Namgail 2009

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1hesis committee
Superisor
Pro. dr. l.l.1. Prins
Proessor o Resource Lcology
\ageningen Uniersity
Co-superisor
Dr. S.L. an \ieren
Associate proessor, Resource Lcology Group
\ageningen Uniersity
Other members
Pro. dr. G.R. de Snoo, \ageningen Uniersity
Dr. B.A. Nolet, NIOO-KNA\, Maarssen
Dr.ir. l.l. de Iongh, Leiden Uniersity
Dr. L.S. Bakker, NIOO-KNA\, Maarssen

1his research was conducted under the auspices o the Graduate School o Production
Lcology and Resource Conseration
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1sewang Namgail
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Submitted in partial ulillment o the requirements or the degree o doctor
at \ageningen Uniersity
by the authority o the Rector Magniicus
Pro. dr. M.J. Krop
in the presence o the
1hesis Committee appointed by the Doctorate Board
to be deended in public
on 1uesday 1 Noember 2009
at 13:30 in the Aula, \ageningen

1sewang Namgail
Geography o mammalian herbiores in the Indian 1rans-limalaya: patterns and
processes. Pages: 122.

1hesis, \ageningen Uniersity, \ageningen, NL ,2009,
\ith reerences and summaries in Dutch and Lnglish

ISBN 98-90-8585-524-8
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Chapter 1. lerbiore species richness: Introduction 1
Chapter 2. Aboeground biomass and phytodiersity along altitudinal 19

gradients at dierent spatial scales in the Ladakh 1rans-limalaya
1sewang Namgail, Gopal S. Rawat, Charudutt Mishra, Sipke L. an \ieren
& lerbert l. 1. Prins,

Chapter 3. Biogeography o mammalian herbiores in Ladakh: distribution 35
in relation to enironmental and geographical barriers
1sewang Namgail, Sipke L. an \ieren & lerbert l. 1. Prins ,in reiew,

Chapter 4. \inter habitat partitioning between Asiatic ibex and blue 5
sheep in Ladakh, Northern India
1sewang Namgail ,2006,. Journal o Mountain Lcology, 8: -13.

Chapter 5. Lects o herbiore species richness on blue sheep's niche dynamics 6
and distribution in the 1rans-limalaya
1sewang Namgail, Charudutt Mishra, Christine B. de Jong, Sipke L. an \ieren
& lerbert l. 1. Prins ,2009,. Diersity and Distributions, 15: 940-94.

Chapter 6.Coexistence o Ladakh urial and blue sheep at multi-spatial scales in 81
the 1rans-limalayan mountains
1sewang Namgail, Sipke L. an \ieren, Charudutt Mishra & lerbert l. 1. Prins

Chapter . Mammalian herbiore species-richness: Synthesis 95

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Lcology is one o the youngest and least settled branches o biology. Larnest analytical
inestigations in ecology started only in the 1960`s with Robert MacArthur working on
the Lotka-Volterra`s model. Seeral ecologists including MacArthur were startled by the
uneen distributional pattern o biodiersity on earth. 1his pattern, although described
by naturalists long beore ecology as a science was conceied, remains to be understood
thoroughly. Larly ecologists adopted a reductionist approach to look at biodiersity
pattern on the planet. But that changed in the last couple o decades with ecologists
adopting a more pluralistic and macro-ecological approach. 1his thesis is a step into that
direction.
It became apparent in the last ew decades that species distributional patterns are
dynamic and depend on the coniguration o resources in the enironment. Species
colonize areas, exploit resources and expand their populations, proided the resources
are abundant and their birth rates exceed death rates. 1hus, animal distributional ranges
change through space and time. I we look at this dynamicity in the animal kingdom,
nothing can beat the rate at which man`s distribution changed in the early phases o
human history. lor instance, Luropeans inaded the Polynesian islands, depleted
resources and let or other archipelagoes within a decade. But human distribution
became more static once we resorted to agriculture, which enabled production o surplus
ood in a particular area.
1he settled lie and mass production o ood also allowed us to indulge in
recreational actiities. Consequently, human needs and aspirations hae sky-rocketed.
1oday, when we think about the lutchinsonian niche hyperolume, no species beit this
more than we do. 1he list o our resource requirements is growing exponentially with
technological deelopments, because it is associated not only with surial but also with
anity. So today when we ask what is the single most important actor determining
human distribution, or some lanky and scrawny models walking down the ashion-
ramps o Paris, it deinitely will not be ood, but cosmetics like lipsticks, mascara, nail-
polish, so on and so orth. 1hese can be produced in one corner o the world and get
transported to another within a short time, thereby precluding the need to redistribute
human population according to the aailability o these resources.
Non-human animal distribution howeer remains luid een today, depending on
aailability o ood, water and shelter. 1heir distributions thus depend on the presence o
these resources, and gien the resource state and biological actors o an area, we can
predict their distribution in that area. But things get complicated and murky once one
asks how animal assemblages are ormed and maintained through time. lere one
simultaneously need to deal with distribution o more than one species, and the niche
space o an animal gets additional dimensions such as sympatric competitors, predators
and parasites eeding on them. Moreoer, we need to integrate the eolutionary history
o the constituent species o an assemblage.
1hereore, enironmental actors, eolutionary history, dispersal, competition,
acilitation, predation and parasitism are some o the most important actors determining
the ormation and maintenance o animal assemblages. Neertheless, gien that some
species surie in oreign terrain, once introduced there, enironmental actors seem less
important, suggesting that geographical barriers or competitors had stopped them rom
adancing into those areas. lurthermore, when we talk o the biotic enironment o a
species, herbiores orm a unique taxon whose distribution is limited not only by ood
resources but also by predators at higher trophic leels, thereby urther complicating the
issue. So what are the most important biotic and abiotic actors that determine the
ormation and maintenance o herbiore assemblages I plunged headlong into a enture
our years ago with this ery question in mind. I chose the mountainous rangelands o
ii
the Indian 1rans-limalaya to address this question, and at the end o the project, I hae
emerged with scores o new questions while trying to ind an answer. lurthermore, I
lash out banners with answers like it depends on the taxon under inestigation, spatial
and temporal scales. In any case, i I managed to spark an interest amongst ecologists on
the ascinating assemblage o mammalian herbiores in the 1rans-limalaya, and thereby
stimulate urther research, I would hae accomplished an important goal.

iii
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Namgail, 1. ,2009,. Geography o mammalian herbiores in the Indian 1rans-limalaya:
patterns and processes.

1he loss o mammalian herbiores rom grazing ecosystems has become a major
concern, and eorts to stem such losses are stymied by lack o inormation on the
proximate and ultimate actors inluencing their distributions and diersity patterns. 1his
research inestigated the distribution, species-richness patterns and underlying
mechanisms in mammalian herbiores o the 1rans-limalayan region o Ladakh, India.
It adopted a multi-spatial approach to understand these issues in the little-known
herbiore assemblage o the region. Since egetation is the most important actor that
determines the distribution o herbiores, irst I researched the distribution and
abundance patterns o ascular plants along an altitudinal gradient at dierent spatial
scales. Both plant species-richness and aboeground biomass showed a hump-shaped
relationship with altitude. Such a relationship in case o species-richness is expected, but
it is contrary to my expectation o a negatie linear relationship, in case o abundance. I
relate this unexpected pattern to the limited precipitation and perasie liestock grazing
at lower altitudes in this dry alpine enironment. I then inestigated the biogeography o
mammalian herbiores, and ound that they orm geographical groups on the basis o
their eolutionary histories. Subsequently, I assessed the niche relationship between
Asiatic ibex &DSUD LEH[ VLEHULFD and blue sheep 3VHXGRLV QD\DXU the most common large
herbiores in Ladakh, to see whether local leel processes like competition generate
spatial pattern o herbiore species-richness. 1he results showed that blue sheep
constrains the distribution o ibex, which implies that competition amongst natie
species does play a role in structuring large herbiore assemblages in the region.
Recognising the lack o inormation on large herbiores` niche ariation across
assemblages, I also studied blue sheep`s niche width in relation to herbiore species-
richness. It became apparent that the species` niche aries across assemblages with
dierent number o sympatric species, which could negatiely inluence the animal`s
reproductie perormance and population. linally, I asked i the distributional range o
the endangered Ladakh urial 2YLV YLJQHL YLJQHL is constrained by the abundant blue sheep,
and ound that these two species associate randomly at large geographical scales, but co-
occur at the landscape leel as a result o local habitat-leel resource partitioning. 1hese
results contribute towards understanding the mechanisms responsible or the ormation
and maintenance o large herbiore assemblages in the 1rans-limalaya and other
mountainous regions o the world.

.H\ ZRUGV Large herbiores, mammalian herbiores, mountain ungulates,
biogeography, chorotype, species richness, diersity, biome, ecotone, Ladakh urial, blue
sheep, Asiatic ibex, marmot, hare, pika, ole, ascular plants, altitudinal gradient

iv
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Lery research project ends with lot o memories, and this is the section to reie them
all, and express gratitude to those who made them memorable. lour years ago, I set out
on a quest o understanding herbiore diersity in the 1rans-limalaya. 1he journey was
rather tortuous that needed assistance at eery turn. lere I thank all those who helped
me, making the journey smooth and enjoyable. \hile carrying out this research, I had the
priilege o meeting some o the brightest and dynamic people I hae eer known.
Constraint o space would not allow me to list them all. Some o them contributed
immensely, and here I make a humble eort to express my gratitude to them in words.
My research carrier started in 199, when I worked on insects pests o poplar
trees under the superision o Pro. P.K. 1ewari at the Panjab Uniersity, Chandigarh. I
thank her or getting me interested in research. 1hen I shited my ocus on bigger
animals: large mammals. I was ortunate to get an opportunity to work with Dr. Joe lox
in Norway. Joe you gae me a break when I needed one desperately. \ou led me through
murky waters, and gae resh impetus that enabled me to do a Ph.D. \ours is one o
those contributions that words would be too eeble to express gratitude or. 1hank you
so much in any case. Dr. \ash Veer Bhatnagar is another person that played a crucial role
in my early research carrier. \ash Veer, I still remember the time when you gited me a
copy o the Mountain Monarchs`, which I cherish to this day. 1he earthwatch days` in
Ladakh were absolute un. 1hanks so much or all your support, guidance and
encouragement oer the years.
1he discourse or a PhD started with a trip to \ageningen in 2001. No, it was
neither the tulips nor the windmills, not een the cheese that brought me to the
Netherlands, but the erudite insight o Pro. lerbert Prins that brought me here.
lerbert, things didn`t start the way we wanted, but start it did, and I consider mysel
lucky to get this opportunity to work with you and the group. \ours has been a
contribution as big as the limalaya, so the word o gratitude has to be equally big, and I
don`t know i exists in the lexicon. \ou taught me how to turn a question on its head and
look on the other side o the picture. I am glad that you guided me away rom the species
chauinist approach. Moreoer, your intellectual insight, expert guidance and humour
made this journey ar easier than expected. I cherish our conersations on the mountain
trails and at campsites oer tea. 1hanks or being a constant source o inspiration.
My other superisor is Dr. Sip an \ieren, who apart rom intellectual input
taught me how to surie in the Netherlands. Sip, your contribution has been immense
at eery stage, right rom the proposal writing to the time o acknowledgement writing.
\ou readily soled all problems, be it inancial or logistical, and put a smile on my ace in
diicult times. 1hanks or listening patiently to all my petty questions, or belieing in me
and letting me do the things that I wanted, although you made sure that I inished in
time. I enjoyed working with you on desk as well as in the ield. 1ogether we enjoyed the
icy heights o the 1rans-limalaya and mudlats o the Netherlands. Both you and
lerbert made the last our years enjoyable and memorable. 1hank you both or
supporting me all through my ups and downs.
Dr. Charudutt Mishra is another person who made a huge contribution to this
research project, not only in terms o academic input but also logistical and inancial. le
has been a great riend and mentor through all these years. Charu, irst o all, thanks or
initing me to Mysore. It was great to work with you and others at NCl. 1hanks or
your guidance and the stimulating discussions that went a long way in urthering my
research carrier. \ou went out o your way and helped me in raising additional unds.
1hanks so much or eerything.
I also thank my riend and colleague Dr. Sumanta Bagchi, who was a great
company in the ield. \orking with you was a joy. 1hanks or all the scholarly
v
discussion and collaboration. 1he competition in the canas tent at 400 m was un! I
also enjoyed my time in the ield with Dr. Prana 1riedi. 1hanks Prana or all the
support and encouragement. \our humorous and jolly nature always put a smile on my
ace, although we didn`t spend much time together. It was un to watch you motiating
young wildlie aspirants. Look orward to more un in the mountains!
During this research, I spent a considerable amount o time at the \ildlie
Institute o India, and seeral people made my stay enjoyable and ruitul. I especially
thank Dr. G.S. Rawat or identiying the plants and helping me in acquiring permission
to use the laboratory acilities. I also thank P.R. Sinha, the Director, or allowing me to
use the library. I thank Dr. Karthik Vasudean or the memorable time in the ield as
well as at the institute. 1hanks or all the goodwill and encouragement oer the years. I
also thank Suresh Kumar or all the help and discussions. Suresh and Manju also made
my stay in the south enjoyable, thanks and loe to you both. I also thank Dr. S.
Sathyakumar or helping me during this and preious research projects.
During this research, I was ortunate to get a chance to work with Pro. \oram
\om-1o rom the 1el Ai Uniersity, Israel. \oram, thanks much or introducing me
to ornithology. It was a joy to work with you, and look orward to collaborating urther. I
also receied considerable support rom Dr. Otto Pister, who readily shared his
knowledge o the birds and mammals o Ladakh. 1hanks Otto or writing
recommendation letters to support my grant proposals. I also thank Dr. lenry Noltie
rom the Royal Botanic Garden, Ldinburgh, I beneitted rom his ast knowledge o
limalayan plants. 1hanks are also due to Pro. Rory Putman and an anonymous reiewer
or their comments on the research proposal, which helped in designing the study.
I thank Dr. George Schaller or being a constant source o inspiration. le readily
agreed to be an unoicial adisor, and I immensely beneitted rom arious discussions
in the ield. 1hanks George or sending all the reerence materials oer the years. I thank
Dr. Asad Rahmani, Director Bombay Natural listory Society, or all the support and
encouragement. I am grateul to Dr. 1om McCarthy at the International Snow Leopard
1rust or all the support. I also thank Dr. A.J.1. Johnsingh or inspiring. I sincerely thank
Mr. A.K. Sriastaa, Mr. Jigmet 1akpa, Mr. 1ariq Shawl, Mr. Salim Ul-laq, Dr. C.M.
Seth and all the Rangers rom the Department o \ildlie Protection, Ladakh or
permitting me to work in arious protected areas and proiding logistical support,
wheneer I needed.
I extend my thanks to all the organisations that unded my research in Ladakh.
1he most important ones are the \ildlie Conseration Society, Ruord Small Grants
loundation, International Snow Leopard 1rust, \idlie Protection Society o India, lord
loundation, Nature Conseration loundation, India, \ageningen Uniersity, the
Netherlands and the Uniersity o 1romso, Norway. I thank all the sta in these
organisations that helped me in one way or another.
Apart rom academic, logistical and inancial, one also needs adequate emotional
and moral support to complete a research project, and I am lucky to hae amazing
amily, riends and relaties, who always stood by me in diicult times. lirst o all, I
thank my parents, brothers and sisters or accepting my nomadic lie, and belieing that
what I am doing is good or me as well as or the society. All o you hae been ery
supportie. Demise o Aaba Phuntsog and Aama Zangmo created a huge acuum in my
lie. It was diicult to come to terms with their demise. 1hey had nursed and groomed
me, and I eel extremely sorry that my nomadic liestyle didn`t allow me to spend time
with them when they needed me.
I thank Ven. Sonam Kunga or all his support and encouragement oer the years.
I am extremely thankul to Skarma Namtak or all his care and concern about my
personal as well as proessional lie. 1hanks Skarma, you helped and supported me all
vi
these years. 1hanks or all your beneolent assistance. I also thank all my aunts who hae
always been concerned about my wellbeing, eagerly waiting or me to end the nomadic
lie. But little do they know that this liestyle gies me joy, not all the time though!
I am ortunate to hae a home away rom home. Paul and Rigmor are my
parents, riends and mentors. 1hey always made sure that I elt at home, here in the
Netherlands and in Norway. Paul and Rigmor, I don`t know how to thank you
adequately. \ou gae me strength and conidence when I lacked them. 1usen takk or all
the warmth, care and loe. I also thank Odd and Lllinor or their care, loe and
hospitality. I`ll neer orget the memorable time spent with you at Malsale. I also thank
John and lelga or all the support and encouragement.
JK and Shi always made me eel at home wheneer I isited Chandigarh. It was
a joy to be with you. I enjoyed going with you to the obscure places in limachal Pradesh
reporting on bizzare rituals. I tried my best to stop wheneer I was in Delhi, although
you always kept on complaining that I don`t isit. 1hanks much and loe to you both, lie
without you would hae been quite boring. I miss Athar, and hope that the Chachu`
will be with him soon. I thank my other riends, Maj. R. Bisht and Maj. S. Dhadwal.
Bishta and Dhada, time spent with you couldn`t hae been more exciting. 1hanks or
sharing an interest to sere the country.
I thank Sanji Gorka or all the good times we had together, both in the ield and
in Bangalore. 1hanks Sanji or accommodating wheneer I dropped by. \ou listened
patiently and soled all my computer related problems. \ou salaged important data on
seeral occasions. I also thank Sarika or all the good times in Chandigarh. I thank
Nawang, Gitanjali, Sara, Swati and Gopi or their help and hospitality in Delhi. I also
thank Bindu or sharing interests and good company in the ield. I am thankul to Chris
or all the support and or remaining as a good riend.
Dr. Kyle 1omlinson and Miss Jia \u made my stay here in the Netherlands more
memorable. 1hank you both or help and or making my time at oice and home
pleasant. 1he rierside picnics were antabulous. I express heartelt thank to Mrs. Aartje
Van Strijland Mum` or all the care, loe, aection and the innumerable cups o coee
and bowls o soup. Mum, I really enjoyed my stay in your home, which really is a loely
place with wonderul neighbours and serene enironment. 1hanks to larry, Chris and
lenrriete or making it more interesting.
I enjoyed my time spent in the company o Paul and Gisella. 1hank you or
initing me to your parents` place. It was a wonderul trip, biking up and down the
undulating terrain o Limburg. Lie without you would hae had been less interesting at
Borneseesteg. 1hanks or all the help and support, and or listening to my stories. lope
Borneys` remains open longer so that we can guzzle some more beer. I thank lendrik
Sardjoe ,1oko, and his amily or all the support. lendrik thanks or the innumerable
cups o tea and the mobile in need. I also thank lrederique or all the help and support. I
also thank her parents and brother or initing me to the south and showing me the
Dutch countryside.
I am lucky to be associated with an amazing bunch o people at Nature
Conseration loundation, Mysore. I especially thank Madhu and Sridhar or all their help
and suggestions. 1hanks to Aparajita or the innumerable discussions, sumptuous meals
and o course booze. I also thank Diya or all the help and encouragement and or the
wonderul time in ield. I also thank Rohan or all the help and support during my stay in
Mysore. I am grateul to Raghuath or helping with all the map works. I thank Rakhee,
Vena, 1anuja, Nain and Mr. Vishwanath or helping me with paper work. 1hank you all
or ably soling eery glitch. A special thank to Suma or the innumerable cups o tea,
coee and buttermilk plus that unathomable smile on the ace. My stay at Mysore was
vii
made special and memorable by Aaron, Ambika, Ayesha, Riyaz, Rishi, Rohit, Raji,
Anand, Chandrima, Paithra, Coralie, Sumetro, Neema, Krupakar-Senani, Rana,
Narayan, Debapriyo and Kein. I thank you all or the great company.
At Resource Lcology Group in \ageningen, I thank lermann, Gerda and
Patricia or all their help at arious stages o the project. I beneitted immensely rom
discussions with Ignas, lred and lrank. 1hank you all or helping me, especially during
the proposal-writing phase. Christine de Jong readily agreed to analyse aecal pellets o
my study animals. 1hanks much Christine, it was o great help. \illemian Schouten was
o great help in the beginning, so I record here my thanks to her too.
I thank Daniel or translating the summary o the thesis into Dutch. Dank u! I
truly appreciate Arend`s gesture o lending his old bike when I trudged around. 1hanks
Arend. I oice my thanks to Ral, Lrika and her parents or showing me the stunning
Zeeland. I also record my thanks to Luridice or her great company in the beginning. I
thank Andrew, Steen, Ron, Milena, Pim and 1om or all the adices and support.
1hanks Andrew or considering to collaborate on a study on snow leopards, although it
did not materialize due to unoreseen circumstances, it did in a way acilitate the initiation
o the present work. I gained a lot rom discussions with Geerten, Nicole, Jasja, 1homas,
Mariaan, Anne, Ral, Rudy, Daniel, Robert, 1om, Kyle, Jasper, lenjo, \olanda, larshid,
Anil, Lduardo, Priya, Vincent, Ldward, Audri, Cornelis, Xaier, Lmanuel, Benson and
1essema. 1hank you all. I especially enjoyed the reewheeling talks during the breaks.
I express my gratitude to Claudius an de Vijer or extending all possible
support during arious phases o the research. 1hanks Claudius or all the aours and
beneolence. I also thank Marion Rodenburg at the CSA or helping me with the isa
and residence permit related issues. 1hanks Marion or replying to all my mails, and
making sure that things went in the right direction. I thank Simon Delany at the \etland
International or all his support and goodwill.
Back in Ladakh, I thank all my riends: Sonam Joldan, Rigzin Sandup, Nawang
Dorjey, Skarma, Dorjey Cheetah and Phuntsog Paldan or all the un, wheneer I was in
Ladakh. I also thank Dorjey Angchuk, 1sewang Dorjey, Jorphel and 1ashi Mahe or
their great company and hospitality in lanle during the ield work. I also thank all other
workers at the Indian Astrophysics Obseratory, lanle or their care and hospitality. It
was a truly amazing stay there with all modern amenities in the middle o nowhere. I
thank 1ashi Morup or all the great discussions and or sharing his inormation on
Changthang. I also thank Rinchen \angchuk, Nisa Khatoon and Pankaj Chandan or
their help and support in Ladakh. I am indebted to sta at the Snow Leopard 1rails and
\ama 1reks or organising arious expeditions.
I thank our ield coordinator Karma Sonam in Ladakh. le made inaluable
contribution to this project. le helped not only in the ield but also in entering data.
1hanks Karma or being so supportie. I also thank Mark and Carola, M.Sc. students or
their help and company in the ield. I thank Sushil Dorje, 1hinles, Sheru, Sherpa,
Chuldim and arious others who helped a lot during the study on blue sheep in Spiti.
1hank you all. I`ll be back one day to rock some more with lute and harmonica. Various
other people helped me in the ield in Ladakh and I thank them all, especially 1sewang
Morup, Mipham, Namgyal, Lobzang, 1hinles-I, 1ashi Gyatso, 1hinles-II, Gialson and
Morup. \a julley!

