Joslin (1973). These improvements can be attributed to the major conservation measures taken in 1972 and thereafter, including the shifting from Gir o f 63 Maldhari nesses (pastoral settlements) along with their livestock population, total exclusion o f migratory cattle, declaration o f 258 k m 2 o f National Park, fire control, development o f water-holes and better protection measures. However, factors such as d r o u g h t would reverse this trend. It is clear that any policy formulated for Gir will have lions as the central theme, because o f their highly endangered status in the world. Considerable effort

1990). Better conditions o f grass cover in National Park, coupled with low chital density there, would have resulted in less competition for food for sambar and hence explain the lower incidence o f debarking. Mortality o f this magnitude in the plant populations o f Gir has several m a n a g e m e n t implications. The plant communities include a high p r o p o r t i o n o f species which exist at extremely low densities whose loss would greatly impoverish these communities. Gir Lion Sanctuary has made a remarkable recovery since 1967 in terms o f forage production, increase in herbivore and lion numbers ( K h a n et al., 1990) and

Table 5. Proportion available (Pio), proportion debarked (Pie) and 95% Bonferroni confidence limits for different girth classes in Sanctuary West and National Park a Girth classes (cm)

n

Pio

D

Pie

Confidence limits for Pie

Preference rating

Sanctuary West 0-20 21~,0 41~i0 61-80 81-100 101-120

228 280 231 87 30 6

0.264 0.324 0.267 0:100 0.034 0.006

5 8 9 9 8 9

111 84 23 13 7 1

0.464 0.351 0.096 0.054 0.029 0.004

4 4 2 3 2 1

0.379 0.270 0.046 0.015 0.000 0.006

555 -< Pl < 0.549 183 _
244 616 362 850 842 970

+ 0 --

National Park 0-20 2140 41~0 61-80 81-100 101-120

200 213 256 129 31 14

0.237 0.252 0.303 0.153 0-036 0.016

2 6 6 0 7 6

67 57 13 5 0 0

0.471 0.401 0.091 0.035 0-0 0.0

8 4 5 2

0.361 0.293 0.027 0.005

623 214 866 472

976 585 133 872

+ + ---

< Pl -< 0.581 < P2 < 0.509 < P3 < 0.155 < P4 < 0.075

0 0

o n, Number of individuals sampled; D, number of individuals debarked; --, avoided; +, preferred; O, debarking proportional to availability.

154

J. A. Khan, IV. A. Rodgers, A. J. T. Johnsingh, P. K. Mathur Table 6. Percentage of trees in different damage categories in Gir~

Plant species

Low

Medium

n

D

D

%

D

Boswellia serrata Cassia fistula Ehretia laevis Emblica officinalis Flaucortia indica Grewia tiliaefolia Millusa tomentosa Sapindus emarginatus Schrebera swietenioides Tectona grandis Wrightia tinctoria Xeromphis spinosa X. uliginosa

53 26 8 68 7 50 24 3 10 1 016 289 116 8

1 1 1 10 1 1 1 1 1 42 231 85 5

1 1 0 5 1 0 1 0 1 20 21 18 0

1.8 3.8 0.0 7.3 14.2 0 4.1 0 10.0 1.9 7.2 15.5 0-0

Total

1 678

381

69

4-1

147

Severe %

D

% Overall damage %

0 0

0 0

0 0

0 0

1

12.5

0

0

3 0 1 0 1 0 17 94 29 1

4.4 0 2.0 0 33.0 0 1-6 32.5 25-0 12.5

2 0 0 0 0 0 5 116 38 40

2.9 0 0 0 0 0 0.49 40.1 32.7 50.0

1.8 3.8 12.5 14.7 14.2 2.0 4.1 33.3 10.0 4.1 79.9 73.2 62-5

8.7

165

9.8

22.7

" n, Number of individuals sampled; D, number of individuals debarked. may be required to maintain the present level of wild herbivores, and manipulation of the vegetation to suit the herbivore populations. Management action required immediately includes weed control, fire management, patch cutting (Dinerstein, 1979) and further water-hole development. The ground cover in Sanctuary West and East is dominated by weeds such as Achyranthes aspera Linn., resulting from excessive cattle grazing which needs to be controlled to increase food availability for chital. This species would benefit considerably from patch cutting with adequate provision of water, especially inside National Park where the vegetation cover has increased significantly during the last 20 years, and where the density of chital is at present quite low (Khan et al., 1990). Changes in the overall composition of these communities will depend on how the different tree species regenerate. Differential regeneration suggests that the composition might change even if no management intervention takes place. Continuous accurate monitoring of ungulate densities in Gir is therefore essential to evaluate the impact of changes in plant composition and cover. The damage from debarking is not at present a serious concern for managers, since severe damage is restricted to only three species, but these should be monitored during further studies. ACKNOWLEDGEMENTS We wish to express our sincere thanks to Mr H. S. Panwar, Director, Wildlife Institute of India, for providing us with the logistic support to carry out research in Gir. Thanks are also due to Sanat Chavan,

Ashok Kumar, S. K. Pant, Bharat Pathak and Uday Vohra for providing valuable suggestions and help. We would also like to thank Ajith Kumar for his help in data analysis and Rashid and Salim in manuscript preparation.

REFERENCES

Berwick, S. H. (1974). The community of wild ruminants in Gir Forest. PhD dissertation, Yale University. Brander, A. A. D. (1923). Wild animals in Central India. Edward Arnold, London. Byers, C. R., Steinhorst, R. K. & Krausman, P. R. (1984). Clarification of a technique for analysis of utilizationavailability data. J. Wildl. Manage., 48, 1050-3. Champion, H. G. & Seth, S. K. (1968). A revised survey of the forest types of India. Manager of Publications, Government of India, New Delhi. Dinerstein, E. (1979). An ecological survey of the Royal Karnali Bardia Wildlife Reserve, Nepal, Part II. Habitat/animal interactions. Biol. Conserv., 16, 265-99. Hodd, K. T. B. (1969). The ecological impact of domestic stock on the Gir Forest. IUCN Tech Mtg, l lth, 259455. Joslin, P. (1973). The asiatic lion: a study of ecology and behaviour. PhD thesis, University of Edinburgh. Khan, J. A., Rodgers, W. A., Johnsingh, A. J. T. & Mathur, P. K. (1990). Gir Lion Project. Ungulate-Habitat ecology in Gir. Project Report, Wildlife Institute of India, Dehra Dun. Neu, C. W., Byers, C. R. & Peek, J. M. (1974). A technique for analysis of utilization-availability data. J. Wildl. Manage., 38, 541-5. Patel, R. I. (1984). Forest flora of Gujarat State. Forest Department, Gujarat State, Baroda. Schaller, G. B. (1967). The deer and the tiger. University of Chicago Press, Chicago. Sinha, S. P. (1987). The ecology of wildlife with special reference to Gir lion in Gir Lion Sanctuary and National Park. PhD thesis, Saurashtra University, Rajkot.