1sewang Namgail
\ageningen, 2009

CHAP1LR J

Herbivore species-richness: introduction

1sewang Namgail

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Anonymous

Resource Lcology Group, Department o Lnironmental Sciences, \ageningen
Uniersity, Droeendaalsesteeg 3a, 608 PB \ageningen, 1he Netherlands
&
Nature Conseration loundation, 306,5 IV-Cross, Gokulam Park, Mysore-50002,
Karnataka, India

Chapter 1
2
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Biodiersity on earth is uneen with some biogeographic units harbouring more species
than others, a pattern obsered long ago by naturalists ,looker, 1853, Sclater, 1858,
\allace, 186,. 1he most prominent o this discrepancy in species richness being
exempliied by the latitudinal gradient in species diersity with more species in the tropics
and less in the temperate and polar regions ,Pianka, 1966, Rohde, 1992,. \ithin these
broad geographical zones, some biomes support more and some less number o species.
1hereore, one obious question is what determines such spatial pattern o species
diersity. 1o answer such questions, studies hae been conducted on dierent taxa at
seeral spatial and temporal scales. 1hese studies hae indicated that actors such as
competition ,1ilman, 1994,, habitat diersity ,Shmida & \ilson, 1985, 1ews et al., 2004,,
time-or-speciation or eolutionary time ,Stephens & \iens, 2003,, area ,Prins & Ol,
1998, Losos & Schluter, 2000,, spatial scaling o resource use by species ,Ritchie & Ol,
1999,, climatic oscillations ,Dynesius & Jansson, 2000,, geometric constraints ,Colwell &
Lees, 2000,, optimal precipitation,soil ertility ,Ol et al., 2002, and seeral other
parameters ,Pianka, 1983, may inluence species richness across the planet.
Despite such research eorts, hitherto there is no consensus on the primary
mechanisms ,lawkins et al., 2003,. It, howeer, became apparent that species assembly
patterns hinge on the spatial scale o inestigation, as dierent actors play important
roles at dierent scales ,Lein, 2000, \illis & \hittaker, 2002, Chase & Leibold, 2003,
Rahbek, 2005,. lor example, climatic and historical actors play a major role in
determining species richness at continental scales ,\hittaker et al., 2001,, while
interspeciic competition and habitat structure become more important at local and
regional scales ,luston, 1999, Amarasekare, 2003, 1ews et al., 2004,. 1hus, space has
been considered to be piotal to many ecological processes.
1he studies on spatial pattern o biodiersity are howeer biased towards birds,
ish, insects and plants, and remains little known or large herbiores ,but see Prins &
Ol, 1998, Ol et al., 2002, Klop & Prins, 2008, Cromsigt et al., 2009a, Cromsigt et al.,
2009b,. Large herbiores are represented by a wide ariety o taxa, and occupy a range o
ecosystems across the planet. 1hey inluence the structure and unctioning o ecosystems
by altering the dynamics o plant communities ,Naeem & Li, 199, Augustine &
McNaughton, 1998, Ritchie & Ol, 1999,. Large herbiores also constitute important
prey o large carniores, thereby maintaining species diersity at higher trophic leels.
1hus, large herbiores play an important role in ecosystem unctioning, which proides
compelling reasons to study diersity patterns o large herbiores that will help in
maintaining their diersity in grazing ecosystems.
In understanding species-richness pattern o an area, the primary question that
needs to be answered is, do local assemblages represent smaller subsets o the regional
species pool I so, what processes determine such non-random distribution o co-
occurrences \hy do some areas contain more species than others do Are there speciic
determinants o these patterns Larly ecologists had come to the conclusion that species
diersity o a region relected its age ,Pianka, 1983,, and they did not clearly distinguish
between regional and local diersity, but more recently ecologists partitioned the regional
rom the local diersity ,Rickles & Schluter, 1993,, and took on diersity as an ecological
problem that they could sole by resorting to insights rom population biology.
loweer, to understand the species diersity at a global scale, ecologists need to abandon
the parochial iew o local determinism and recognise that ecology, eolution, geography
and history are dierent acets o a single set o processes and the patterns they generate
,Brown, 1985,.
Introduction
3
MacArthur ,192, proposed a simple model to explain species richness D. as a unction
o resource aailability and niche relationships among species: D. ~ Dr,Dv;;1-C),
where Dr represents the diersity o resources, Dv the niche width o each species, C
describes the number o potential competitors in niche space, and is the mean
competitie coeicient or mean niche oerlap. 1his model predicts that assemblages
dier in species richness in three dierent ways, ,1, an area with greater resource
diersity ,larger Dr, can maintain more niches and thereore contain higher number o
species ,lig. 1b,, ,2,
assemblages where
species exploit smaller
portions o the total
niche space ,smaller
Dv, contain more
species ,lig. 1c,, and
,3, greater oerlap o
niches ,larger o,
acilitates co-existence
o more species ,lig.
1d,. 1his model can
sere as a useul
starting point in
understanding species
richness in local and
regional assemblages o
large herbiores
lig. 1. A simple model

o species richness, D
each species uses a
portion v o the
aailable resources ,R,,
oerlapping with
adjacent species by an
amount o. More species may occur in one assemblage than another because E a greater
range o resources is present ,larger R,, because F species are specialized ,smaller
aerage v,, G greater oerlap in resource use among species ,larger aerage o,, ,ligure
adopted rom MacArthur, 192,.

1LFKHWKHRU\DQGKHUELYRUHGLYHUVLW\
Although large herbiore diersity on a continental,global scale may be determined by
climatic actors ,Ol et al., 2002,, biotic interactions ,Prins & Ol, 1998, and habitat
eatures ,Ben-Shahar & Skinner, 1988, become more important at local and regional
scales. lacilitation and competition are important biotic actors potentially goerning
herbiore species richness ,Arsenault & Owen-Smith, 2002,. 1he ormer may enhance
species richness in communities in high productie areas in the 1ropical systems
,luisman & Ol, 1998,, but perhaps is less important in structuring herbiore
assemblages in less productie ecosystems where competition tends to be the dominant
orm o interaction ,Mishra et al., 2002, Mishra et al., 2004, Namgail et al., 200b,.
loweer, large herbiores in these ecosystems may aoid strong competition by
dierging in their diet and,or habitat use ,see Namgail et al., 2004,, as these are the most
important resource dimensions along which species segregate in order to coexist
R
More species as a result of higher specialization (smaller u)
More species as a result of greater overlap (larger o)
More species as a result of greater resource range (larger R)
R
(a)
(b)
(c)
(d)
Chapter 1
4
,Schoener, 194,. 1hus, biotic actors are more important in the more productie
tropical ecosystems, while the physical enironment plays a signiicant role in the
deelopment and maintenance o communities in less productie ecosystems toward the
poles ,MacArthur, 192,.
A plethora o studies on niche separation between large herbiores hae been
carried out ,e.g., Jenkins & \right, 1988, Voeten & Prins, 1999, Namgail et al., 2004,.
Nonetheless, the most crucial actors determining assembly patterns remain elusie. Most
o the preious studies were conducted in single assemblages, and it is not known as to
how the niche o a species can ary across herbiore assemblages with arying number o
sympatric species. Based on niche theory, one might expect the habitat-niche width o a
species to decline with the number o sympatric species during summer ,lig. 2,, and the
diet-niche width to increase with species richness due to aailability o a greater range o
plant species in high altitude drier enironments during this growth season. In contrast,
both diet and habitat widths are expected to decline with number o sympatric species in
an assemblage during winter due to plant senescence and snow coer ,lig. 2,.

,a, ,b,

lig. 2. A conceptual model o the expected relationship between niche ,diet and habitat,
width and sympatric herbiore species-richness during summer ,a, and winter ,b,.

lerbiore diersity, at local scale, is a unction o the number o niches ,Rosenzweig,
1995,, and competitie interactions amongst sympatric species promote local species
richness through modiying niche relations ,Amarasekare, 2003,. 1his implies that
energetic constraints set a limit to the number o species that an area can support
,MacArthur, 192, Ol & Ritchie, 1998, Ritchie & Ol, 1999,. Such limitations allow the
coexistence o only those species, which exhibit trade-os in resource use in response to
competition ,Chase & Leibold, 2003, Dayan & Simberlo, 2005,. In other words, co-
occurring species are superior in exploiting alternatie resources ,Pianka, 1983,, and
diergence along any one niche dimension promotes species coexistence and thus
richness in a community ,MacArthur & Leins, 196, Kneitel & Chase, 2004,.
1he oerarching role o competition in structuring ecological communities has,
howeer, been disputed in the recent years, especially in the wake o dierential
responses o species to enironmental gradients ,\iens, 19, Strong et al., 1984,
Putman, 1996,. It has been suggested that abiotic actors play an equally important role in
organising ecological communities ,Dunson & 1rais, 1991,. In any case, recognising the
importance o both biotic and abiotic actors in organising species assemblages,
contemporary ecologists hae adopted a more pluralistic iew ,Schoener, 1986, Dunson
& 1rais, 1991,.
Introduction
5
lood and habitat are the most important axes along which species segregate ,Schoener,
194,. 1he dierential use by species along these niche axes is associated with dierences
in body size among herbiores ,Prins & Ol, 1998,. Because larger herbiores eat diets
o lower nutritional alue, their nutritional requirement per unit body mass is lower than
that o smaller herbiores ,lanley, 1982, Owen-Smith, 1988,. On the other hand, smaller
herbiores can orage selectiely due to their narrower muzzles ,Jarman, 194, Drescher
et al., 2006,. Non-ruminants can consume relatiely lower-quality bulk- orage, while
ruminants are able to extract nutrients rom intermediate-quality diets, due to their
eiciency in digesting ibrous plants ,Van \ieren, 1996,. Such allometric relationships
and physiological and,or morphological adaptations lead to dierential use o diets ,e.g.,
plant communities, plant species and plant parts, among large herbiores ,Jarman, 194,.
1his type o niche dierentiation is possible in high-productiity enironments,
where range o resources ,'U in MacArthur`s model, is larger, but is perhaps less
important in areas o lower plant productiity and diersity ,smaller 'U,. In the latter
situation, species may need to strike a balance between the need to maintain a suicient
intake rate or growth and reproduction, and the need to partition resources to enable
coexistence with potentially competing species. In such systems, one can expect a
relatiely greater dietary oerlap ,R in MacArthur`s model, amongst herbiores, this is
established by recent studies in the less productie ecosystem o 1rans-limalaya ,larris
& Miller, 1995, Bagchi et al., 2004, Mishra et al., 2004,. At the same time, high dietary
oerlap and resource limitation make competition an important interaction among
species in areas with smaller 'U ,De Boer & Prins, 1990,, and this again is supported by
recent studies ocusing on liestock-wild herbiore relationships in the 1rans-limalaya
,Mishra et al., 2002, Bagchi et al., 2004, Mishra et al., 2004,.
Apart rom the biotic enironment, the physical habitat also inluences the
distribution o large herbiores as alluded to earlier ,Ben-Shahar & Skinner, 1988,. In act
broad-scale distributional patterns o large herbiores are determined mainly by abiotic
actors, which act as constraints within which biotic mechanisms operate ,Bailey et al.,
1996,. 1hereore, dierences in large herbiore distributions in response to heterogeneity
in physical enironment may also lead to resource partitioning, and acilitate coexistence
o sympatric species. Physical enironment become important especially in mountainous
regions where the interaction between altitudinal gradients and topographic eatures
enhances the landscape heterogeneity, which proide multiple niche axes or resource
partitioning by large herbiores ,Schaller, 1998, Geist, 1999,. lor example, large
herbiores use dierent anti-predator habitats, which acilitate their co-existence in the
same area ,Lingle, 2002, Namgail et al., 2004,. 1hus, assemblages in areas with dierse
habitats may contain a higher number o large herbiore species with dierent habitat
requirements.

0HWDSRSXODWLRQWKHRU\DQGKHUELYRUHGLYHUVLW\
Animal populations are spatially structured, especially in patchy enironments, where
species hae discrete populations in habitat islands. Under such scenarios, certain
populations hae better reproductie perormance, and act as source populations that
maintain metapopulation dynamics. Ler since its conception ,Leins, 1969,,
metapopulation theory played a signiicant role in understanding population dynamics o
animals. Metapopulation theory is especially releant in understanding population
dynamics at the edge o a species range, where populations o species tend to be more
ragmented due to unsuitability o the enironmental actors ,Brown, 1984,. 1hereore,
the presence-absence o a species in an area essentially relects the balance between
colonization and extinction ,MacArthur & \ilson, 196, Rickles, 198, luston, 199,.
Chapter 1
6
Lnironmental conditions generally closely correspond to a species` ecological
requirements at the centre o its range ,Brown, 1984,. 1owards the periphery o its range,
the enironmental suitability or a species is expected to decline. 1hus, the probability o
occurrence o a species ,i.e., density, at regional scales is expected to decline rom the
centre o its range to the periphery due to decline in enironmental suitability as well as
its recolonization potential ,MacArthur, 192,. Gien this underlying spatial gradient in
actors inluencing species population density, dierent species in an assemblage,
constrained by their respectie biogeographic histories, perhaps respond dierently to
actors such as spatial heterogeneity and biotic relationships in any gien area.
Larly ecologists thought herbiore assemblages as interacting species without any
reerence to space ,1ilman & Kareia, 199,, but that changed once spatial ecology
including the metapopulation theory deeloped ,lanski, 1998,. 1oday, landscapes are
iewed as patches o habitats where indiidual species persist through migration between
dierent populations. Local species richness is oten inluenced by processes like
immigration and emigration that operate at a regional scale ,lanski, 1998,. 1hese
processes are the cornerstones o the metapopulation theory. 1he dispersal moement o
animals ensures genetic exchange amongst populations, thereby maintaining genetic
diersity in a species. 1hus, or a gien regional species-pool, the species diersity at a
local scale ,alpha diersity, is determined by the moement o indiiduals amongst
populations ,Rickles & Schluter, 1993,. It is, howeer, to be noted that metapopulation
dynamics are not addressed explicitly in this thesis, and this inormation here seres as
mere inormation about processes underlying herbiore species-richness patterns.
Although it is ery important, especially in the context o the large herbiore assemblage
in the 1rans-limalaya, and I come back to the subject in the synthesis section.

0DFURHFRORJLFDOWKHRU\DQGKHUELYRUHGLYHUVLW\
Although MacArthur`s model and metapopulation based models are useul in explaining
species richness patterns at local and regional scales, it does not account or large-scale
determinants such as eolutionary history and broad scale biogeography, which are also
important in understanding species richness patterns, especially at continental scales
,Rickles, 2004,. Species undergo adaptie radiations by eoling phenotypes that are
successul in a noel enironment ,Ropiquet & lassanin, 2005,, which ultimately lead to
speciation. \hen a species ails to eole in response to the changing enironment, it
goes extinct, thereby paing way or another species.
1hereore, apart rom ecological actors, historical actors are also important in
explaining species distribution at macroecological scales ,Rickles & Schluter, 1993,. lor
instance, the continental scale biogeographic patterns on the planet are the result o plate
tectonics, consequent orogenesis, glaciations and climatic oscillations. 1he inluence o
these on large scale biogeography can be studied by examining the aunal relationship
between dierent continents using ossil records ,Briggs, 2003,. 1he other actors that
are important in explaining continental scale biogeography are dispersal, icariance,
extinction and eolution ,Rickles & Schluter, 1993,.
Despite its importance in understanding the diersity pattern early in the history
o ecology, the ield o biogeography has been relegated being labeled as a descriptie
science. Part o the reason that this ield remained less deeloped is because ecologists
took a reductionist approach looking at community patterns and processes at a local scale
,Brown, 1985,. But with ecologist taking a more macroecological and holistic approach,
this important ield o biology is being integrated into the mainstream ecology ,Brown &
Lomolino, 1998,.

Introduction
7
+HUELYRUHGLYHUVLW\LQ/DGDNK
1he 1rans-limalaya encompassing the 1ibetan plateau and its marginal mountains ,F2.5
million km
2
, represents a ast and unique grazing ecosystem in the world as it has a
ascinating assemblage o wild herbiores. Small sections o this huge plateau extend into
the Indian 1erritory at seeral locations in the north and north-eastern parts o the
country, and are collectiely called the Indian 1rans-limalaya, o which almost 0 is
represented by the Ladakh region o the north Indian state o Jammu and Kashmir.
Ladakh is located at the intersection o two biogeographic realms YL] Palaearctic and
Oriental, and contains elements rom these realms. But most o the herbiores are
Palaearctic, a trend also obsered in the regions` birds ,Namgail & \om-1o, 2009,. It
een harbours species such as cape hare /HSXVFDSHQVLVrom the Lthiopian realm. Ladakh
also represents a biome transition between the ast plains o the 1ibetan Plateau and the
rugged mountains o lindukush-Karakoram ranges ,lig. 3,, and thus harbours a
relatiely dierse assemblage o mammalian herbiores with biogeographic ainities to
these biomes ,lig. 4,. 1he regional herbiore species-pool comprises twenty mammalian
herbiores representing 6 amilies and 11 genera ,lox et al., 1991, Pister, 2004,.

lig. 3. Location o Ladakh with regard to major mountain ranges in south and central
Asia ,source Google Larth,. 1he dotted line marks the approximate boundary between
the Palaearctic and the Oriental Biogeographic realms.

1he large herbiores o Ladakh ,2 kg, include eight wild ungulates, YL]1ibetan gazelle
3URFDSUD SLFWLFDXGDWD 1ibetan antelope 3DQWKRORSV KRGJVRQL Blue sheep 3VHXGRLV QD\DXU
Ladakh urial 2YLV YLJQHL YLJQHL, Asiatic ibex &DSUD LEH[ VLEHULFD, 1ibetan argali 2YLV DPPRQ
KRGJVRQL Kiang (TXXV NLDQJ and wild yak %RV PXWXV ,Namgail, 2009, see lig. 4,, two
marmot species: limalayan marmot 0DUPRWD EREDN and the long-tailed marmot 0
FDXGDWD and two hare species: 1ibetan woolly hare /HSXVRLRVWROXVand cape hare Some o
the herbiores like Ladakh urial and 1ibetan gazelle are conined to small geographical
areas, while others like blue sheep are more widely distributed. Apart rom these large
herbiores, there are seeral small mammalian herbiores such as pikas 2FKRWRQDspp. and
Chapter 1
8
oles $OWLFROD spp. ,Pister, 2004, Bagchi et al., 2006,, about which ery little is known.
1he large herbiores hae ragmented distributions, and many are endemic and rare with
global conseration importance ,Namgail, 2009,. 1he mechanistic understanding o the
species-richness patterns o these herbiores is thereore important rom both academic
and applied conseration iewpoints.
Due to the ragmented nature o the species` distributions, and their dierential
responses to the enironmental actors ,Mallon, 1983, Namgail et al., 2004,, there is a
spatial ariation in large herbiore species-richness across Ladakh with some parts
harbouring higher number o species compared to other areas. Len at smaller spatial
scales, some alleys support more and others less species. lor instance, the Rumchung
alley in the lemis ligh Altitude National Park supports three Caprinae species, while
the Puyul alley in the proposed Gya-Miru \ildlie Sanctuary has only two species ,see
Chapter 5,. 1hus, it is apparent that local assemblages represent small incomplete subsets
o the regional species pool.
I inestigated this spatial pattern, searching or plausible causes responsible or
this pattern. Such a study is crucial or a mechanistic understanding o herbiore diersity
pattern o Ladakh as well as other 1rans-limalayan rangelands. 1he object o this thesis
is thus twoold: 1, Understanding the distributional patterns o herbiore species-
richness pattern, 2, Lxploring plausible causes operating at local as well as regional scale
that inluence such spatial pattern o herbiore species-richness. litherto, most o the
studies on large herbiores hae been carried out in single assemblages, and the
mechanisms that operate at regional scale ,macroecological scales, hae not been
addressed adequately. Since the actors that aect species distribution at dierent spatial
scales are important in consering biodiersity in the long-run, it is crucial to examine
them in detail.
C
a
s
p
i
a
n
s
e
a
Cape haie
Asiatic ibex
Longtaileu maimot
Lauakh uiial Tibetan gazelle
Wilu yak
Tibetan antelope
Tibetan aigali
Blue sheep
Kiang
Tibetan woolly haie
Bimalayan maimot
lig. 4. Study area and species positioned according to their biogeographic ainities ,note
the conspicuous absence o Oriental species, the cape hare in the bottom let corner o
the map is rom the Lthiopian Biogeographical realm, which is not shown on the map,.
1he broken line marks the approximate boundary between the Palaearctic and the
Oriental Biogeographical zones.
Introduction
9
7KHVWXG\DUHDDQGVSHFLHV
Ladakh ,32
o
to 36
o
N and 5
o
to 80
o
L, is the largest proince o the northern Indian
state o Jammu and Kashmir, and encompasses F 80,000 km
2
,lig. 5,. It is bounded by
Karakoram range in the north and the Great limalaya in the south, and is bounded by
the 1ibetan plateau in the east and the lindukush Mountains on the west ,see lig. 3,.
1he greatest extent o the region is northwest to southeast coering F350 km. It is one
o the remotest regions in India, and one o the highest inhabited regions in the world. It
has the lowest human population density ,3 persons,km
2
, in India. 1he inhabitants are
mostly nomadic pastoralists in the east, and agro-pastoralists in the west, although this
trend is changing ,Namgail et al., 200a,. 1he inhabitants in the east and central parts o
Ladakh are predominantly Buddhists with cultural ainities with 1ibet, while those in the
west are mainly Shia Muslims.
Ladakh is located at the junction o two biogeographic realms: Palaearctic and
Oriental and has elements rom these biogeographic zones. But most o the mammalian
herbiores in the region are Palaearctic in origin with ery ew smaller herbiores such as
the Royle`s pika 2FKRWRQD UR\OHL rom the Oriental. 1here is also one Lthiopian species:
cape hare as mentioned earlier. Such a trend was also obsered in the region`s aiauna
,Namgail & \om-1o, 2009,. On a smaller scale, it is also located at the junction o two
biomes: 1ibetan plateau and the lindukush-Karakoram Mountains ,including the
Ladakh and Zangskar ranges, as alluded to earlier, and thus harbours species that hae
the core o their ranges in these biomes. 1he relatiely high species richness o large
herbiores in the region could perhaps be related to this juxtaposition o biomes.

lig. 5. Ladakh region in the state o Jammu and Kashmir, Indian 1rans-limalaya.
Chapter 1
10
&OLPDWH
1he climate is characterised by extreme aridity with an aerage annual precipitation o F
500 mm, and the region has been classiied as a cold desert. Being located in the rain-
shadow o the Great limalaya, it receies only ew monsoon clouds. 1hereore the
precipitation is minimal, mostly in the orm o snow in winter, which is associated with
the extra-tropical disturbances o mid-latitudes known as \estern Disturbances` ,Dhar
& Mulye, 198,. 1he southwestern part o the region, howeer, gets slightly more
precipitation compared to the east and the northeast, and thus is greener ,lartmann,
1983,. 1his discrepancy is indicated by high snowall on Zuzhi La` ,La means pass in
Ladakhi,, and less on Chang La and Khardung La, which remain open to traic during
winter except during exceptionally heay snowall. 1he climate is characterised by great
extremes o cold ,upto -35
o
C during winter, and heat ,upto -35
o
C during summer,, and
there is a great diurnal dierence in temperature with cool nights and hot days. 1he
humidity luctuates between 50 and 60 rom May to October at arious places ,Murti,
2001,. ligh-elocity winds blow all through the year, and absence o high humidity helps
to keep the atmosphere clear.

9HJHWDWLRQ
Champion and Seth ,1968, classiied Ladakh as a dry alpine scrub that completely lacks
orest coer, except relict populations o juniper -XQLSHUXVspp. stands in some parts such
as Zangskar and Sham. Some other common trees are willows 6DOL[ spp. and poplars
3RSXOXV spp. that are restricted to areas along riers and streams. 1here are also some
shrubs such as +LSSRSKDH UKDPQRLGHV and 0\ULFDULD JHUPDQLFD that grow along streams.
Although the egetation o Ladakh is sparse compared to the egetation o similar
altitudes in the limalayan range, it is o immense signiicance as it represents elements
rom seeral phytogeographic realms such as the Palaearctic and Oriental. Various
botanists hae described the egetation o Ladakh as dry bushes, alpine steppe, dry alpine
scrub and alpine stony deserts ,Rawat & Adhikari, 2002,. Neertheless, there is a great
plant diersity and more than 600 ascular plants hae been reported rom the region
,Kachroo et al., 19,.

6RLO
1he soils o Ladakh are sandy or sandy-loam, and are generally characterized as poor in
organic matter and nitrogen content ,Murti, 2001,. 1he pl ranges between and 11
,Bhat, 1965,. In the absence o any substantial leaching o minerals rom the soil, bases
are continuously being added to the soil complex, which result in high pl alue, this
together with the absence o organic matter turn the soil toward alkalinity ,Rawat &
Adhikari, 2002,. Calcium content is relatiely high, but the soil is poor in magnesium.
1here are, howeer, adequate plant nutrients in the soil due largely to low precipitation
and hence less weathering o rocks. \ithin Ladakh there are ariations in soil
composition. lor instance, the Nubra and Dras alleys are morainic, and the soils in the
Suru alley in Kargil are black due to grey shale, the Puga alley in eastern Ladakh has
sulphur and borax deposits.

*HRPRUSKRORJ\
One o the unique eatures o the geology o Ladakh is the presence o a huge plutonic
mass, the Ladakh Batholith, which is characterised by a long ,F350 km, ridge with peaks
up to 6200 m asl. 1he northern slope o this batholith is exposed in the Nubra alley,
and is in juxtaposition with the Khardung and Shayok olcanics ,Bhutani et al., 2009,.
1he southern side o the Indus Valley comprises a series o interlue ridges o the Indus
Molasse that are deeply incised by its tributaries, the largest o which is the Zangskar
Introduction
11
rier. 1he present day limalaya was once occupied by the 1ethys sea, which separated
India rom Lurasia ,1erra & lutchinson, 1934,. In the mid-Locene, the Indian plate
collided against the Asian plate, and the latter buckled under the ormer and uplited the
1ethys seabed, orming the mighty 1ibetan Plateau between Asia and the Indian sub-
continent, a process which still continues.
Ladakh displays a geomorphology that is heaily inluenced by reworking o rost
shattered rock and Quaternary glacial deposits through snowmelt, glacial runo and mass
moement processes. Major damming o riers has been identiied as haing had a large
inluence on the geomorphology that is currently isible in the region. Particularly
notable are the lake sediments, such as those ound at Lamayuru and Leh, dated to
around 35-40 ka BP, where lacustrine deposits are presered at the distal ends o alluial
ans that discharge into the Indus Valley.
3DODHRHQYLURQPHQW
Ladakh was glaciated during much o the Quaternary period, which is demonstrated by
the presence o wide, U-shaped glaciated alleys and extensiely glaciogenic sediments
along the upper course o major riers ,Pant et al., 2005,. \ell-deeloped lateral
moraines presered at many places ,e.g., Baralacha La and Shayok and Nubra alleys,
clearly indicate that this region experienced at least two glacial adances during the
Quaternary period. 1he Last Glacial Maximum in the limalaya as well as in western
1ibet, howeer, was less extensie due to weak monsoon.
\ith the retreat o major alley glaciers, lacustrine and luial enironments
dominated the region. 1he lacustrine enironment was restricted and preailed in areas
where the terminal moraines or landslide debris could block the riers and create lakes.
1he terminal phase o lacustrine successions in the region was dominated by are and
rhythmite sedimentation. Similarly, the transormation rom meandering to braid-
meandering riers suggests dwindling in hydrological regime. 1he snowline also retreated
leading to the deelopment o hanging glaciers.

3URPLQHQWVWXG\VSHFLHV
Amogst the eight wild ungulates in Ladakh, our belong to the sub-amily Caprinae.
1hese are the 1ibetan argali, Ladakh urial, Asiatic ibex and blue sheep. 1he 1ibetan argali
is the largest wild sheep in the world, standing 3.5 to 4 eet at the shoulder with the horn
measuring 90-100 cm. 1he animal is light-brown with its rump, throat, chest and belly
white. 1he Ladakh urial is a small wild sheep o about 80 cm high at the shoulder. 1he
coat-color o the upper parts o the body is ruous-grey, while the underparts and legs are
whitish. 1he Asiatic ibex is a majestic wild goat o 80-100 cm high at shoulder. 1he adult
males hae long pointed beards and scimitar-shaped horns with prominent ridges on the
rontal surace. 1he blue sheep is a unique mountain ungulate that is placed between
sheep and goat as it displays sheep as well as goat-like characteristics. lor example, it has
a lat tail with bare central surace, lacks acial glands, and during combats the rams rear
up and clash their horns like goats, while they lack a beard and rub their aces to the
rump o their rials, which are characteristic eatures o sheep.
Apart rom these there are two antelopes: 1ibetan gazelle and 1ibetan antelope.
1he 1ibetan gazelle is a small antelope weighing about 15 kg. It has a greyish-brown
body and short, black-tipped tail in the center o a heart-shaped white rump-patch. 1he
1ibetan antelope is a graceul animal adapted to the highlands o 1ibet. 1he animal is
conined to Aksai Chin and the Chhang Chhenmo areas o northeastern Ladakh. Apart
rom these there is a boid: \ild yak, and an equid: 1ibetan wild ass, as mentioned
earlier. \ak is a sturdy and bulky ungulate with high lung capacity and thick coat, which
are adaptations to the high altitude enironment o Ladakh and 1ibet. \ild yaks are
Chapter 1
12
mostly black with greyish muzzle, but some are aberrantly brown. 1he 1ibetan wild ass
or .LDQJ is the largest wild ass in the world with some stallions standing 1.4 m tall and
weighing upto 400 kg. 1he colour is auburn with white belly, chest and legs, and the
sexes are alike.

7KHWKHVLV
Mammalian herbiores are highly mobile, especially in patchy enironments where they
need to moe constantly to acquire their preerred plant species and other resources,
although less than the mammalian carniores. 1hereore, species assembly patterns in
mammalian herbiores need to be looked at multiple spatial scales, because actors that
operate at dierent spatial scales may interact. Keeping this in iew, this study was
carried out at dierent spatial scales. Chapter 2 presents the results o a biogeographic
analysis o the ascular plants o Ladakh in relation to altitude. 1his chapter also sheds
light on the distribution pattern o aboeground biomass in relation to altitude, which
has not been assessed beore, neither in the 1rans-limalaya nor, to my knowledge,
elsewhere.
Chapter 3 deals with the geographical communities o the mammalian herbiores
o Ladakh in a spatially explicit way. It explores whether the mammalian herbiore
species in the Ladakh 1rans-limalaya orm chorotypes or groups sharing distributions,
and i so, what explains the congruence in their distributions. It highlights the act that
mammalian herbiore distribution at a regional scale is determined largely by niche
conseratism. 1he chapter urther shows that riers are not eectie geographical
barriers and thus do not inluence the zoogeography o Ladakh.
I then moed on to assemblage leel mechanisms to see how they can explain the
herbiore species-richness pattern in Ladakh. Since the 1rans-limalayan rangelands are
less productie with ery low graminoid biomass compared to other natural grazing
ecosystems in the world ,Mishra, 2001,, I expected competition to be an important actor
that inluences the herbiore assembly structure and in turn the species-richness pattern.
lurther, since blue sheep is the most abundant wild ungulate in Ladakh, and thus
probably the dominant species, I inestigated the niche relationship between this species
and other sympatric species with similar body size to assess the role o competition in
range dynamics o large herbiores. Chapter 4 presents the results o an inestigation o
the nature o interaction between the Asiatic ibex and blue sheep in winter, which is the
pinch period, when herbiore species are more likely to compete.
Chapter 5 assessed the niche dynamics o blue sheep, again at a regional scale.
1his chapter emphasises the inluence o the number o sympatric species on the niche
utilization o species. It demonstrates that the habitat-niche width o a species declines in
response to the number o sympatric species, whereas the diet-niche width has a hump-
shaped relationship with sympatric species richness.
Chapter 6 examines i the blue sheep constrains the range expansion o the
Ladakh urial, which is conined to narrow tracts along two rier alleys. I studied the co-
occurrence pattern o the two species at geographical, landscape and habitat leels. 1he
results indicate that there is a high potential or competition between these two species,
especially during winter when the blue sheep descends to aoid snow coer at higher
altitudes. 1he chapter underscores the act that species co-occur at large spatial scales but
segregate at small spatial scales.
linally, in Chapter , I discuss the results o this research in light o the current
discourse on herbiore species-richness patterns and the underlying mechanisms. I
synthesise by drawing on arious chapters o this thesis as well as rom my pre-PhD
research on the large herbiores o Ladakh. I also underline the implications o the
results or mammalian herbiore conseration in Ladakh and other dry alpine
Introduction
13
ecosystems. I highlight urther gaps in inormation and research prospects that will
oster mechanistic understanding o large herbiore assembly rules in the 1rans-
limalaya and other mountainous regions o the world.

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17
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Phytogeography of Ladakh
19
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Chapter 2
20
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1he Ladakh region o the Indian 1rans-limalaya harbours more than 600 ascular
plants, representing 190 genera and 51 amilies, but their distributional patterns ,both
horizontal and ertical, remain little explored. \e studied the spatial pattern o ascular
plant species-richness in relation to altitude at a geographical as well as a local scale in the
region. \e used Generalized Linear and Generalized Additie Models to assess the
relationships. 1he relationship between aboeground biomass and altitude was also
assessed in our protected areas. \e ound a hump-shaped relationship between ascular
plant species-richness and altitude at both local and geographical scales. 1he species
richness at the local scale in eastern Ladakh ,aerage altitude ~ 4900 m, peaked between
5000 m and 5200 m aboe sea leel. 1he cubic smooth spline indicated that the plant
species richness at the geographical scale peaked around 4000 m asl. Vegetation coer at
this scale peaked at an altitude o about 4300 m. Contrary to our expectation, we ound a
unimodal relationship between aboeground biomass and altitude. \e suggest that such
a trend is engendered by excessie grazing by domestic liestock and less precipitation in
alley bottoms due to rainshadow eects and high eaporatie losses because o high
temperature.
.H\ZRUGVVascular plants, species richness, aboeground biomass, altitude, Rapoport`s

rule, mid-altitude eect, macroecology
21
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1he spatial ariation o biodiersity on earth is an interesting natural phenomenon. 1he
declining species diersity rom the tropics to the temperate regions ,Rapoport`s
Latitudinal Rule, is one o the most prominent pattern, which has widely been studied
,Pianka, 1966, Rickles & Schluter, 1993,. Larlier, plant species were thought to be more
dierse at lower sites and decline in diersity with altitude. 1hus, the altitudinal gradient
in species richness in mountainous regions was thought to be analogous to the
Rapoport`s Latitudinal Rule ,Steens, 1992,, but more recent studies hae contested this
iew, and hae shown that biodiersity in the mountainous regions does not peak at
lower altitudes but somewhere in the middle o the gradient ,Rahbek, 1995, Rahbek,
199,, although some did obsere a negatie linear relationship ,Odland & Birks, 1999,.
1he discrepancy was largely attributed to biases engendered by dierences in sampling
area and eort ,Rahbek, 1995,.
A mechanistic understanding o the altitudinal gradient in species richness is
crucial or the management o natural resources largely in the light o the current climatic
changes. 1he altitudinal gradient in plant diersity has been the centre o attention o
plant ecologists in the last couple o decades, and studies were carried out in all the
mountainous regions o the world, including the limalayas ,Bhattarai & Vetaas, 2003,
Oommen & Shanker, 2005,. Altitudinal gradient is one o the most decisie actors or
spatial pattern o phytodiersity, because it presents changes in aailability o abiotic
actors such as heat, moisture and nutrients, which determine the growth o plants.
Neertheless, there are alternatie hypotheses that account or this pattern, including area
,Rosenzweig, 1995,, hard boundaries ,Colwell & lurtt, 1994, and climate ,Currie &
Paquin, 198,. But hitherto almost all the studies on the subject were carried out in the
tropics and temperate regions, and inormation remains rudimentary rom the alpine
systems, which in act are ideal or studying biodiersity patterns along altitudinal
gradients as the enironment in these systems can change within short distances.
Aboeground plant biomass is an important component o grazing ecosystems
and determines the stocking density o herbiores. It is thereore crucial to understand
the spatial pattern o aboeground biomass or eectie and sustainable management o
rangelands or wild as well as domestic herbiores. Neertheless, although the spatial
pattern o net primary productiity and its relation with climatic actors was studied in
1rans-limalayan mountains ,\ang et al., 2009, and other natural grassland ecosystems
,Sala et al., 1988, Lpstein et al., 199, Jobbagy et al., 2002,, the pattern o aboeground
biomass along an altitudinal gradient has not been explored in a spatially explicit way,
neither in the 1rans-limalaya nor, to our knowledge, elsewhere. Altitude, as alluded to
earlier, is an important gradient along which aboeground biomass is expected to decline
due to declining temperature and nutrients, which get washed down ,Rastetter et al.,
2004,. But there has been no apparent eort to document this with empirical eidence,
and it is less clear i such a trend can be expected in drier enironments o deserts with
little complexity in egetation structure.
1he Ladakh region ,32
o
to 36
o
N and 5
o
to 80
o
L, o the Indian 1rans-limalaya
is a high altitude cold desert and supports a unique assemblage o lora adapted to
extreme enironmental conditions ,Rawat & Adhikari, 2005a,. 1he region has unrialed
geographical and topographic eatures, and an altitudinal range rom 2800 to ,60 m
aboe sea leel, which proides an array o habitats or alpine plants. Ladakh is located at
the intersection o two major biogeographical zones: Palaearctic and Oriental, and hence
has elements rom these realms. Kachroo HWDO ,19, reported 611 ascular plants rom
the region, representing 190 genera and 51 amilies. 1he spatial pattern o plant
distribution and plant community structure in this area is howeer poorly known, and
Chapter 2
22
some inormation on these aspects hae started to come in only recently ,see Kala &
Mathur, 2002, Rawat & Adhikari, 2002, Klimes, 2003,.
1he main objectie o this study was to understand the ascular plant distribution
and abundance along an altitudinal gradient at dierent spatial scales in Ladakh. \e were
interested to know whether these parameters are monotonically or unimodally related to
altitude. Based on inormation rom literature, we predicted 1, a hump-shaped
relationship between ascular plant species-richness and altitude, 2, a negatie
relationship between aboeground biomass and altitude.

0DWHULDOVDQGPHWKRGV
6WXG\DUHD
Ladakh is situated in the Indian 1rans-limalaya, coering an area o F80,000 km
2
. It is
unique in terms o geography, geology and edaphic eatures ,Kachroo et al., 19,. 1he
egetation community is dierse due to these and the great altitudinal range mentioned
earlier. Ladakh is characterized by high, steep and rugged mountains separated by deep
alleys into which the Indus and Zangskar riers and their tributaries low. 1he climate is
characterized by long, cold periods without aourable conditions or growth in winter
and short growth seasons in summers, and there is a marked diurnal ariation in
temperature. 1he soil is nutrient-poor, consisting o sandy clay and at places coarse
grael or murram, and become more alkaline in some parts. 1he region supports ery
sparse egetation coer consisting o low shrubs, stunted grasses and herbs ,Chaurasia &
Singh, 199, Rawat & Adhikari, 2005b, largely due to the short growing season, harsh
climatic conditions and poor soil.
Ladakh is bounded by the Great limalayan range on the south and the
Karakoram range on the north. 1he ormer range blocks most o the monsoon bearing
clouds, making the region arid. \inter precipitation is mostly in the orm o snow, which
is associated with the extra-tropical disturbances o mid-latitudes known as \estern
Disturbances` ,Dhar & Mulye, 198,. 1he south-western part o the region, howeer,
gets slightly more precipitation compared to the east and the north-east, and thus is
greener ,lartmann, 1983,. 1he egetation o eastern Ladakh is largely dominated by
lemicryptophytes, ollowed by 1herophytes and Chamaephytes ,Klimes, 2003,.
Champion and Seth ,1968, described the egetation o Ladakh as dry alpine scrub` that
is characterised by complete absence o orest coer, except relict patches o juniper
-XQLSHUXVSRO\FDUSRV stands in some parts o Zangskar and Sham. Other prominent trees are
willow 6DOL[ spp. and poplar 3RSXOXV spp. that are restricted mainly to cultiated areas
along riers. 1here are also other shrubs such as seabuckthorn +LSSRSKDH UKDPRLGHV
WXUNLVWDQLFDand 0\ULFDULDHOHJDQV that grow along stream and rier banks.
Ladakh harbours a relatiely rich assemblage o wild mammals such as 1ibetan
gazelle 3URFDSUD3LFWLFDXGDWD1ibetan antelope 3DQWKRORSVKRGJVRQLBlue sheep 3VHXGRLVQD\DXU
Ladakh urial 2YLV YLJQHL YLJQHL, Asiatic ibex &DSUD LEH[ VLEHULFD, 1ibetan argali 2YLV DPPRQ
KRGJVRQL 1ibetan wild ass or Kiang (TXXV NLDQJ and wild yak %RV PXWXV 1he prominent
predators include the endangered snow leopard 8QFLDXQFLD and 1ibetan wol &DQLVOXSXV
FKDQFR 1he mountain slopes ,or the rangelands, are used by the local communities or
grazing a ariety o domestic liestock including yak, sheep, goat, horse, donkey, cow and
dzo ,hybrid between yak and cow,. 3DVKPLQD or cashmere wool, obtained rom a local
breed o goat called &KDQJUD, is the most aluable product rom these rangelands. 1he
local people in the western part are agropastoralists with major emphasis on agriculture,
while those in the east are mainly nomadic pastoralists ,Namgail et al., 200,. Ladakh has
the lowest human population density in India with less than 3 persons,km
2
.

23
6DPSOLQJPHWKRGV
3ODQWVSHFLHVGLYHUVLW\
Sampling or the local scale inestigation was carried out in July 2004 ,during peak
standing biomass, in the lanle Valley o eastern Ladakh. 1wo transects ,each coering
1000 ertical meters between 4500 and 5500 m asl with no ariation in aspect, were
walked on a hillside, and plant species richness was sampled using a line-intercept
method ,Muller-Dumbois & Lllenberg, 194,. \e placed 20 m line intercepts at interals
o 50 ertical meters, and recorded plant species ,or any other substrate such as soil or
rock, at eery 0.5 m interal along the line intercept. 1he species richness o graminoids
and non-graminoids along each transect were recorded separately. 1he highest altitude
sampled was the top o the hill.
Data or assessing the species richness at a regional scale were collected between
July and September 200. \e droe rom Rangdum in the west to lanle in the east
,approx. 600 km, altitudinal range, 2800 to 6000 m asl,, and 1urtuk in the north to
Sarchhu in the south ,approx. 300 km, altitudinal range, 2860 to 4900 m asl,, at an
aerage speed o 50 km,hour, stopping eery one hour and laying 20 m transects ,similar
to the local scale method, on randomly selected adjacent slopes. \hen the road was
rough, which slowed us down, we increased the time interal between samplings to aoid
spatial autocorrelation. \e also aoided sampling on human-modiied landscape. Plants
were identiied in the ield using ield guides ,e.g., Kachroo et al., 19, Polunin &
Stainton, 1990, Aswal & Mehrotra, 1994,. 1hose species that could not be identiied in
the ield were brought to the lerbarium at the \ildlie Institute o India or
identiication.
1o assess the relationship between species richness pattern and climate at the
regional scale, we obtained climatic ariables: precipitation in the wettest quarter o the
year ,abbreiated in the igures as Pre\etQ,, mean annual precipitation ,AnnPrec,,
precipitation in the driest quarter ,PreDriQ,, annual mean temperature ,AnMe1e,, mean
temperature o the coldest quarter ,M1eCoQ,, mean temperature o the warmest quarter
,M1e\aQ, at 1 km
2
resolution rom the \orldClim database ,lijmans et al., 2005,.
1hese ariables were extracted rom all the transect locations in a GIS enironment
,ArcView, 9, LSRI, 1996,
3ODQWELRPDVV
Data or assessing the relationship between aboeground biomass and altitude were
collected between July and September 200. Sampling was done in our established and
proposed protected areas: the proposed Ridzong \ildlie Sanctuary ,34
o
20` N,
o
04`
L,, lemis National Park ,34
o
06` N,
o
23` L,, the proposed Gya-Miru \ildlie
Sanctuary ,33
o
40` N,
o
4` L, and the Changthang \ildlie Sanctuary ,33
o
19` N, 8
o
29`
L,. A transect was laid on an altitudinal gradient at eery 200 m, alternately on either side
o a alley, starting at the alley-mouth. Lach transect was then diided into 50 m ertical
segments, and a 2 x 2 m plot was sampled at eery 50 m intercept. Vegetation was
clipped to the ground leel in these quadrats and stored in paper bags. 1hey were then
sun-dried and weighed to the nearest 0.1 g to estimate aboeground biomass. \e
aoided sampling in human-modiied landscape and along rier and stream banks, which
generally support high biomass o woody species due to high moisture rom them.

9HJHWDWLRQFRYHU
\e estimated egetation coer ,in coer classes o ten percent, in association with
altitude and slope. Data or this were collected in August 2006. \e explored the area
between Lamayuru and Sarchhu in the Zangskar mountains. lirst we walked along the
upper reaches o the Zangskar Rier, oer the Sisir-la Pass ,400 m, and Singge-la Pass
Chapter 2
24
,4825 m,, and then along the 1sarap, Lingti and Kargyak riers at about 4200 to 4800 m,
oer the Shingo-la Pass ,5050 m,, and inally along the Sangpo and Jankar riers at about
4500 to 300 m. Locations and altitudes were determined through a global positioning
system ,Garmin GPS 12, and an altimeter ,Origo Accusense Multi-sensor,. In total we
walked about 250 km at an aerage altitude o 4800 m asl.

6WDWLVWLFDODQDO\VLV
lirst we analysed the data using Generalized Linear Models, but when we assessed the
dierence between a logarithmic link and an identity link unction ,assuming normal
distributions o error, by drawing diagnostic Q-Q plots ,Q stands or quantile, o the
residual ariation, the Poisson Model had a slightly better it, perhaps due to ewer
counts. 1hereore, we used Generalized Additie Models ,GAM, assuming a Poisson
distribution. \ithin the GAM ramework, we used a cubic smooth spline so that any
abrupt change in egetation distribution could be captured ,lastie & 1ibshirani, 1990,.
1his is especially appropriate because Ladakh represents two biogeographic proinces:
1ibetan plateau in the east and the lindukush-Karakoram mountains ,including Ladakh
and Zangskar ranges, in the west, and there might be abrupt changes in lora.
Graminoids and non-graminoids were analysed separately at the local scale. \e regressed
species richness against the climatic ariables to ealuate the inluence o climate on
species richness. All these tests were carried out in Statistica .
\e used Canonical Correspondence Analysis ,CCA, to explore the relationship
between plant species distribution and enironmental ariables, putting all species in a
continuous enironmental perspectie. 1his method is useul in measuring the amount o
ariation in the species distribution data that can be explained by dierent explanatory
ariables. 1he statistical signiicance o this relationship was assessed using a permutation
test, which has an added adantage o the ability to check which ariables contribute
most to the relationship. 1he analysis was carried out using the CANOCO sotware
ersion 4.5 ,1er Braak, 1986,.

5HVXOWV
A total o 91 plant species were recorded during the inestigation at the regional scale,
while only 28 species were recorded during sampling at the local scale in lanle. Plant
species richness aried along the altitudinal gradient at both geographical and local scales.
Species richness had a hump-shaped relationship with altitude, peaking at mid-altitude at
both spatial scales ,ligs. 1 and 2,.
6SHFLHVULFKQHVV
Local scale
Plant species in 4 transects were recorded to assess the relationship between species
richness and altitude at the local scale. Both graminoids and non-graminoids had hump-
shaped relationships with altitude ,lig. 1,. 1he smooth spline with our degrees o
reedom or these unctional groups indicated a sharp hump between 5000 and 5200 m.
It also indicates a steep decline in species richness aboe 5200 or both unctional
groups. lor the graminoids there was also a sharp increase in species richness beore it
slumps, while or non-graminoids the increase and decrease were more gradual ,lig. 1,.

25
,a, ,c,
y = -2E-06x
2
+ 0.0193x - 48.216
R
2
= 0.4543
0
0.1
0.2
0.3
0.4
0.5
0.6
0.7
0.8
4400 4900 5400 5900
Altitude (m)
G
r
a
m
i
n
o
i
d

S
p
e
c
i
e
s

R
i
c
h
n
e
s
s

,b, ,d,
4200 4400 4600 4800 5000 5200 5400 5600 5800
Altitude (m)
-1.0
-0.8
-0.6
-0.4
-0.2
0.0
0.2
0.4
0.6
0.8
1.0
1.2
1.4
P
a
r
t
i
a
l

r
e
s
i
d
u
a
l

4200 4400 4600 4800 5000 5200 5400 5600 5800
Altitude (m)
-0.6
-0.4
-0.2
0.0
0.2
0.4
0.6
0.8
1.0
1.2
P
a
r
t
i
a
l

r
e
s
i
d
u
a
l

lig. 1. Unimodal relationship between non-graminoid ,a, b, and graminoid ,c, d, species
richness ,Simpson`s index, and altitude at a local scale. In igure b and d, the solid lines
are the cubic smooth splines and the dashed lines are 95 conidence limits.

Geographical scale
\e sampled plants rom 55 transects spanning entire Ladakh. 1wenty-nine plant species
were encountered most requently during the sampling at this scale ,1able 1,. 1he plant
species richness had a hump-shaped relationship with altitude ,lig. 2a,, and peaked at
4000 m asl ,lig. 2b,, and dominance was lowest around this altitude ,lig. 3,.

a, b,
y = -1E-07x
2
+ 0.0011x - 1.3415
R
2
= 0.3902
0.4
0.5
0.6
0.7
0.8
0.9
1
2500 3500 4500 5500
Altitude (m)
S
p
e
c
i
e
s

R
i
c
h
n
e
s
s

2000 2500 3000 3500 4000 4500 5000 5500 6000
Altitude (m)
-0.3
-0.2
-0.1
0.0
0.1
0.2
0.3
P
a
r
t
i
a
l

r
e
s
i
d
u
a
l
lig. 2. Scatterplot o relationship between plant species richness ,Simpson`s index, and
altitude at a regional scale. In igure b the solid line is the cubic smooth spline and the
dashed lines are the 95 conidence limits.

y = -1E-06x
2
+ 0.011x - 27.476
R
2
= 0.2535
0
0.1
0.2
0.3
0.4
0.5
0.6
0.7
0.8
0.9
4400 4900 5400 5900
Altitude (m)
N
o
n
-
g
r
a
m
i
n
o
i
d

S
p
e
c
i
e
s

R
i
c
h
n
e
s
s
Chapter 2
26
On aerage, Caprioliaceae and Crassulaceae had higher altitudinal range, while Iridaceae
had the lowest altitudinal range ,lig. 4,. &DUJDQD YHUVLFRORU .UDVFKHQLQQLNRYLD FHUDWRLGHV and
7DQDFHWXP WLEHWLFXP are some o the most common species that occurred at high altitude
drier areas, while $UWHPLVLD PDULWLPD (SKHGUD JHUDUGLDQD and (FKLQRSV FRUQLJHUXV occurred at
lower regions o Ladakh ,lig. 5,.
y = 1E-07x
2
- 0.0011x + 2.3415
R
2
= 0.3902
0
0.1
0.2
0.3
0.4
0.5
0.6
2500 3000 3500 4000 4500 5000 5500
Altitude (m)
D
o
m
i
n
a
n
c
e

lig. 3. Dominance pattern o plant species along an altitudinal gradient at a geographical
scale in Ladakh

A
m
a
r
y
l
l
a
i
d
a
c
e
a
e
B
e
r
b
e
r
i
d
a
c
e
a
e
B
o
r
a
g
i
n
a
c
e
a
e
C
a
m
p
a
n
u
l
a
c
e
a
e
C
a
p
r
i
f
o
l
i
a
c
e
a
e
C
a
r
y
o
p
h
y
l
l
a
c
e
a
e
C
o
m
p
o
s
i
t
a
e
C
o
n
v
u
l
v
u
l
a
c
e
a
e
C
r
a
s
s
u
l
a
c
e
a
e
C
r
u
c
i
f
e
r
a
e
C
u
p
r
a
s
s
a
c
e
a
e
E
u
p
h
o
r
b
i
a
c
e
a
e
G
e
n
t
i
a
n
a
c
e
a
e
G
e
r
a
n
i
a
c
e
a
e
G
r
o
s
s
u
l
a
r
i
a
c
e
a
e
I
r
i
d
a
c
e
a
e
L
a
b
i
a
t
e
a
e
L
e
g
u
m
i
n
o
s
a
e
Plant family
2600
2800
3000
3200
3400
3600
3800
4000
4200
4400
4600
4800
5000
5200
A
l
t
i
t
u
d
e

(
m
)

lig. 4. Altitudinal range o some common amilies o plants in the Ladakh 1rans-
limalaya.

27
1able 1. Plant species that were encountered in more than 50 o the transects at the
regional scale.
Species lamily Abbr. in CCA

$FRQRJRQXPWRUWXRVXP Polygonaceae Aco-tor
5RVDZHEELDQD Rosaceae Ros-web
3RWHQWLOODIUXWLFRVD Rosaceae Pot-ru
6DXVVXUHDWDUD[LFLIROLD Compositae Sau-bra
6DXVVXUHDEUDFWHDWD Compositae Sas-tar
(FKLQRSVFRUQLJHUXV Compositae Lch-cor
$UWHPLVLDEUHYLIROLD Compositae Art-bre
$UWHPLVLDPDULWLPD Compositae Art-mar
:DOGKHLPLDVWROLF]NDL Compositae \al-sto
7DQDFHWXPWLEHWLFXP Compositae 1an-tib
&LFHUPLFURSK\OOXP Papilionoideae Cic-mic
&DUDJDQDYHUVLFRORU Papilionoideae Car-er
$FDQWKROLPRQO\FRSRGLRGHV Phimbaginaceae Aca-lyc
$QGURVDFHPXFURQLIROLD Primulaceae And-muc
(SKHGUDJHUDUGLDQD Lphedraceae Lph-ger
/LQGHORILDORQJLIORUD Boraginaceae Lin-lon
5KHXPZHEELDQXP Polygonaceae Rhe-web
'UDFRFHSKDOXPKHWHURSK\OOXP Labiateae Dra-het
+HUDFOHXPSLQQDWXP Umbellierae ler-pin
*HUDQLXPVLELULFXP Gentianaceae Ger-sib
.UDVFKHQLQQLNRYLDFHUDWRLGHV Chenopodiaceae Kra-cer
$UHQDULDEU\RSK\OOD Caryophyllaceae Are-bry
2[\WURSLVPLFURSK\OOD Leguminosae Oxy-mic
$VWUDJDOXVUKL]DQWKXV Papilionoideae Ast-rhi
3HGLFXODULVFKHLODQWKLIROLD Scrophulariaceae Ped-che
&DUH[PHODQDQWKD Gramineae Car-utr
6WLSDRULHQWDOLV Gramineae Sti-ori
(O\PXVQXWDQV Gramineae Lly-nut

1able 2. 1he regression o climatic ariables and plant species richness. Pre\etQ ~
Precipitation in the \ettest Quarter o the year, AnnPrec ~ Mean Annual Precipitation,
M1eCoQ ~ Mean 1emperature o the Coldest Quarter, AnMe1e~ Annual Mean
1emperature,

Variable Beta Standard
error beta
B Standard
error B
t,24, P-leel
Pre\etQ -2.31 0.419 -0.010 0.002 -5.520 0.001
AnnPre 2.28 0.390 0.004 0.000 5.839 0.001
M1e\aQ 0.91 0.50 0.003 0.002 1.054 0.302
AnMe1e -0.892 0.09 -0.003 0.002 -1.258 0.220

Chapter 2
28
-0.6 1.0
-
0
.
4
0
.
6
Barren
Car-utr
Aca-lyc
And-muc
Sti-ori
Aco-tor
Sau-bra
Eph-ger
Ech-cor
Lin-lon
Art-bre
Art-mar
Rhe-web
Cic-mic
Ely-nut
Dra-het
Her-pin
Sas-tar
Ger-sib
Ros-web
Pot-fru
Car-ver
Kra-cer
Are-bry Wal-sto
Oxy-mic
Ast-rhi
Ped-che
Tan-tib
Altitude
PreDriQ
PreWetQ
AnnPre
MTeCoQ
MTeWaQ
AnMeTe
$ERYHJURXQGELRPDVV
\e estimated aboeground biomass rom 65 randomly selected quadrats in the our
protected areas that spanned a range o climatic and geographical regimes. 1he
relationship between aboeground biomass and altitude was also hump-shaped ,l ~
9.803, P ~ 0.003, lig. 6,.

lig. 5. CCA ordination o some common plant species in Ladakh and the enironmental
ariables. See 1able 1 or species identity. ,Pre\etQ ~ Precipitation in the \ettest
Quarter o the year, AnnPrec ~ Mean Annual Precipitation, PreDriQ ~ Precipitation in
the Driest Quarter, AnMe1e~ Annual Mean 1emperature, M1eCoQ ~ Mean
1emperature o the Coldest Quarter, M1e\aQ ~ Mean 1emperature o the \armest
Quarter,.

y = -2E-05x
2
+ 0.1369x - 288.59
R
2
= 0.2328
0
5
10
15
20
25
30
35
3200 3700 4200 4700 5200
Altitude (m)
B
i
o
m
a
s
s

(
g
)

lig. 6. Non-linear relationship between aboeground plant biomass ,dry weight, and
altitude in Ladakh.
29
1he mean aboeground biomass ,dry weight, or the entire region was 5.4 g m
-2
,range 1
to 16.4 g dry weight,. 1he mean aboeground biomass in the proposed Ridzong \ildlie
Sanctuary was 3.9 g m
-2
, lemis National Park was 4.3 g m
-2
, Gya-Miru \ildlie Sanctuary
was 6.4 g m
-2
, and that in Changthang \ildlie Sanctuary was 5.6 g m
-2
.
9HJHWDWLRQFRYHU
Vegetation coer aried as a unction o altitude and slope angle. It peaked in areas at an
altitude o 4200 m and zero to 10 degree sloped angle ,lig. ,. Low altitudes ,3600 m,
and ery high areas ,5000 m, had ery little egetation coer ,10,. Similarly, ery
steep areas ,45
o
, had little egetation coer, which is in line with our biomass data.

5200
4800
4200
3600
3200
10
45
0
10
20
30
40
50
60
V
e
g
e
t
a
t
i
o
n

c
o
v
e
r

(
%
)
A
l
t
i
t
u
d
e
(
m
)
S
l
o
p
e
(
o
)
50-60
40-50
30-40
20-30
10-20
0-10

lig. . Vegetation coer ,, as a unction o slope angle and altitude during peak o
egetation growth in the mountainous region o Ladakh, India.

1he CCA showed that amongst the most commonly encountered species, the
distribution o $UWHPLVLD PDULWLPD (SKHGUD JHUDUGLDQD (FKLQRSV FRUQLJHUXV and /LQGHORILD
ORQJLIORUD occur in areas with high precipitation ,lig. 5,. .UDVFKHQLQQLNRYLD FHUDWRLGHV
7DQDFHWXPWLEHWLFXP&DUDJDQDYHUVLFRORUand $VWUDJDOXVUKL]DQWKXVoccur in drier, high altitude
habitats. 1emperature is not a good determinant o plant species distributions ,1able 2,
lig 5,.

'LVFXVVLRQ
1he hump-shaped relationship between ascular plant species-richness and altitude
obsered in Ladakh conorms to the general trend obsered in studies on this subject
,Rahbek, 1995,. Such a pattern could be related to seeral enironmental and biological
actors that ary along an altitudinal gradient. lor example, the lower number o plant
species at lower altitude could be related to the competitie exclusion o some species by
other plant species with higher adaptation and tolerance towards heat, moisture and high
Chapter 2
30
soil nutrients, while the less species richness at higher altitudes may be related to the
instability o substrates on the steep slopes in higher areas as well as to lower temperature
,Klimes, 2003,. It is, howeer, to be noted that since altitude represents a combination o
seeral climatic ariables closely correlated with numerous other enironmental
properties, it is diicult to single out important actors indiidually. Local topography,
soil conditions and snow deposition pattern that ary along an altitudinal gradient can
inluence plant species diersity along such gradients ,Chapin & Korner, 1995, Rawat &
Adhikari, 2005b,.
1emperature is known to decrease monotonically with altitude with a lapse rate
o 0.6
o
C per 100 m, while precipitation has a more erratic pattern. Gien these, and
assuming that they hae positie eects on species richness, one could expect a
monotonic inerse relationship between ascular plant species-richness and altitude. But
we ound a hump-shaped relationship, suggesting that climatic actors hae less inluence
on the relationship, and local microtopography and soil conditions are more important as
reported by Rawat and Adhikari ,2005b, in 1sokar Basin, Ladakh. Neertheless,
precipitation seems to hae an eect at the regional scale as regression analysis showed a
strong relationship between precipitation and species richness at this scale, but
temperature did not aect the relationship, as also indicated by less contribution o this
ariable in the CCA o species distribution. Very high moisture leels also lead to
dominance by ew moisture-loing plants, which exclude other plants ,Ol et al., 2002,,
thereby negatiely aecting the plant species richness. 1his was indicated by the negatie
relationship between plant species richness and precipitation during the wettest quarter
o the year in Ladakh.
Although the climatic actors might determine the relationship between altitude
and plant species richness at the regional scale, biotic actors might become more
important in determining the relationship at a local scale. lor instance, biological actors
such as competing species might competitiely exclude plant species, with less tolerance
towards high temperature and nutrient leels, rom the lower slopes. Alternatiely, since
the species richness peaked at the mid-altitudes and then declines precipitously as the
mountain-top is approached, it could also be related to hard boundaries: mountain tops
and alley bottoms ,see Colwell & lurtt, 1994,.
1he unimodal relationship between plant biomass and altitude is contrary to our
expectation. Generally, one would predict a higher plant biomass at lower altitudes as the
plant coer is higher at the base o a mountain due to greater aailability o moisture and
nutrients ,Rastetter et al., 2004,, but we ound less plant biomass at lower altitudes,
which is intriguing. One possibility is that plants at low altitudes could be sparse and
stunted due to trampling and eeding by the domestic liestock. It is also to be noted that
because o the high temperature, the limited moisture in the air ail to precipitate at lower
altitudes, which makes these areas dry and perhaps less egetated. Moreoer, the high
mountains cause a rainshadow in the deeper alleys.
Pastoralists also collect species like &DUDJDQD sp. rom the lower slopes or
uelwood, which leads to soil erosion ,Rawat & Adhikari, 2005b, and perhaps to lower
plant diersity. 1hese leguminous plants are known to inluence ertility as its root-
nodules hae nitrogen-ixing bacteria. lurther they help retaining moisture in the soil.
1hereore, extraction o this plant might hae led to the impoerishment o lora at
lower slopes. Anthropogenic pressures such as excessie liestock grazing and collateral
actiities are known to inluence the phytogeography in the alpine grazing ecosystems. A
discussion on this issue has been started in the limalayan region ,Saberwal, 1996, Mishra
& Rawat, 1998,, and the potential role o liestock grazing in creating spatial pattern o
plant distribution and abundance need to be studied in detail, because the liestock
population on the rangelands o Ladakh is increasing apace ,Namgail et al., 200,.
31
Vegetation coer peaked at around 4300 m. Such a trend could be related to the high
moisture content at higher altitudes due to low temperature and thus high precipitation,
as alluded to earlier. Rawat and Adhikari ,2005b, reported an increase in plant coer with
altitude until the nial zone, and related the trend to a high precipitation and thus high
soil moisture content.
1he main conclusions o the study are as ollows: plant species richness peaked at
mid-altitude at both local and geographical scales. At the local scale the graminoids had a
narrower peak than the non-graminoids. Precipitation aected the relationship between
altitude and species richness at the geographical scale, but temperature did not inluence
the relationship. Contrary to the general belie, the plant biomass also had a hump-
shaped relationship with altitude. Perhaps, excessie grazing by liestock at the lower
altitudes, low precipitation at low altitudes might hae caused such a spatial pattern,
either singly or in tandem. 1o our knowledge, this is the irst study on the relationship
between aboeground biomass and altitude, and has both theoretical and conseration
implications. lurthermore, this is the irst study o this kind in Ladakh at multiple spatial
scales, and thus establishes a basis or urther experimental work and mechanistic
understanding o plant distributional patterns in the region.

Acknowledgements
1his study was inancially supported by the \ageningen Uniersity, the Ruord Small
Grants loundation, and the \ildlie Conseration Society. \e thank Karma Sonam or
his assistance in the ield. \e thank the Director, \ildlie Institute o India or all the
help to 1N during his stay in Dehradun.

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Saberwal, V. K. ,1996, Pastoral politics: Gaddi grazing, degradation, and biodiersity
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33
1er Braak, C. J. l. ,1986, Canonical Correspondence Analysis - a New Ligenector
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\ang, \. l., lang, J. \., Pan, \. D. & Ji, C. J. ,2009, Aboeground biomass in 1ibetan
grasslands. -RXUQDORI$ULG(QYLURQPHQWV 91-95.

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1
Resource Lcology Group, Centre or Lcosystem Studies, \ageningen Uniersity,
Droeendaalsesteeg 3a, Lumen-100, 608 PB \ageningen, 1he Netherlands
&
2
Karnataka, India
Chapter 3
36
$EVWUDFW
1he 1rans-limalayan region o Ladakh harbours 20 species o mammalian wild
herbiores representing 6 amilies and 11 genera. \e studied their distributional patterns
and asked i they orm chorotypes or groups o species with similar distributions. \e
sampled presence-absence o 20 mammalian herbiores rom 39 quadrats or operatie
geographical units ,OGU, 20 x 20 km,. A probabilistic similarity technique was used to
look or herbiore species with distributions more similar than expected at random.
Baroni-Urbani & Buser`s similarity index was applied to the presence-absence data o
herbiores in the OGUs, and the Unweighted Pair-Group Method using Arithmetic
Aerage ,UPGMA, was used to classiy the species according to their distributions.
Statistical signiicances o the groups obtained were assessed by G-test o independence.
\e used the Canonical Correspondence Analysis ,CCA, to explore the relationship
between enironmental ariables and the chorotypes. \e obtained six chorotypes
including two large ones, o which one was composed o nine species, and occurred in
the open high plains o eastern Ladakh. 1he other comprised six species and occurred in
the rugged mountains o north and south-western Ladakh. Seenty six percent o the
ariation in species distribution was explained by enironmental ariables. Some o the
enironmental ariables were spatially structured, and a partial constrained ordination
showed that eight percent o the ariability in species distribution is explained by non-
spatial enironmental ariation, 55 by spatially structured enironmental ariation and
13 by pure spatial ariation. 1he remaining 24 o the ariation was unexplained by
the ariables used, suggesting the importance o riers, high mountains and other
historical actors as geographical barriers hindering animal moements.

.H\ZRUGV Mountain ungulates, rodents, lagomorphs, chorotype, UPGMA,
biogeography, 1rans-limalaya
Zoogeography of Ladakh
37
,QWURGXFWLRQ
1he distributional patterns o species and the underlying processes are important aspects
o biogeography ,Myers & Giller, 1995,. Species either occur alone, responding
dierently to enironmental gradients or they share ranges. In the latter case, speciic
groups can be classiied into biogeographical units corresponding to enironmental
,Gmez-GonzalezHWDO, 2004, as well as historical actors ,Myers & Giller, 1995,. Studies
on congruence or shared geographical ranges hae been carried out extensiely in seeral
taxa ,Marquez et al., 199, Gmez-Gonzalez et al., 2004, Oertli et al., 2005,, but is poorly
understood in mammalian herbiores, especially large herbiores such as ungulates.
Gien that large herbiores occupy almost 50 o the earth`s surace, and many o them
hae historically been useul to humans, more so since the dawn o ciilisation
,Diamond, 199,, there is an urgent need to understand their biogeography.
Analytical biogeography, applying hypothetico-deductie approach, is an
objectie way o studying distributional patterns o animals ,Myers & Giller, 1995,. 1he
distributional associations o species, termed Chorotype` ,Baroni-Urbani et al., 198,,
hae oten been studied by probabilistic approaches using similarity indices ,Real &
Vargas, 1996, Armenteras et al., 2003,, which are deemed crucial or understanding the
distribution o animal assemblages at large geographical scales. Cross-taxon analysis o
congruence or shared distributions o species is also helpul in ealuating the suitability
o some taxa or guilds as surrogate or others while prioritizing conseration areas with
limited time and budget ,\illiams & Gaston, 1994, Balmord & Long, 1995, \arman et
al., 2004,. 1his is especially releant in gathering inormation on biodiersity in
mountainous regions where species inentorying is constrained by terrain ruggedness and
inaccessibility.
1he mountainous region o Ladakh in northern India supports a relatiely dierse
assemblage o 20 mammalian herbiorous species representing 6 amilies and 11 genera
,lox et al., 1991a, Pister, 2004,. Most o these are Palaearctic, a trend also obsered in
the region`s birds ,Namgail & \om-1o, 2009,. Perhaps because the tropical lowland
organisms do not oercome high mountain passes o the 1rans-limalaya ,see Janzen,
196,. In any case, the mammalian herbiore assemblage includes twele wild ungulates
including endangered species like the 1ibetan antelope 3DQWKRORSV KRGJVRQL and Ladakh
urial 2YLVYLJQHLYLJQHLand also seeral rodents and lagomorphs ,Pister, 2004,.
Although some studies were carried out to understand the status and distribution
o wild ungulates in Ladakh ,lox et al., 1991a, Mallon, 1991, Chundawat & Qureshi,
1999, Bhatnagar & \angchuk, 2001,, inormation on their precise distributional ranges
are lacking. On the basis o limited ecological inormation, the distributions o these
ungulates were mapped through habitat suitability modeling ,Chundawat & Qureshi,
1999,, which still need to be ground-truthed across entire Ladakh. 1he distribution and
diersity patterns o smaller herbiores such as rodents and lagomorphs, howeer,
remain unknown except or some prominent places in the protected areas. lurthermore,
een in the case o ungulates, apart rom ew studies on auteecology and coexistence o
species pairs, hitherto no apparent eort has been made to understand the assembly
patterns at a multispecies leel. Such inormation are, howeer, crucial or a
comprehensie understanding o the Zoogeography o Ladakh.
Macro-leel distributional patterns o species and the underlying mechanisms
were oten inestigated in distinct ecosystems and communities ,lolland et al., 1991,
Risser, 1995,, but remain obscured or transition areas where one biome meets another.
1ransition areas, known as ecotones, are thought to be richer in biodiersity as they are
more dierse in habitats, thereby supporting a greater number o species. Lcotones are
thus thought to be dynamic biodiersity-hotspots where noel and unique eolutionary
orms are generated ,Kark & an Rensburgt, 2006,.
Chapter 3
38
Ladakh ,32
o
to 36
o
N and 5
o
to 80
o
L, is located at the junction o two biomes:
the 1ibetan plateau in the east and lindukush-Karakoram mountains ,including Ladakh
and Zangskar ranges, in the west. 1he ormer is characterised by ast open plains
interspersed with rocky outcrops, while the latter is characterised by rugged and broken
terrain. 1hus, Ladakh`s mammalian herbiores are perhaps adapted to two types o
terrain eatures rom these biomes ,Mallon, 1991, Pister, 2004,. loweer, apart rom
anecdotally, it is not known as to whether the species belonging to the two biomes orm
either distinct geographical groups corresponding to their eolutionary histories or
indiidual species rom dierent biomes respond independently to enironmental
gradients. Although some studies hae been carried out on local scales, as mentioned
earlier, assessing the coexistence o species pairs ,Namgail et al., 2004, Namgail, 2006b,,
there has been no large scale study encompassing multi-species at larger spatial scales.
1hereore, the present study inestigated the association,dissociation o mammalian
herbiores with dierent biogeographic ainities at the entire Ladakh leel. Recognising
the apparent lack o knowledge on smaller herbiores in Ladakh, we also gathered
inormation on their distributions and included them in the analysis.
As ar as the ecology o the wild ungulates are concerned, only a ew studies hae
been carried out ,Namgail et al., 2004, Bhatnagar et al., 2006b, Namgail, 2006b, in
Ladakh, as alluded to earlier, although inormation is also aailable rom studies on some
species like the blue sheep 3VHXGRLV QD\DXU and Asiatic ibex &DSUD LEH[ rom the
neighbouring Spiti alley, which addressed the interaction between these and the
domestic liestock ,Johnsingh et al., 1999, Bagchi et al., 2004, Mishra et al., 2004,.
Anecdotal inormation and limited studies on single or species-pairs addressing habitat
use hae suggested that the large herbiore species in the region partition space on the
basis o altitude, slope angle and distance to cli ,Mallon, 1991, lox et al., 1992,
Chundawat & Qureshi, 1999, Namgail et al., 2004, Namgail, 2006b,. Chundawat and
Qureshi ,1999, identiied these ariables as the most important actors determining the
distribution o wild ungulates in Ladakh. 1here is not much inormation on the habitat
use o wild yak %RVPXWXV1ibetan antelope and the 1ibetan gazelle 3URFDSUDSLFWLFDXGDWDin
Ladakh, but studies rom the 1ibetan plateau showed that these animals partition
resources along an altitudinal gradient ,larris & Miller, 1995,.
1he scanty dietary inormation on these ungulates suggest that amongst the our
Caprinae species, 1ibetan argali 2YLV DPPRQ KRGJVRQL and blue sheep include a higher
proportion o graminoids such as &DUH[ spp. and 6WLSD spp. in their diets, while the
Ladakh urial and Asiatic ibex incorporate a higher proportion o non-graminoids in their
diet ,Johnsingh et al., 1999, Mishra et al., 2004, also see Van den 1empel & De Vrij,
2006,. 1he 1ibetan gazelle and 1ibetan antelope are known to eed mainly on non-
graminoids such as orbs except the male 1ibetan antelope during summer ,larris &
Miller, 1995,. 1he 1ibetan wild ass and the wild yak are known to eed predominantly on
graminoids such as 6WLSD &DUH[ and .REUHVLD ,larris & Miller, 1995, Schaller, 1998,.
Nonetheless, the inormation on habitat and diet o the smaller herbiores remain
elusie, as alluded to earlier, although they are urgently needed, as many o these smaller
herbiores ace persecution by humans due to a preconception that they degrade
pastures ,Bagchi et al., 2006,.
So what does all these niche utilisation patterns o large herbiores and the
landscape coniguration in Ladakh tell us about their distribution in the region Can the
distribution o landscape eatures explain the distributional patterns o these large
herbiores on the basis o their niche preerences Do species rom dierent biomes
respond dierently to the enironmental conditions in Ladakh or they adapt to noel
conditions, thereby conerging in their resource use pattern irrespectie o their
eolutionary histories I so, then do they share distributions Can we also say something
39
about the distribution and diersity patterns o the smaller herbiores on the basis o the
distribution o the larger ones ,2 kg, like the ungulates. 1hese were some the questions
that stimulated us to start this inestigation. 1hese are interesting research questions that
need to be answered in order to understand the oerall herbiore assembly pattern in a
transition area like Ladakh.
Gien the paramount importance o topographical eatures in the distribution o
these herbiores ,Chundawat & Qureshi, 1999, Namgail et al., 2004,, we largely ocused
on the role o physical actors in understanding the congruence, or lack thereo, in
distribution o herbiores rom the two biomes. Although egetation is more important
than climatic ones in determining the distribution o herbiores, especially at smaller
spatial scales, we did not use egetation as an explanatory actor in this study because
there is no reliable egetation map or the region. 1he egetation distribution in Ladakh
is ery sparse, and there are no trees except along some rier banks ,Kachroo et al., 19,
Rawat & Adhikari, 2005b,. 1his and the barren nature o the landscape make it diicult
to get remotely sensed data on egetation. \e, howeer, inestigated the inluence o
climatic actors such as temperature and precipitation, as proxy actors, that can explain
the large herbiore distributions.
1hus, gien the act that there is a topographical gradient, and species rom
dierent biogeographic zones might respond dierently to such gradients leading to
association o some species, while dissociation o others as obsered between species
pairs, we hypothesised that mammalian herbiores o Ladakh orm geographical groups
on the basis o their biogeographic ainities, and physical and climatic ariables like
altitude and slope angle determine such grouping patterns. \e explore the importance o
major riers in determining geographical distribution o mammalian herbiores in
Ladakh, and also discuss the possible role o mountain ranges in their distributions.

6WXG\DUHD
Ladakh is the biggest proince, encompassing an area o F 80,000 km
2
, in the north-
Indian state o Jammu and Kashmir. 1he region has some o the highest mountain
ranges in the world, and the largest stretch o glaciers beyond the polar region ,Namgail,
2003,. Altitude in the region ranges rom 3000 to 600 m, proiding a ariety o habitats
or mammalian herbiores. 1rans-limalaya, o which Ladakh is a part, came into being
as a result o the collision between the Indian and Lurasian plates F35 million years ago
,DeweyHWDO, 1988,. 1he subsequent buckling o the Indian plate under the Lurasian one
uplited the limalaya and the 1ibetan plateau, leaing its legacy on the present
distribution o lora and auna. 1he tectonic moement and orogenic processes
inluenced the climate o the 1rans-limalaya making it progressiely drier ,Ramstein et
al., 199,, and a predominantly endemic assemblage o herbiores deeloped in the
region ,Schaller, 1998,. lollowing the initial rise o the limalaya, seeral species
especially o the amily Boidae underwent adaptie radiations by eoling ecological and
phenotypic diersities, occupying the newly created mountainous niches ,Ropiquet &
lassanin, 2005,.
Ladakh has three mountain ranges: Zangskar, Ladakh and Karakoram that run
almost parallel to one another and a network o riers and streams drain these ranges.
1he soil is sandy or sandy-loam, and are generally characterized by poor in organic matter
and nitrogen content ,Murti, 2001,. 1here is no orest coer, and the egetation is desert-
like, consisting o low shrubs and herbs ,Rawat & Adhikari, 2005a,. Some trees like
poplar 3RSXOXVspp. and willow 6DOL[spp. grow along rier-alleys. 1he rangelands o the
region are also characterised by low graminoid biomass ,Mishra, 2001,. Ladakh is a ery
remote region in India, and remains cut-o rom the rest o the country or almost seen
months o the year due to heay snow on the high passes. Due to the inhospitable
Chapter 3
40
enironment and low productiity, the human population is low with a density o less
than three persons,km
2
, which is the lowest or India.
'DWDFROOHFWLRQ
\e collected presence-absence data o mammalian herbiores rom 39 quadrats or
operatie geographical units ,OGUs, o 20 x 20 km
2
. 1his quadrat-size was chosen to
strike a balance between coering a maximum geographical area and capturing the
essence o possible inluence o enironmental actors on the shared distributions
amongst species. Although, we sureyed 95 quadrats, the remaining 56 were not analysed
as we could not surey them thoroughly or smaller mammals, and there was little
inormation in the literature. It was especially diicult to establish the absence o small
herbiores such as pikas 2FKRWRQD spp. and oles $OWLFROD spp., which are less isible
compared to the larger ones: ungulates. 1he 39 quadrats used, howeer, spanned entire
Ladakh and thus had a representatie sample rom each landscape unit: rugged terrain
and open plain.
Although we opportunistically accumulated inormation on species presence-
absence in the 39 quadrats oer a period o seen years beginning in early 2000, proper
sureys in the OGUs were carried out between May 2006 and June 2008. Animals were
located rom trails and antage points. Apart rom direct obserations, presence o body
parts such as horns o ungulates was recorded as presence o a species ,de Vrij & an den
1empel, 2006,. 1he occurrence o ossorial herbiores such as marmots and pikas were
determined also by locating burrows ollowed by intensie search to establish their
presence. Inormation on herbiore occurrence in the quadrats was gathered also by
interiewing wildlie oicials and researchers working in dierent areas. \e triangulated
such inormation by carrying out a questionnaire surey through post, whereby people
rom illages in or nearby a quadrat illed out a orm on presence-absence o animals and
sent it back to us. Apart rom these, we also used data rom published sources ,e.g., lox
et al., 1991a, Mallon, 1991, Pister, 2004,. Such multiple sources o data ensured a
thorough sampling rom each quadrat.
1o look at the relationship between enironmental ariables ,physical and
climatic, and chorotypes, we recorded mean altitude ,n ~ 50, and slope angle ,n ~ 50, an
index o terrain ruggedness, rom each quadrat. 1o detect geographical trends in the
distributions o chorotypes, we also took into consideration the geographical longitude
and latitude o the quadrats. 1he climatic ariables ,mean annual temperature and mean
annual precipitation, at 1 km
2
resolution were obtained rom the \orldClim database
,lijmans et al., 2005,. \e obtained these ariables in a GIS ,Geographic Inormation
System, ArcView, New \ork, USA, enironment ,LSRI, 1996,.
1here are two major riers: Indus and Zangskar that could inluence the
zoogeography o Ladakh. 1he ormer runs rom east to west diiding Ladakh into two
almost equal hales, while the Zangskar runs rom south to north, diiding Ladakh in
western and eastern parts, beore it drains into Indus. 1hereore, we also assessed their
importance as geographical barriers. lor this, we recorded the number o species that
occur on one side o the rier but not on the other, and ealuated whether these riers
restrict the range o mammalian herbiores in Ladakh.
6WDWLVWLFDO$QDO\VLV
&KRURW\SHV
1o classiy species according to their distributions, we used the similarity index o
Baroni-Urbani & Duser ,196, to each pair o species a and b as

41

C B A D C
C D C
B
+ + +
+
=
.
.
Lqn. 1
where $ is the number o quadrats where only species D is present, % is where only
species Eis present, &is where both are present, and 'is the number o quadrats rom
which both are absent.
A dendrogram o the biogeographical relationship between species was obtained
by applying the Unweighted Pair-Group Method using Arithmetic Aerage ,UPGMA, to
the matrix o alues o Baroni-Urbani & Buser`s similarity index. 1his procedure is an
agglomeratie one that produces the lowest distortion in relation to the original distances
between samples ,Sneath & Sokal, 193,. 1he statistical signiicance o each similarity
alue was ealuated using the statistical table in Baroni-Urbani & Duser ,196,. In this
way a matrix o signiicant similarities was generated by substituting the similarity alues
with a -`, -`, or 0` sign when the similarities were, respectiely, higher, lower or similar
than would be expected at random. \e then conerted the similarity matrix o species
locations to a secondary matrix o similarity coeicients.
\e used G-tests to statistically test the signiicance o each group in the
dendrogram ,McCoy et al., 1986,. 1his method distinguished species that occur together
more requently than expected at random rom those that are artiacts o the cluster
analysis. 1he null hypothesis was that no geographical groups or chorotypes are ormed
by the 1rans-limalayan mammalian herbiores. lor each comparison, we took a sub-
matrix, ,corresponding to a sub-group in the dendrogram, o signiicant similarities,
which we diided into three zones: A and B corresponding to each group compared, and
zone AxB corresponding to the intersection o both groups.
\e considered species to orm chorotypes or show similarity in distribution
when signiicantly similar species ,-, tended to be located in zones A and B, but not in A
x B. I, on the other hand, the signiicantly less similar species ,-, occurred in zone A x B,
but not in A or B, we considered the two groups o species to be strongly segregated.
1he parameters D\ ,A x A, and D\ ,B x B, measure the internal homogeneity o each
group, whereas DS measures strong segregation between groups. A group o species was
considered to be a weakly segregated chorotype when D\0 and G\ was statistically
signiicant, and a strongly segregated one when DS0 and GS was statistically signiicant.
1his procedure determines whether the clusters identiied hae any biogeographical
meaning, or are the artiacts o the analysis haing to orm clusters.
2UGLQDWLRQPHWKRG
1he geographical communities or chorotypes could be inluenced by ecological as well as
historical actors. \e used the Canonical Correspondence Analysis ,CCA, to explore the
relationship between the identiied chorotypes and the enironmental ariables, putting
all species in a continuous enironmental perspectie. 1his method is useul in measuring
the amount o ariation in the species distribution data that can be explained by dierent
explanatory ariables. 1he statistical signiicance o this relationship was assessed using a
permutation test, which has an added adantage o the ability to check which ariables
contribute most to the relationship. 1he analysis was carried out using the CANOCO
sotware ,1er Braak, 1986,.
Since enironmental ariables could be spatially structured, and conound the
results, we perormed the so-called partial constrained ordinations suggested by ,Borcard
et al., 1992,, and controlled or a second set o ariables by using them as coariates.
1hereore, we carried out our types o CCA: ,1, between species and enironmental
ariables, ,2, between species and enironmental ariables with spatial ariables ,latitude
Chapter 3
42
and longitude, as coariates, ,3, between species and spatial ariables and ,4, between
species and spatial ariables with enironmental ariables as coariates.
5HVXOWV
\e sampled 20 mammalian herbiores rom the 39 OGUs. 1hese comprised eight
ungulates, seen lagomorphs and ie rodents ,1able 1,. lity percent o the species ,n ~
20, were present in more than 10 OGUs. Blue sheep was the most common mammalian
herbiore occurring in more than 25 OGUs, ollowed by the Asiatic ibex &DSUD LEH[
VLEHULFD and long-tailed marmot 0DUPRWD FDXGDWD ,lig. 1,. 1he similarity between species
distributions are shown by the dendrogram created through UPGMA ,lig. 2,. 1he matrix
o signiicant similarity between species distributions are presented in 1able 2. 1he
signiicant segregations o species based on their distributions are presented in 1able 3.
&KRURW\SHV
A total o six chorotypes were identiied ,lig. 2,. Chorotype I was composed o only one
species: 1ibetan antelope 3DQWKRORSVKRGJVRQLwhich had signiicantly dierent distribution
rom all other herbiores. Chorotype II was strongly segregated rom all others, and
comprised six species: long-tailed marmot ,abbreiated as Lo1 Mar,, Asiatic ibex ,Asi
Ibx,, Large-eared pika ,LaL Pik, 2FKRWRQDPDFURWLV, Nubra pika ,Nub Pik, 2FKRWRQDQXEULFD,
Cape hare ,Cap lar, /HSXVFDSHQVLV and Ladakh urial ,Lad Uri, lig. 2, 1able 3,. Chorotype
III again comprised one species: wild yak ,\il \ak, with a distribution unique to itsel.
Chorotype IV was ormed by the Royle`s pika ,Roy Pik, 2FKRWRQD UR\OHL and Royle`s
mountain ole ,RoM Vol, $OWLFROD UR\OHL, while Chorotype V comprised one species:
1ibetan gazelle ,1ib Gaz, 3URFDSUD SLFWLFDXGDWD although it was weakly segregated rom
others.

0
5
10
15
20
25
30
B
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ik
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l
a

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N
u
b

P
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Species
N
u
m
b
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o
f

O
G
U
s

lig. 1. Number o operatie geographical units ,OGUs, occupied by mammalian
herbiores in the Ladakh 1rans-limalaya. See 1able 1 or species identity.

43

Chorotype VI was the largest o all with nine species that included Silery mountain ole
,SiM Vol, $OWLFROD DUJHQWDWXV, 1ibetan wooly hare ,\ol lar, /HSXV RLRVWROXV, limalayan
marmot ,lim Mar, 0DUPRWDEREDN, 1ibetan wild ass ,\il Ass, (TXXVNLDQJ, Ladakh pika
,Lad Pik, 2FKRWRQD ODGDFHQVLV, Plateau pika ,Pla Pik, 2FKRWRQD FXU]RQLDH, Stoliczka`s
mountain ole ,StM Vol, $OWLFRODVWROLF]NDQXV, 1ibetan argali ,1ib Arg, 2YLVDPPRQKRGJVRQL
and blue sheep ,Blu She, lig. 2,.

1able 1. Mammalian herbiores in the Ladakh 1rans-limalaya and their abbreiation
used in the study.
2UGHUlamily 6FLHQWLILFQDPH &RPPRQQDPH $EEU

$UWLRGDFW\OD
Boidae &DSUDLEH[VLEHULFD Asiatic ibex Asi Ibx
2YLVDPPRQKRGJVRQL 1ibetan argali 1ib Arg
2YLVYLJQHLYLJQHL Ladakh urial Lad Uri
3VHXGRLVQD\DXU Blue sheep Blu She
3DQWKRORSVKRGJVRQL 1ibetan antelope 1ib Ant
3URFDSUDSLFWLFDXGDWD 1ibetan gazelle 1ib Gaz
%RVPXWXV \ild yak \il \ak
3HULVVRGDFW\OD
Lquidae (TXXVNLDQJ 1ibetan wild ass \il Ass
5RGHQWLD
Sciuridae 0DUPRWDFDXGDWD Long-tailed marmot Lo1 Mar
0DUPRWDEREDNKLPDOD\DQD limalayan marmot lim Mar
Muridae $OWLFRODUR\OHL Royle's mountain ole RoM Vol
$OWLFRODDUJHQWDWXV Silery mountain ole SiM Vol
$OWLFRODVWROLF]NDQXV Stoliczka's mountain ole StM Vol
/DJRPRUSKD
Leporidae /HSXVRLRVWROXV 1ibetan woolly hare \ol lar
/HSXVFDSHQVLV Cape hare Cap lar
Ochotonidae 2FKRWRQDFXU]RQLDH Plateau pika Pla Pik
2FKRWRQDODGDFHQVLV Ladakh pika Lad Pik
2FKRWRQDPDFURWLV Large-eared pika LaL Pik
2FKRWRQDQXEULFD Nubra pika Nub Pik
2FKRWRQDUR\OHL Royle's pika Roy Pik

Chapter 3
44

lig. 2. Classiication dendrogram o the distribution o mammalian herbiores in Ladakh.
Species in bold represent chorotypes o single species and roman numerals indicate
chorotype number, see 1able 1 or species identity. S is strong and \ is weak segregation
amongst species distributions. P 0.01, P 0.005.
2UGLQDWLRQRIVSHFLHV
Lnironmental ariables inluenced the distribution o the chorotypes as their
relationship established by CCA was statistically signiicant ,l-ratio ~ 4.5, p ~ 0.002,.
1hey explained 63 o the ariation in geographical distributions o species. 1he irst
axis ordered mammalian herbiores along a gradient o decreasing altitude and increasing
slope angle, while the second axis ordered species along a gradient o decreasing
precipitation and increasing temperature. 1his ordering indicated that Asiatic ibex, long-
tailed marmot, Ladakh urial, Nubra pika and cape hare occur in steeper areas at lower
altitude with high temperature ,lig. 3,. Large-eared pika and Royle`s mountain ole occur
in areas with high precipitation, while 1ibetan argali, 1ibetan wild ass, limalayan
marmot, 1ibetan wooly hare, Stoliczka`s mountain ole and silery mountain ole occur
in latter and drier areas at high altitude with less temperature ,lig. 3,. Precipitation and
altitude contribute little along the two ordination axes ,lig. 4,, suggesting that these
ariables are spatially structured. In igure 4, species are arranged by increasing altitude
and decreasing slope angle along axis one, while they are ordered along a gradient o
decreasing temperature and precipitation on the second axis.
Some o the species are related dierently to enironmental ariables ater
remoing the eect o spatial structure. lor example, in the irst analysis without the
spatial ariable as coariate ,lig. 3,, blue sheep was grouped with 1ibetan argali, 1ibetan
wooly hare and limalayan marmot, but it is associated more closely with Ladakh urial
and cape hare ater the spatial structure was eliminated ,lig. 4,.
Blu She
Tib Arg
StM Vol
Pla Pik
Lad Pik
Wil Ass
Him Mar
Wol Har
SiM Vol
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RoM Vol
Roy Pik
Wil Yak
Lad Uri
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Asi Ibx
LoT Mar
Tib Ant
W**
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47
1able 4. Results o the CCAs perormed excluding the eects o dierent set o
ariables.
Lxplanatory ariables Coariables explained 3
Lnironmental None 63 0.01
Lnironmental Spatial 8 0.01
Spatial None 8 0.01
Spatial Lnironmental 13 0.01

A CCA between species and spatial ariables accounted or 8 o the ariation in the
species distribution data ,1able 4,. lrom lig. 5, three prominent groups can be
distinguished: ,1, Large-eared pika, Nubra pika, Ladakh urial and cape hare occur in
northern part o Ladakh, ,2, Asiatic ibex, long-tailed marmot, Royle`s pika and Royle`s
mountain ole are distributed in the southwestern part o Ladakh, while ,3, 1ibetan
gazelle, Ladakh pika, plateau pika, 1ibetan argali, 1ibetan wooly hare, Stoliczka`s
mountain ole, silery mountain ole, limalayan marmot and 1ibetan wild ass occur in
the eastern part o Ladakh. Blue sheep has a broad distribution with no apparent
geographical trend as indicated by its position near the intersection o the two axes.
\hen enironmental ariables were used in the CCA as coariates, spatial
ariables accounted or 13 o the ariation in species distribution data ,1able 4, lig 6,.
Species ordered along the two axes leads to almost the same groups as in lig. 5,
suggesting that species distribution is not completely explained by spatial ariation in
enironmental data. 1he percentage o total ariation in the species distribution
accounted or by each component o the partial constrained ordination was as ollows:
eight percent by non-spatial enironmental ariation, 55 by spatially structured
enironmental ariation and 13 by pure spatial ariation. 1he remaining 24 o the
ariation was unexplained by these ariables.
-1.0 1.0
-
0
.
8
0
.
4
Blu She
Lad Uri
Asi Ibx
Tib Arg
Wil Ass
Tib Gaz
LoT Mar
Him Mar
Wol Har
Cap Har
SiM Vol
RoM Vol
StM Vol
Pla Pik
Lad Pik
Nub Pik
LaE Pik
Roy Pik
ALT
SA
TEMP
PREC

lig. 3. lirst two axes o CCA ordination o the species and enironmental ariables ,with
no coariables,. See 1able 1 or species identity. ,SA ~ Slope angle, Alt ~ Altitude,
PRLC ~ Precipitation, 1LMP ~ 1emperature,.
Chapter 3
48
5LYHUVDVJHRJUDSKLFDOEDUULHUV
1wo species o lagomorphs: cape hare and Nubra pika, which constitute 29 o this
taxon ,n ~ , ound in Ladakh, occurred only in quadrats north o the Indus Rier, while
all other mammalian herbiores occurred on both sides o the rier. Similarly, ie
ungulates ,62 o the taxon ound in Ladakh, n ~ 8,, two rodents ,40, n ~ 5, and our
lagomorphs ,1, n ~ , occurred only in quadrats east o the Zangskar Rier, while all
others occurred on both sides o the rier.

'LVFXVVLRQ
1he biogeographical characteristics o Ladakh 1rans-limalaya is unique with an
interesting assemblage o mammalian herbiores adapted to dierse habitats. 1he present
study has shown that the mammalian herbiores in the region orm groups based on
their biogeographic ainities, which was hitherto presumed but not tested. 1he 1rans-
limalaya is thought to be the cradle o Caprinae eolution ,Schaller, 19,, and Ladakh
1rans-limalaya is one o the ew areas in central Asia that harbour the highest number
o species belonging to this taxon. 1hus, Ladakh is an important biogeographic unit rom
both ecological and eolutionary perspecties. Although biogeographical analysis o
mammals hae been carried out elsewhere ,Badgley & lox, 2000, l. leikinheimo et al.,
200,, this study, to our knowledge, is the irst biogeographical analysis exclusiely o
mammalian herbiores including ungulates and has important implications or a
mechanistic understanding o the distribution o wild ungulates in the 1rans-limalaya
and other alpine rangeland areas.

-0.8 0.8
-
0
.
6
0
.
8
Blu She
Lad Uri
Asi Ibx
Tib Arg
Wil Ass
Tib Gaz
LoT Mar
Him Mar
Wol Har
Cap Har
SiM Vol
RoM Vol
StM Vol
Pla Pik
Lad Pik
Nub Pik
LaE Pik
Roy Pik
ALT
SA
TEMP
PREC

lig. 4. lirst two axes o CCA ordination o the species and enironmental ariables
,explanatory, with spatial ariables ,coariates,. See 1able 1 or species identity. 1he
abbreiations o enironmental ariables are same as in lig. 1.
49
It seems that there are two main biogeographic units o mammalian herbiores in the
region, as indicated by the strong segregation between chorotype II comprising six
species and all the other species, nine o which belong to chorotype VI. Chorotype II
belongs to the north and south-western rugged parts o Ladakh, while species in other
chorotypes correspond to the eastern plain o Changthang. 1hese conorm to the
anecdotal inormation on their distributional patterns in Ladakh ,lox et al., 1991a,
Mallon, 1991,. 1he enironmental ariables that best characterise Chorotype II are
rugged terrain with steep slopes, typical o north and south-western part o Ladakh. In
chorotype II, Asiatic ibex and long-tailed marmot are more similar in their distributions
than the rest o the members. \hen looked at their global distributions, both seem to
hae relatiely wide and oerlapping distributions in central Asia, with the ibex haing
somewhat wider distribution. 1he areas in Ladakh with the core o their ranges are
characterised by rugged, relatiely lower and thus higher temperature. Ibex distribution in
central Asia seems to be determined largely by topography, as it is known to use steep
clis as escape terrain ,lox et al., 1992, Namgail, 2006b,, but the role o terrain
ruggedness in the habitat selection o long-tailed marmot is not known. 1here could be
another actor that is causing the conergence o their distribution, which can only be
known by studying their habitat utilization patterns in detail.
1here are three chorotypes each with a single species, which hae ery restricted
distributions unique to themseles. 1hese are the 1ibetan gazelle, 1ibetan antelope and
wild yak. Both wild yak and 1ibetan gazelle aced precipitous decline in their populations
in Ladakh during the last century, excessie hunting and liestock grazing hae been
listed as the most important causes o these declines ,Bhatnagar et al., 2006a, Namgail et
al., 200, Namgail et al., 2008,. 1he 1ibetan antelope, on the other hand, did not hae a
wide distribution in Ladakh in the past too. 1he animal is known or its long latitudinal
migration, and perhaps it does not hae a bigger population in Ladakh because there is
not enough open areas in a north-south orientation in this region, as it is bounded by the
Karakoram range on the north and the Great limalayan range on the south.
Chorotype IV is ormed by the Royle`s pika and Royle`s mountain ole. 1here is
no inormation on the ecology o these species in Ladakh, except anecdotal inormation
about their distribution in ew areas ,Pister, 2004,. 1heir global distribution is, howeer,
restricted to the limalayan region ,Smith & Boyer, 2008,, where their distributions span
an altitudinal range between 2500 and 4300 m asl. lurther, mountain ole is endemic to
India and is the only small mammlian herbiore o Ladakh that is listed as Near
1hreatened` on the IUCN red list o threatened animals, as all others are listed as Least
Concern` ,IUCN, 2008,. Anecdotal inormation suggests that both Royle`s pika and ole
occur in rocky habitats in Ladakh ,Pister, 2004, at lower altitudes with slightly higher
precipitation ,pers. obs.,, perhaps due to the spatial structuring in enironmental
ariables ,see discussion below,. 1he role o temperature and precipitation in their
distributions is also indicated by the CCA results.
All other species are clubbed into chorotype VI, in which some species are more
similar in distribution than others, although not signiicantly. All the species in this
chorotype hae the core o their ranges on the 1ibetan plateau except or three species:
silery mountain ole, limalayan marmot and Ladakh pika, which are distributed mainly
in the marginal mountains o the plateau ,Molur, 2008, Molur & Shreshtha, 2008, Smith
& Johnston, 2008,. \ithin this chorotype, woolly hare and limalayan marot are more
similar to each other than they are to others. Globally both species hae similar
distributions along an altitudinal gradient ,Molur & Shreshtha, 2008,, thus altitude might
be a actor that is leading to the congruence in their distributions in Ladakh, which needs
to be explored. lurther, within this chorotype, blue sheep stands apart rom others,
indicating that although it shares habitat eatures with other members o the chorotype,
Chapter 3
50
its distribution is idiosyncratic in some ways. 1his could be related to its ubiquity, as it is
the most abundant and widely distributed wild ungulate in Ladakh as indicated by the
CCA analysis ,see also lox et al., 1991a, Namgail, 2006b,.
1he congruence pattern o mammalian herbiore distribution in Ladakh indicates
that their distributions are inluenced by both current ecological and historical actors, as
the enironmental actors did not explain all the the geographical groups obtained in the
cluster analysis. A substantial ariation in the species distributional data was also
explained by the spatial ariables, suggesting that some enironmental ariables hae a
spatial structure and consequently their eect is conounded by the geography. 1he
results o the partially constrained ordinations suggest that altitude and precipitation hae
a latitudinal and,or longitudinal component. \hen the topographical and climatic maps
o Ladakh are examined closely, it is apparent that south-western part o Ladakh receies
more precipitation ,lartmann, 1983,, and the eastern part is higher than the rest.

-1.0 2.0
-
1
.
0
1
.
5
Blu She
Lad Uri
Asi Ibx
Tib Arg
Wil Ass
Tib Gaz
LoT Mar
Him Mar
Wol Har
Cap Har
SiM Vol
RoM Vol
StM Vol
Pla Pik
Lad Pik
Nub Pik
LaE Pik
Roy Pik

lig. 5. lirst two axes o CCA ordination o the species and spatial ariables. See 1able 1
or species identity.

1he 24 o the ariation in species distribution that remained unexplained suggests that
there are actors that inluence their distributions but were not included in this analysis,
as alluded to earlier. Such actors could be both current ecological and historical actors.
1he most plausible such actors are egetation, humans and their endeaours, riers and
high mountains as geographical barriers. One o the major caeats o this study is the
lack o inormation on the egetation, but this would not hae inluenced the oerall
conclusion o the study or a couple o reasons: 1, physical habitats explained most o
the ariation on distributional pattern o herbiores, especially the larger ones, 2, the
climatic ariables used in the study would hae accounted or the ariation in the
51
egetation distribution to some extent, as they inluence the egetation growth and
regeneration ,Chapter 2,.
1here are more mammalian herbiores in the eastern than in the western part o
Ladakh ,Pister, 2004,, as also shown by the cluster and CCA analysis. One wonders why
there is a discrepancy in species richness across the landscape. One o the main actors
accounting or this could be the topographical eatures, as indicated by the CCA. Because
the eastern part is latter and open that might acilitate ree moement o animals, while
the western part is more rugged with myriad glaciers that may restrict animal moement.
But there is no denial that anthropogenic pressures might also hae played a role, as
dierent species might respond dierently to gradients created by humans. 1he largest
chorotype occurred in the eastern part o Ladakh, which is the centre o 3DVKPLQD or
cashmere wool production in India. 1he people in this part o Ladakh are predominantly
nomadic pastoralists and rear a ariety o liestock including yak, horse, sheep and goat.
Could it be that there are more species o wild herbiores here than in the western part
because these liestock act as a buer or the wild herbiores against predators, implying
that the predators in the west, with declining liestock population ,Raghaan, 2003,, hae
a more controlling eect on wildlie populations. 1he leel o hunting by humans has
been reported to be higher in the western part ,Namgail, 2006a,, which might exacerbate
the aorementioned negatie inluence o predators. lurther discussion on this topic is
beyond the scope o this paper, but such issues need to be addressed to hae a
comprehensie understanding o the species richness pattern o mammalian herbiores
in Ladakh.
1he role o riers and high mountains in the herbiore species richness pattern in
Ladakh has neer been assessed. Gien that 24 o the ariability in the distributional
patterns o mammalian herbiores remain unexplained it is possible that the present
assemblage o large herbiores in Ladakh are shaped by these geographical eatures.
Some species like the cape hare and Nubra pika occurred only in quadrats north o the
Indus rier, which indicate that this rier could be restricting their southward expansion.
lurther, most o the species that hae the core o their ranges on the 1ibetan plateau
occured only on the eastern side o the Zangskar rier, suggesting that it might be a more
important determinant o Ladakh`s zoogeography. 1he rier`s inluence on animal
distributions might, howeer, be conounded by the terrain coniguration ,animals not
being able to adapt to rugged terrain,, as there is an interace between western rugged
terrain and eastern plains that runs almost parallel to the rier, although located approx.
30 km east o the rier.
1hereore, we looked or species that occur in the area between the rier and the
terrain interace to disentangle their eects, and ound one ungulate: 1ibetan argali and
three lagomorphs in this area, suggesting that Zangskar might constrain the distributional
ranges o these species, but not that o the others, which might be constrained by the
topography. \et, literature suggests that 1ibetan argali is also adapted to open areas
,NamgailHWDO, 2004,, and there is only one small population o this species within 10 km
rom the rier that colonised the area in 198 ,loxHWDO, 1991b,. 1hus, the riers do not
seem to inluence the distribution o large herbiores in Ladakh, perhaps because animals
cross the riers during winter when they reeze, although they might restrict the ranges o
some o the less mobile species such as smaller herbiores.

Chapter 3
52
-2.0 1.0
-
1
.
5
1
.
0
Blu She
Lad Uri
Asi Ibx
Tib Arg
Wil Ass
Tib Gaz
LoT Mar
Him Mar
Wol Har
Cap Har
SiM Vol
RoM Vol
StM Vol
Pla Pik
Lad Pik
Nub Pik
LaE Pik
Roy Pik

lig. 6. lirst two axes o CCA ordination o the species and spatial ariables ,explanatory,
and enironmental ariables ,coariates,. See 1able 1 or species identity.

1he unexplained ariation in the species distribution, thereore, could largely be
accounted or by historical actors associated perhaps with orogenesis. 1he rise o the
limalaya as a result o the collision o the Indian and Lurasian plates led to the
ormation o ormidable mountains in the region, which might hae acted as barriers,
conining species with similar modes o dispersal to similar areas. 1he limalaya is still
geologically actie, and the mountains are rising rapidly due to block aulting ,Valdiya,
1991,. 1hus, the geographical grouping o mammalian herbiores in Ladakh could, to
some extent, be attributed to the high and ormidable mountains that restrict their
moements. 1his perhaps is ouched by the act that the open plains o eastern Ladakh,
where animals can moe reely, is more dierse in mammalian herbiores compared with
the rugged areas o the western part.
lurthermore, the role o high and ormidable mountains in herbiore
distributions is perhaps also indicated by the absence o large herbiores in Ladakh rom
the Oriental region, because the limalayan range hindered their northward expansion
,see Chapter ,. Similarly, there are less species rom west Asia, and more rom the
1ibetan plateau in Ladakh`s regional herbiore species-pool, which is perhaps due to the
geographical barrier comprising a jumble o mountains to the west o Ladakh, that
respresent the end o seeral mountain ranges such as the Kunlun Shan, 1ian Shan and
the limalaya. Although these are just a ew acts to support the plausible role o high
mountain ranges inluencing Ladakh`s zoogeography, a thorough discussion on the issue
is beyond the scope o this paper. 1he results o this study merely sere as baseline
inormation or urther studies that will enhance our understanding o biogeography,
phylogeny and eolution o mountain ungulates.

53
Conservation implications
1he populations o many mammalian herbiores in Ladakh are declining precipitously
,Bhatnagar et al., 2006a, Namgail et al., 2009,. Amongst the eight wild ungulates, two are
listed as Lndangered`, two as Vulnerable` and one as Near 1hreatened` on the IUCN
red-list o threatened animals ,IUCN, 2008,. 1he status o less prominent taxa like
lagomorphs and rodents remains little known, although some species are known to be
persecuted by humans as pest control measures ,Bagchi et al., 2006,. 1his calls or
enhanced research, monitoring and conseration programs. Neertheless, the harsh
enironment and the diicult terrain make it diicult to deelop such programs. 1hus,
there is an urgent need or innoatie ways to study and consere this unique but
threatened group o animals. Our study has shown that there is cross-taxon congruence
in the distribution o the mammalian herbiores o Ladakh. Under such a scenario, larger
and easily located taxa like ungulates can be used as surrogates or other less prominent
taxa such as pikas and oles while planning conseration strategies and prioritising areas
or conseration o these high-altitude animals.

Acknowledgements
1he study was inancially supported by the \ildlie Conseration Society ,\CS,, the
Ruord Small Grants loundation and the \ageningen Uniersity. \e thank the Jammu
and Kashmir \ildlie Department or granting the necessary permission to surey
arious protected areas in the region. \e thank Otto Pister or sharing his inormation
on the distribution o mammalian herbiores in Ladakh. \e thank Charudutt Mishra and
\ash Veer Bhatnagar or their support. lield assistance o Karma Sonam and Mipham
are thankully acknowledged.

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1rans-limalaya. vrirovvevtat Mavagevevt, 39, 490-496.
Namgail, 1., lox, J. L. & Bhatnagar, \. V. ,2009, Status and distribution o the Near
1hreatened 1ibetan argali in Ladakh, India: eect o a hunting ban. Or,, 43,
288-291.
Namgail, 1. & \om-1o, \. ,2009, Lleational range and timing o breeding in the birds
o Ladakh: the eects o body mass, status and diet. ]ovrvat of Orvitbotog,, JS0,
505-510.
Oertli, S., Muller, A., Steiner, D., Breitenstein, A. & Dorn, S. ,2005, Cross-taxon
congruence o species diersity and community similarity among three insect taxa
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Dissertation, Saurashtra Uniersity.
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motion and land-sea distribution on Lurasian climate change oer the past 30
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Chapter 3
56
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Interaction between ibex and blue sheep
57
&+$37(5
:LQWHUKDELWDWSDUWLWLRQLQJEHWZHHQ$VLDWLFLEH[DQGEOXHVKHHSLQ
/DGDNKQRUWKHUQ,QGLD
1sewang Namgail

SDUWVRIWKH+LPDOD\DUHPDLQYLUWXDOO\XQNQRZQ]RRJHRJUDSKLFDOO\VSHDNLQJ7KHSUHFLVH
OLPLWV RI VSHFLHV KDYH VHOGRP EHHQ GHILQHG DQG WKH IDVFLQDWLQJ HFRORJLFDO SUREOHPV UDLVHG ZKHQ
WZRUHODWHGVSHFLHVRFFXS\WKHVDPHUDQJHKDYHEHHQODUJHO\LJQRUHG

George B. Schaller

Uniersity, Droeendaalsesteeg 3a, Lumen-100, 608 PB \ageningen, 1he Netherlands
&
Nature Conseration loundation, 306,5, IV Cross, Gokulam Park, Mysore - 50 002,
Karnataka, India
Chapter 4
58
$EVWUDFW
Asiatic ibex &DSUDLEH[VLEHULFDand blue sheep 3VHXGRLVQD\DXUare the most abundant wild
ungulates in the Ladakh Region o the Indian 1rans-limalaya. Both species use rugged
terrain to escape predation, and the competitie exclusion principle suggests that the
distribution o one species may be aected by the presence o the other. I ealuated
habitat use by these mountain ungulates in the Shun Gorge, at the eastern boundary o
ibex distribution in the Zangskar Mountains, Ladakh, India. I hypothesised that due to
their high ainity toward clis as a predator escape strategy, ibex and blue sheep oerlap
in their habitat use, especially in winter when they are likely to be conined by snow
coer. Resource selection indices and chi-square statistics reealed that both ibex and
blue sheep preer habitat close ,1-50 m, to clis. 1he two species were also similar in
their use o habitat in terms o slope angle, except that ibex aoided gentle slopes ,15
o
,
and blue sheep aoided ery steep slopes ,45
o
,. Both used habitats in terms o eleation
and snow coer non-selectiely except that blue sheep aoided ery low areas ,4000 m,,
and ibex aoided snow-ree areas. I suggest that there is high potential or competition
between the two species, and the presence o one species may negatiely inluence the
distributional pattern o the other.

.H\ZRUGVAsiatic ibex, &DSUDLEH[blue sheep, 3VHXGRLVQD\DXUresource selection, habitat
partitioning, Zangskar, Ladakh, 1rans-limalaya

59
,QWURGXFWLRQ
Large herbiores with similar ecological requirements are expected to partition resources
to coexist, especially in areas where the resources are in short supply ,\iens, 19, De
Boer & Prins, 1990,. Understanding the mechanism o such resource partitioning is a
central issue in community ecology ,Rickles, 1990,. Resource partitioning is
accomplished through eolutionary diergence o resource use by co-occurring species
,\alter, 1991,, apparently in response to competition. Neertheless, some ecologists
argue against the importance o competition in the dierential use o resources by
sympatric species, and contend that predation may also lead to niche dierentiation
,lairston et al., 1960, lolt, 19, Repasky, 1996,, as predatory risk proides an axis
along which habitat partitioning can occur ,lolt, 1989,. Neertheless, in areas where
competition does structure communities, species should segregate along at least one
resource dimension in the lutchinsonian niche hyperolume ,Schoener, 194,.
As per the competitie exclusion principle, when two species co-occur in an area
and depend on the same limited resources, the dominant species competitiely exclude
the subordinate one, as exempliied by Gause`s experimental work ,Gause, 1934,.
1hereore, the range o a species can be reduced by the presence o other species with
similar ecological requirements ,Connell, 1961,. Studies on resource partitioning by
ecologically similar wild ungulates were carried out extensiely in North America and
Arica ,Lamprey, 1963, Jarman & Sinclair, 199, lanley & lanley, 1982, Jenkins &
\right, 1988, Voeten & Prins, 1999,, but there is little inormation on the niche
separation amongst wild ungulates o the 1rans-limalayan ecosystem o south Asia.
\ildlie managers in this mountainous region, thereore, rely on inormation generated
rom studies in the tropical and sub-tropical regions, which may not necessarily be
applicable to this dry alpine ecosystem with a unique assemblage o large herbiores.
In this study, I looked at the dierential habitat use by two mountain ungulates:
Asiatic ibex &DSUD LEH[ VLEHULFD ,hereater ibex, and blue sheep 3VHXGRLV QD\DXU in the
Zangskar Mountains o Ladakh, India. 1hese are the most abundant wild ungulates in the
region with blue sheep`s population ,F 11,000 indiiduals, reaching almost double the
number o ibex ,lox et al.,1991,. Blue sheep also has a wider distribution in the region,
mostly in the eastern part and ibex has a relatiely narrower distribution in the western
part ,lox et al., 1991,. lox et al. ,1992,, howeer, reported a relatiely higher abundance
o ibex in central Ladakh ,despite its dryness and low egetation,, which they attributed
to the lack o snowpack that might had encouraged ibex populations to oerwinter in the
region. 1he two species constitute the most important prey species o the highly
endangered snow leopard 8QFLDXQFLD,Mallon, 1991, Oli et al.,1993,.
Ibex and blue sheep hae similar anti-predator habitat requirements, as both use
rugged terrain to escape predation ,Bhatnagar, 199, Namgail et al., 2004,. Preliminary
obserations on the habitat use by these species also reealed that they are similar in the
use o habitat ariables such as altitude, slope angle, rock type etc. ,Mallon, 1991,.
1hereore, it is possible that they compete or some resources, and the presence o one
species could negatiely aect the other species. 1here is anecdotal inormation on their
distributions that support such contentions, e.g., across large mountainous tracts o
northwestern 1ibetan Plateau, the distributions o these species demarcate oten
abruptly, thereby suggesting competitie exclusion o one species by the other ,Schaller,
1998,.
Blue sheep may use large boulders and scree o slate talus as escape terrain in
parts o the 1ibetan Plateau with low aailability o clis ,larris & Miller, 1995,.
lurthermore, in other areas where the two species co-occur, blue sheep are reported to
explore the open slopes near clis more oten than ibex ,\egge, 1989, pers. obs.,. 1he
ormer is thereore more lexible in its habitat use, and may hae a competitie adantage
Chapter 4
60
oer ibex. 1he Shun Gorge marks the eastern boundary o ibex distribution in the
Zangskar Mountains o the Indian 1rans-limalaya ,Namgail, 2004,, while along the
limalayan Range to the south its eastern limit terminates abruptly at the Sutlej Deile
,lox et al., 1992,. Based on the competitie exclusion principle, one can thereore
speculate that competition with blue sheep may limit its eastward distribution. A study
was thus carried out in the Shun Gorge to assess the degree o oerlap and possible
competition between these ecologically similar species. Based on their similar anti-
predator habitat requirements, I predicted a high oerlap between the two species in the
use o habitat, especially in winter when snow coer is likely to restrict them to limited
grazing grounds.

0HWKRGV
6WXG\DUHD
1he Shun Gorge ,33
o
N,
o
L, is located in the Zangskar Range, Ladakh, India, and
encompasses F0 sq. km. As other parts o Zangskar, it remains cut o rom the rest o
Ladakh in winter ,No.-Apr.,, when the only motorable road rom Kargil is blocked by
heay snow. 1hus during winter, it can only be accessed through a trek o F15 days oer
the rozen Zangskar, Lungnak and 1sarab riers. 1opographically, the area is
characterised by rugged terrain, with low rier blus along the 1sarab Rier. Lleation
ranges rom 3500-5000 m. Precipitation is mostly in the orm o snow during winter
,No.-leb.,, and during the study period, I recorded about a meter o snow on the upper
slopes.
No ungulates are ound in the area except the study species and domestic yak,
horse, sheep and goats. 1he sheep and goats were herded on the rier blus near the
illages, and were sometimes taken to the side-alleys, depending on the depth o snow.
1he yaks and horses entured out ar away rom the illages but not ar enough to
disturb the ibex and blue sheep, which generally occurred on higher reaches. 1here are
two illages YL]\arshun , households, and Marshun ,3 households, in the study area,
with a total human population o F50 people.
Both wild and domestic ungulates are preyed on by the snow leopardwol &DQLV
OXSXVFKDQFRand to a less extent by lynx /\Q[OLVDEHOOLQDAian predators like the golden
eagle $TXLODFKU\VDHWRV sometimes prey on the lambs o both domestic and wild ungulates.
See Namgail ,2004, or more inormation on wildlie and natural history o Zangskar.

)LHOGPHWKRGV
1he study was conducted between 12 Jan. -20 leb. 2002. 1he length o the study period
was constrained by the remoteness and inaccessibility o the study area. During the study,
I walked on the rozen 1sarab Rier, looking or ibex and blue sheep. 1wo permanent
trails: one upstream ,F5 km, and one downstream ,F3 km, rom Marshun Village were
established on and along the rier. Lach trail was walked at least 15 times during the
study period. 1he obserations were aided by 8x40 binoculars and a 15-45X spotting
scope.
\heneer a group o animals was encountered, I recorded the time, date, species
and habitat characteristics such as distance to cli, slope angle, eleation and snow coer.
All these habitat ariables were isually estimated except eleation, which was determined
rom a 1: 250, 000 topographic map, and the accuracy was oten checked by using a
Global Positioning System ,GPS,. lor estimating the proportions o aailable habitat,
149 random points were plotted on a 1: 250,000 topographic map o the area ,Marcum &
Lotsgaarden, 1980,, which were then located on the ground and the habitat
characteristics at these sites were sampled in the same way as described or habitat use.

61

.vat,ticat vetboa.
1he selection o habitat by ibex and blue sheep were determined by estimating selection
ratios ,ratio o the proportion o habitat use and aailable, or dierent habitat units. 1he
alue o the selection ratio o a habitat unit is proportional to the probability o that unit
being utilized by the study animals ,Manly et al., 1993,. Due to the small size o the study
populations, I needed to include re-sighted groups in the analysis, but the long
obseration interal ,one obseration on a group,day, should minimise the
autocorrelation. Since the aailable habitat was estimated, and indiidual animals were
not identiied, the data conormed to the design I ormat ,1homas & 1aylor, 1990, with
sampling protocol A ,Manly et al., 1993,. lor statistical analyses, the habitat ariables
were classiied into distinct categories, and the selection ratio ,
i
, or each category was
calculated as

i
~ o
i
,
i
Lqn. 1

where o
i
is the proportion o used units in category i, and
i
is the proportion o aailable
resource units in category i. Subsequently, the standard error o a selection ratio was
calculated as

.e ;
i
) ~
i
\ 1 ,v
i
- 1,v
-
- 1 ,v
i
- v
-
Lqn. 2

where v
i
is the used resource units in category i, v
-
is the total number o used units
sampled, v
i
is aailable resource units in category i and v
-
is the total aailable units in
category i.
1o statistically test or habitat selection, i.e., whether sample proportion o used
resource units were signiicantly dierent rom the sample proportion o aailable units,
the modiied

2
: log-likelihood Chi-square statistic

,

2
, or each habitat ariable was
calculated as

2
~ 2 _ v
i
tog
e
v
i
, ;v
i
) - v
i
tog
e
v
i
,;v
i
) Lqn. 3

where ;v
i
) is the expected alue o v
i
, and ;v
i
) is the expected alue o v
i
. I the

2
was
signiicant or a habitat ariable ,i.e., habitat selection,, simultaneous Bonerroni-adjusted
95 conidence interals were calculated or each category o that ariable ,to check
which categories are creating the signiicance, as

i
-
Z
,;2)
.e ;i) Lqn. 4

where is the number o habitat categories and .e ;i) is the standard error o selection
ratio. A habitat was used selectiely, i the conidence limit or that habitat excluded 1. I
selected, a habitat was preerred i the interal was 1, and aoided i 1 ,Manly et al.,
1993,.
Uniariate t-tests were used to statistically test or signiicant dierences in the
use o habitat in terms o distance to cli, slope angle, eleation and snow coer by ibex
and blue sheep. 1he slope angle and eleation were normally distributed, and distance to
cli and snow coer were log and arcsine transormed, respectiely, to ulill the
assumption o normality. Oerlap in habitat use by ibex and blue sheep was calculated
using Pianka`s index ,Pianka, 193,.

Chapter 4
62

=
2 2
.
.
Pik Pij
Pik Pij
Ojk Lqn. 5
where O
;/
is the measure o oerlap between species ; and /, and P
i;
and P
i/
are the
proportions o time spent by species ; and / respectiely on resources iv. Oerlap is
complete when O
;/
~ 1 and absent when O
;/
~ 0.
5HVXOWV
lorty-six obserations on ibex and 0 on blue sheep were made during the study period.
During sureys in the study area, I counted a maximum o 35 ibex and 6 blue sheep.
Assuming that I counted all the animals in the c. 0-km
2
-gorge, the aboe igures translate
to an ibex density o 0.5,km
2
and a blue sheep density o 0.96,km
2
. labitat selection
pattern by the two species is gien in 1able 1, while 1able 2 presents the means ,- SL,
o the habitat ariables used by the two species and their oerlap.
+DELWDWXVHDQGRYHUODS
Ibex preerred habitats close to clis ,1-50 m,

2
~ 8.92, p 0.05,, and their use o
other distance categories were in proportion to their respectie aailabilities ,1able 1,.
Blue sheep showed a similar pattern o habitat use, but used clis ,0 m, signiicantly less
than in proportion to their aailability ,

2
~ 14.2, p 0.05,. Ibex also aoided gentle
slopes ,15
o
,

2
~ 8.59, p 0.05,, while blue sheep aoided ery steep slopes ,45
o
,

2
~ 8.80, p 0.05,, but both species used other slope categories similarly and non-
selectiely ,1able 1,.
1able 1. Lstimated habitat selection indices or Asiatic ibex and blue sheep in Zangskar,
Ladakh, India.
i
estimated habitat selection ratio, .e;
i
) standard error o selection ratio,
i
;t) and
i
;v) 95 lower and upper conidence limits, respectiely.

-
,preerence,,
-
,aoidance,,
0
,use in proportion to aailability,,
]
inerences are less reliable due
to ery ew obserations ,v
i
5,.
Asiatic ibex Blue sheep
Variable
i
.e ;
i
)
i
;t)
i
;v)
i
.e;
i
)
i
;t)
i
;v)
Distance to cli ,m,
0 0.560
0
0.198 0.116 1.004 0.322
-
0.125 0.042 0.602
1-50 1.09
-
0.293 1.053 2.365 1.626
-
0.263 1.03 2.215
51-100 0.560
0
0.262 0.000
]
1.14 1.030
0
0.322 0.309 1.51
100 0.689
0
0.35 0.000
]
1.529 0.92
0
0.351 0.006 1.58
Slopeangle ,deg.,
15 0.395
-
0.23 0.000
]
0.926 0.952
0
0.338 0.195 1.09
16-30 0.945
0
0.19 0.504 1.386 1.341
0
0.202 0.889 1.93
31-45 1.809
0
0.423 0.861 2.5 0.905
0
0.251 0.343 1.46
45 0.448
0
0.22 0.000
]
1.05 0.196
-
0.145 0.000
]
0.521
Lleation ,m,
4000 0.04
0
0.209 0.236 1.12 0.294
-
0.113 0.041 0.54
4001-4150 1.313
0
0.326 0.583 2.043 1.6
0
0.351 0.990 2.562
4151-4300 1.96
0
0.606 0.619 3.333 2.0
0
0.55 0.89 3.365
4300 0.560
0
0.34 0.000
]
1.33 0.490
0
0.23 0.000
]
1.102
Snow coer ,,
0 0.395
-
0.160 0.03 0.53 0.909
0
0.209 0.441 1.3
1-25 2.239
0
1.04 0.000
]
4.584 0.36
0
0.436 0.000
]
1.13
26-5 1.990
0
0.586 0.6 3.303 1.553
0
0.451 0.543 2.563
5 0.833
0
0.194 0.398 1.268 0.890
0
0.13 0.502 1.28
63
Both species used eleation non-selectiely, except that blue sheep aoided habitats at
ery low eleation ,4000 m,
/
~ 23.49, p 0.05, 1able 1,. 1hey also used areas with

arying snow coer non-selectiely except that ibex aoided snow-ree areas ,
/
~ 12.21,
p 0.05,. 1he t-test statistics also reealed a high similarity in habitat use, diering
signiicantly only in the use o slope angle ,W ~ 2.589, S 0.01, 1able 2,, which was
urther conirmed by the relatiely less oerlap ,2
MN
~ 0.88, between the two species in
the use o this ariable ,1able 2,.

1able 2. Mean ,- SL, o habitat use and oerlap ,2
MN
, between ibex and blue sheep in
Zangskar, Ladakh, India.
Statistically signiicant
'LVFXVVLRQ
My results showed that ibex and blue sheep oerlap in their habitat use. Neertheless,
mechanisms other than snow coer restricting the two species to limited grazing grounds
are responsible or such high oerlap, because the two species did not preer the snow-
ree areas as expected under this assumption. During the study period, the two species
were seen eeding in close proximity ,20 m, on three occasions. Such sociality is a
common eature o the ungulate communities o eastern Arica, and has been iewed as a
response to high predation pressure ,Sinclair, 1985,. 1he high oerlap between ibex and
blue sheep in their habitat use could be related to their occurrence in close proximity or
mutual protection against predator.
1he high oerlap in habitat use by these species in the Zangskar Mountains may
also imply: ,a, that resources are abundant and they can coexist without strong
competition ,b, that they dier in the use o their diet ,c, that not enough time has
elapsed or eolutionary diergence o resource use. 1he irst explanation is untenable,
since the 1rans-limalayan Mountains support ery low plant biomass ,Chundawat &
Rawat, 1994, Mishra, 2001,. 1he second explanation is likely to account or the high
oerlap in habitat use, as diet separation allows the co-occurrence o species in the same
habitat ,Schoener, 194, Pianka, 1994,. 1he dietary use and oerlap between them need
to be inestigated to shed light on this aspect. 1he third explanation is also deensible, as
the resource partitioning between co-occurring species is an eolutionary process
,\alter, 1991,.
1he high preerence or habitat close to cli ,1-50 m, by both species is
consistent with the results obtained in other studies on these wild ungulates ,\ilson,
1989, Bhatnagar, 199, Longa, 1998, Namgail et al.,2004,. Such preerences relect the
importance o clis as escape terrain, and perhaps a high predation pressure in the area,
as indicated by a relatiely high abundance o snow leopard signs such as scrapes, spray
marks and droppings ,Namgail, Unpubl. data,. 1he high ainity o these ungulates
toward clis may also make them relatiely less ulnerable to disturbance associated with
liestock grazing, as such terrain types are used less requently by liestock herders ,pers.
labitat ariable %OXHVKHHS $VLDWLFLEH[ tYDOXH p
2
MN
Distance to cli ,m,

42 - 4.905 36 - 5.81 0.24 0.810 0.92
Slope angle ,degrees,
2 - 1.122 32 - 1.634 2.589 0.010 0.88
Lleation ,m,
4143 - 13.211 4092 - 32.56 1.51 0.082 0.95
Snow coer ,,
52.143 - 4.83 53 - 5.496 1.902 0.060 0.98
Chapter 4
64
obs.,. 1he relatiely higher number o these species in Ladakh ,lox et al.,1991, could be
due to a low leel o competition with domestic liestock, which has the capability o
out-competing ibex and blue sheep as shown by theoretical ,Mishra et al., 2002, as well
as empirical studies ,Bagchi et al., 2004, Mishra et al., 2004,.
1he oerlap in habitat use by ibex and blue sheep suggests a high potential or
competition between them, as habitat oerlap can lead to exploitation as well as
intererence competition ,Begon et al., 1996,. But comparatie data rom both sympatric
and allopatric populations, encompassing all seasons need to be collected to demonstrate
competition. loweer, gien the relatiely higher density o blue sheep in the Shun
Gorge, it is possible that this species negatiely inluences the population and distribution
o ibex in the Zangskar Mountains and other areas where they come together.
Covctv.iov
1here was substantial oerlap in habitat use by ibex and blue sheep, which is in contrast
to the prediction o competition theory. Such high oerlap in habitat use by the two
species in a region with inherently low plant productiity, especially in the resource-
limited winter months, suggests a high potential or competition. A thorough
understanding o resource selection o both sympatric and allopatric populations o ibex
and blue sheep is desirable to demonstrate competition.

$FNQRZOHGJHPHQWV
I am thankul to the Uniersity o 1romso, Norway or inancial support and the
necessary ield equipments. I thank Dr. Joseph L. lox or his help and guidance. I
grateully acknowledge the inrastructural support proided by the Nature Conseration
loundation during the preparation o the manuscript. I express sincere gratitude to
Ashley Spearing, Dorjey Gyalpo and Phuntsog Chosphel or their assistance and
encouragement in the ield. Discussions with Drs. \ash Veer Bhatnagar and Charudutt
Mishra helped in improing the manuscript, I thank them both. I also thank Dr. Marco
lesta-Bianchet or his critical comments on the manuscript. Kind cooperation and the
warm hospitality o the illagers o Marshun are also grateully acknowledged.

5HIHUHQFHV
Bagchi S., Mishra C. & Bhatnagar \.V. ,2004,- Conlicts Between 1raditional Pastoralism
And Conseration O limalayan Ibex ,Cara ibirica, In 1he 1rans-limalayan
Mountains. .viv. Cov.err. : 121-128.
Begon M., larper J.L. & 1ownsend C.R. ,1996,- cotog,: vairiavat., Povtatiov. .va
Covvvvitie., Pp. 945, Blackwell Scientiic Publications, Oxord.
Bhatnagar \.V. ,199,- Ravgivg .va abitat |titiatiov , 1be ivata,av be Cara be
ibrica v Piv 1atte, âtiovat Par/. Phd Dissertation, Saurashtra Uniersity, India
Chundawat R.S. & Rawat G.S. ,1994,- Indian Cold Deserts: A Status Report On
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Connell J.l. ,1961,- 1he Inluence O Interspeciic Competition And Other lactors On
1he Distribution O 1he Barnacle Cbtbavatv. tettatv.. cotog,, 42: 10-23.
De Boer \.l. & Prins l.l.1. ,1990,- Large lerbiores 1hat Strie Mightily But Lat
And Drink As lriends. Oecotogia, 82:264-24.
lox J.L., Nurbu C. & Chundawat R.S. ,1991,- Mountain Ungulates O Ladakh, India.
iot. Cov.err. 58: 16-190.
lox J.L., Sinha S.P. & Chundawat R.S. ,1992,- Actiity Patterns And labitat Use O
Ibex In 1he limalaya Mountains O India. ]. Mavvat. 3:52-534.
Gause G.l. ,1934,- 1be trvggte or i.tevce. Baltimore:\illiams & \ilkins.
65
lairston N.G., Smith l.L. & Slobodkin L.B. ,1960,- Community Structure, Population
Control And Competition. .v. ât. 94: 421-425.
lanley 1.A. & lanley K.A. ,1982,- lood Resource Partitioning By Sympatric Ungulates
On Great Basin Rangeland. ]. Ravge Mavage. 35: 152-158.
larris R.B. & Miller J.D. ,1995,- Oerlap In Summer labitats And Diets O 1ibetan
Plateau Ungulates. Mavvatia, 59: 19-212.
lolt R.D. ,19,- Predation, Apparent Competition, And 1he Structure O Prey
Communities. 1beor. Po. iot. 12: 19-229.
lolt R.D. ,1989,- Predation And Competition: 1he Interaction O 1wo 1ypes O
Species Interactions. Oi/o., 54: 256-263.
Jarman P.J. & Sinclair A.R.L. ,199,- leeding Strategy And 1he Pattern O Resource
Partitioning In Ungulates. In: Sinclair, A.R.L. & M. Norton-Griiths ,Lds,
erevgeti, D,vavic. Of .v co.,.tev, Pp. 130-166, Uniersity O Chicago Press,
Chicago
Jenkins, K.J. & \right, R.G. ,1988,- Resource Partitioning And Competition Among
Cerids In 1he Northern Rocky Mountains. ]. .t. cot. 11-24.
Lamprey l.l. ,1963,- Lcological Separation O 1he Large Mammal Species In 1he
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Inluenced By Predator 1ype. M.Sc. 1hesis, Uniersity O 1romso.
Mallon D.P. ,1991,- Status And Conseration O Large Mammals In Ladakh. iot.
Cov.err. 56: 101-119.
Manly B.l.J., Mcdonald L.L. & 1homas D.L. ,1993,- Re.ovrce etectiov , .vivat..
Chapman & lall.
Marcum C.L. & Lotsgaarden D.O. ,1980,- A Non-Mapping 1echnique lor Studying
labitat Preerences. ]. !itat. Mavage. 44:963-968.
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Mishra C., Van \ieren S.L., leitkonig I.M.A. & Prins l.l.1. ,2002,- A 1heoretical
Analysis O Competitie Lxclusion In A 1rans-limalayan Large-lerbiore
Assemblage. .viv. Cov.err. 5: 251-258.
Mishra C., Van \ieren S.L, Ketner P., leitkonig I.M.A. & Prins l.l.1. ,2004,-
Competition Between Domestic Liestock And \ild Bharal P.evaoi. â,avr In
1he Indian 1rans-limalaya. ]. .t. cot. 41: 344-354.
Namgail 1. ,2004,- Zangskar: Mystic Land. avctvar, ..ia, 24: 44-4.
Namgail 1., lox J.L. & Bhatnagar \.V. ,2004,- labitat Segregation Between Sympatric
1ibetan Argali Ori. .vvov oag.ovi And Blue Sheep P.evaoi. â,avr In 1he
Indian 1rans-limalaya. ]. Zoot. ;ova.). 262:5-63.
Oli M.K., 1aylor I.R. & Rogers M.L. ,1993,- Diet O 1he Snow Leopard ,Pavtbera |vcia,
In 1he Annapurna Conseration Area, Nepal. ]. Zoot. ;ova.). 231: 365-30.
Pianka L.R. ,193,- 1he Structure O Lizard Communities. .vv. Rer. cot. .va ,.t. 4:
53-4.
Pianka L.R. ,1994,- rotvtiovar, cotog,, Pp. 486, larpercollins College Publications, New
\ork.
Repasky R.R. ,1996,- Using Vigilance Behaiour 1o 1est \hether Predation Promotes
labitat Partitioning. cotog,, : 1880-188.
Rickles R.L. ,1990,- Lcology, 3rd Ldition. lreeman, New \ork.
Schaller G.B. ,1998,- !itatife Of 1be 1ibetav tee, Pp. 33, Uniersity O Chicago Press,
Chicago.
Schoener 1.\. ,194,- Resource Partitioning In Lcological Communities. cievce, 185:
2-39.
Chapter 4
66
Sinclair A.R.L. ,1985,- Does Interspeciic Competition Or Predation Shape 1he Arican
Ungulate Community -$QLP(FRO54: 899-918.
1homas D.L. & 1aylor L.J. ,1990,- Study Designs And 1ests lor Comparing Resource
Use And Aailability. -:LOGO0DQDJH54: 322-330.
Voeten M.M. & Prins l.l.1. ,1999,- Resource Partitioning Between Sympatric \ild
And Domestic lerbiores In 1he 1arangire Region O 1anzania. 2HFRORJLD, 120:
28-294.
\alter G.l. ,1991,- \hat Is Resource Partitioning -7KHRU%LRO150:13-143.
\egge P. ,1989,- \ild Caprids And 1heir Predators In Khunjerab National Park,
Northern Pakistan. In: :RUOG &RQIHUHQFH 2Q 0RXQWDLQ 8QJXODWHV Abstracts.
Camerino, Italy.
\iens J.A. ,19,- On Competition And Variable Lnironments. $P6FL65: 590-59.
\ilson P. ,1981,- Lcology And labitat Utilization O Blue Sheep 3VHXGRLV 1D\DXU In
Nepal. %LRO&RQVHUY21:55-4.

Species richness and niche dynamics
67
&+$37(5
(IIHFWVRIKHUELYRUHVSHFLHVULFKQHVVRQEOXHVKHHSVQLFKHG\QDPLFV
DQGGLVWULEXWLRQLQWKH7UDQV+LPDOD\D

1sewang Namgail
1,2
, Charudutt Mishra
2
, Christine B. de Jong
1
, Sipke L. an
\ieren
1
, lerbert l. 1. Prins
1

'..,in v .nv ncv {ccvin on v ni.ivc, 1 nv[cv ov vn vcv .in n, v..c. vnv .i.icv v
.o. /{c nv.n .cv.n 1 {ovnv v {c. vvc. o{ v.. vnv vn vinc vn o .o. ^cv.vin v
.v.c o{ 100 ,vv. , 10, 1 .,.cnvi.v, .oc.cv c.c, ..v o{ .ccvion, vnv in nc cnv nvv
.c.vcv .c.cnccn .inccv vvc. o{ .ov.c v.. vnv .c.cn .nv vinc vn., in {v., c.. nvn
onc .ovv {ccv v vincvi on, vnv ,c nc.c ncv. o{ .ncc vnv v..c. v:c .oncncv, vnv .in
.v..c.. on no.c vcn ni.ivc.. Inci ncvn. o{ .v.i.cn.c i. .i v v::c o nc"
R. Meinertzhagen
1
&
2
Karnataka, India
Chapter 5
68
$EVWUDFW
1he distributions o the 1rans-limalayan large herbiores are ragmented, engendering a
spatial heterogeneity in their species-richness. \e capitalised on this natural-experiment
situation to understand the niche dynamics o herbiores in relation to the number o
sympatric species. \e used the blue sheep 3VHXGRLV QD\DXU, a relatiely widely distributed
mountain ungulate, as a model species to address the issue. \e selected three discrete
alleys in three protected areas with almost similar enironmental eatures but arying
wild ungulate species richness, and studied the species` diet and habitat utilization in
them. labitat ariables were obsered in the ield and microhistological aecal analysis
was carried out to determine the habitat and diet width o the animal in the three areas
with dierent ungulate species richness. 1he habitat- and diet-niche widths were
determined using the Shannon`s l` Index. 1he results showed that habitat width o blue
sheep has a negatie relationship with the number o sympatric species. loweer,
contrary to our expectation, there was a hump-shaped relationship between blue sheep`s
diet width and the sympatric species richness, with the diet width being narrower in areas
o allopatry as well as in areas with greater number o sympatric species, and the widest
diet spectrum in areas with moderate species richness. \e suspect that the narrow diet
width in allopatry is out o choice, while it is out o necessity in areas with greater
number o sympatric species due to resource partitioning. \e suggest that interactions
with sympatric species lead to niche adjustment o mountain ungulates, and underscore
the importance o including biotic interactions in species distribution models, which hae
oten been neglected.

.H\ZRUGVLadakh, niche width, 3VHXGRLVQD\DXU, species diersity, 1rans-limalaya

69
,QWURGXFWLRQ
1heoretical and empirical studies in community ecology in the last our decades reoled
around niche related competitie interactions, and the principle o competitie exclusion
gained almost an axiomatic status ,Gause, 1934, lutchinson, 1959, Schoener, 1983,. 1he
basic tenet o competitie exclusion is that Qnumber o species cannot coexist on ewer
than Qresources ,Gause, 1934, lutchinson, 1959,. 1his implies that resource aailability
constrains the number o species occurring in an area ,MacArthur, 192,, and thus there
is a ceiling to species richness in ecological communities ,1erborgh & laaborg, 1980,
1onn et al., 1990,. Resource constraints allow the coexistence o only those species that
show trade-os in niche utilization in response to competition ,Chase & Leibold, 2003,.
1hereore, species may aoid or reduce competition by adjusting their respectie niche
widths in response to their co-inhabitants ,Chase & Leibold, 2003,.
Studies on niche related resource partitioning hae been conducted on seeral
taxa ,1horman, 1982, 1ot, 1985, \heeler & Caler, 1996, 1oda et al., 1999, McDonald,
2002,, but niche relationships are less understood in large herbiores such as ungulates,
largely because o their low population densities and the diiculties associated with
manipulating their populations. Most o the studies on wild ungulates hae addressed
resource partitioning in single assemblages ,Jenkins & \right, 1988, Putman, 1996,
Voeten & Prins, 1999,, but it is not known as to how the niche o a gien species can
ary across assemblages in response to the number o sympatric species. 1his
inormation is crucial or predicting species distributions at a macroecological scale
,Arajo & Luoto, 200,.
Owing to the presence o sympatric species sharing resources, animals in multi-
species assemblages may use only a subset ,i.e., realized niche, o all the resources
aailable ,i.e., undamental niche, in an area ,lutchinson, 195,. Gien this, species can
be packed into assemblages either as a result o increasing the resource range, or
narrowing the niche width o the constituent species ,MacArthur, 192,. In low
productie enironments, since resources are scarce, niche adjustment is expected to be
the predominant way o accommodating additional species. 1hus when a species goes
extinct rom a community, niche space ,habitat supplying resources or a species`
surial, is expected to become acant that can either be occupied by an inading species
or exploited by the extant species leading to readjustment in their niche widths.
1hereore, it is postulated that disappearance o large herbiore species rom
ecological communities leads to increase in the niche width o the extant species. But in
case o diet, loss o a potentially competing sympatric species o large herbiore may
hae an opposite eect on niche width o a gien species, as it may narrow down its
niche by selecting ewer but more nutritious plant species in its diet in the absence o
competitors. Inersely, the species may widen its diet spectrum when a new species
inades the assemblage, as it may need to include less nutritious plants in the diet due to
resource constraint brought by the inading species. 1hus, ood and habitat widths could
hae dierent relationships with the number o sympatric species in an assemblage.
1he mountainous rangelands o the Indian 1rans-limalaya support a relatiely
dierse assemblage o eight wild ungulates ,lox et al., 1991, including our Caprinae
species that represent 40 o this taxon ound in the limalayan region. But their
populations are ragmented, and there is a spatial ariation in the species richness, with
local assemblages representing smaller subsets o the regional species pool. 1he causes o
this ariation and the actors that inluence the organization o herbiore assemblages in
this region are poorly understood, though excessie liestock grazing has been implicated
as one o the causes ,Mishra et al., 2004,.
Using the blue sheep 3VHXGRLV QD\DXU, a relatiely widely distributed animal, as a
model species, we explored how the niche o a wild ungulate aries in response to
Chapter 5
70
sympatric species richness. lor this, we studied its habitat and diet in areas where it
occurs allopatrically and contrasted it with areas where it occurs sympatrically with one:
Ladakh urial 2YLVYLJQHL YLJQHL and two Caprinae species: Asiatic ibex &DSUDLEH[VLEHULFDand
Ladakh urial1hese sympatric speciesare comparable to blue sheep in morphology ,Van
den 1empel & De Vrij, 2006, as well as behaiour ,Schaller, 19,, and thus hae similar
ecological requirements ,Mallon, 1991, Namgail, 2006b,. \e urther explored the
relationship by including additional inormation rom literature on blue sheep`s diet rom
other 1rans-limalayan ungulate assemblages with more than three sympatric species.
\e predicted ,1, an inerse relationship between blue sheep`s habitat width and
the number o co-occurring species ,2, a positie relationship between its diet width and
the number o co-occurring species.
0HWKRGV
6WXG\DUHDDQGVSHFLHV
1he western Indian 1rans-limalaya ,31
o
36` to 34
o
40`N and 5
o
40` to 9
o
30`L, is
classiied as a cold desert. 1he moisture-laden monsoon clouds hardly reach this region
due to the rain-shadow eect o the limalayan range. 1he magnitude o precipitation,
mostly in the orm o snow during winter, is thereore minimal with the mean annual
precipitation rarely crossing 100 mm. 1he temperature ranges rom -30
o
C in peak winter
,Dec-Jan, to -35
o
C in summer ,June-Aug,. Vegetation is characterized by dry alpine
steppe ,Champion & Seth, 1968,, and the plant coer rarely crosses 30 except in
meadows around water bodies such as lakes and riers ,Rawat & Adhikari, 2005,. 1here
are only ew tree species including poplar 3RSXOXV spp. and willow 6DOL[ spp., which are
conined to the rier-alleys. 1he most common egetation includes &DUDJDQD spp.,
$UWHPLVLD spp., /RQLFHUD sp. and $FDQWKROLPRQ sp. Some o the common herbs include
3RWHQWLOODspp., 2[\WURSLVspp., $VWUDJDOXVspp. and 'UDFRFHSKDOXPsp.
1he three study sites where blue sheep occurs allopatrically and with arying
number o sympatric species were as ollows: a, Rongolong ,32
o
20`N, 8
o
02`L, in the
Kibber \ildlie Sanctuary ,hereater Kibber, is located south o Ladakh, and is
administratiely a part o limachal Pradesh, b, Puyul alley ,33
o
43`N,
o
4`L, o the
proposed Gya-Miru \ildlie Sanctuary ,hereater Gya-Miru, and c, Rumchung alley
,34
o
08`N,
o
24`L, o the lemis National Park ,hereater lemis, are administratiely
part o Ladakh, Jammu and Kashmir. Reconnaissance sureys were carried out in these
protected areas prior to the study to ind out alleys with the desired number o ungulate
species but with almost similar enironmental eatures. 1hus, our study site in Kibber
had only one species: blue sheep, Gya-Miru had two species: blue sheep and Ladakh
urial, whereas lemis supported three species: blue sheep, Ladakh urial and Asiatic ibex.
1here are also small populations o limalayan marmot 0DUPRWD EREDN in these areas,
except in Kibber, where it has not been obsered or seeral years ,Mishra, 2001,.
1he blue sheep is a sturdy animal with strong muscular legs that help it in
climbing steep clis, which are used as reuge against predators ,Namgail et al., 2004,. It
grazes on open alpine pastures within an altitudinal range o 3500-5500 m, but keeps
closer to precipitous clis to aoid predation. It, howeer, exhibits an altitudinal
migration by coming down to lower eleations during winter, when upper reaches get
coered with heay snow ,Namgail, 2006b,. 1he animal has been reported to eed largely
on graminoids such as &DUH[.REUHVLD and 6WLSD during summer ,larris & Miller, 1995,
Mishra et al., 2004,. It is distributed all across the 1ibetan plateau and its marginal
mountains, although the population is ragmented and the density aries across its
distributional range ,Schaller, 1998,. 1here is an estimated population o F 11,000
indiiduals in Ladakh, which makes it the most abundant wild ungulate in the region
,lox et al., 1991, Namgail, 2009,.
71
)LHOG0HWKRGV
+DELWDW
Data were collected between May 2005 and Aug. 200. Blue sheep herds were located
rom trails and antage points ,Namgail et al., 2004,. \e searched the mountain slopes
with 8x40 binoculars. Scan sampling was the primary method or animal obserations.
\heneer a group o animals was located, its size and subsequently the habitat ariables:
slope angle, distance to cli and eleation at its location were recorded. Indiiduals were
considered to be solitary or belong to dierent groups when they stood 50 m away rom
another group. Although we also gathered inormation on physical ariables like slope
aspect and slope position, only the aorementioned ariables were used to estimate the
habitat-niche width o the animal, a model selection procedure ,Akaike Inormation
Criterion, identiied these as the most important ariables in blue sheep habitat use
,Namgail, 2006a,. An altitudinal gradient proides dierent habitats or plants, and thus
the egetation diersity and abundance ary along such a gradient in Ladakh ,Chapter 2,,
whereas distance to cli proides a gradient in egetation biomass ,more egetation away
rom clis, and predation risk or blue sheep as the animal uses clis as escape terrain
,Namgail et al., 2004,. Since dierences in aailability o habitat in the three areas could
conound the inluence o ungulate species richness on blue sheep`s niche dynamics, we
recorded the aailable habitats in dierent areas to control or it.

'LHW
Blue sheep`s summer diet inormation or Kibber was aailable rom Mishra et al.,
,2004,. lor the other two areas, we conducted microhistological aecal analysis. \e also
used additional inormation rom larris and Miller ,1995, on blue sheep`s
microhistological aecal analysis-based diet in \eniugou China with six sympatric species
to urther explore the relationship between diet width and species richness. Although
Mishra et al. ,2004, quantiied eeding signs on egetation ater obsering animals
eeding in the ield, gien that the animals could be obsered at close quarters, we beliee
the diet inormation rom the two studies are comparable. Initially, we also did direct
obserations on eeding animals, but gien the workload in the ield: estimating habitat
use, aailability and plant sampling during two seasons in all the dierent study sites, we
ound it less practical in this large scale study. 1hus, one o us handled the laboratory
work, while another estimated habitat use and collected aecal pellets. 1o preent
assigning pellets mistakenly to a dierent species than the one intended to, we collected
them rom bedding sites by waiting or the animals to get up and moe away. A group o
F50 pellets was collected rom each herd o blue sheep. Subsequently, ie pellets were
randomly drawn rom each group to orm one sample or the respectie herd. 1hus there
were 11 samples rom Gya-Miru and 9 rom lemis.
1hese samples were air-dried and stored in paper bags beore boiling in water or
about 1 h and soaking oernight. 1hey were then crushed in the laboratory, and the inner
tissue was separated rom the epidermis and cuticle by mixing a 5 g sub-sample with
water or 1 minute in a \aring blender, and was strained oer a plankton siee ollowing
de Jong et al. ,2004,. 1he residue was then washed again with tap water, transerred into
a petri-dish and allowed to settle. Using a Pasteur pipette, ten random grab samples o
the residue were then taken, and each droplet was put on a glass slide, spread out eenly
and coered with a 2.4 cm coer-slip.
\e prepared separate reerence slides or plants collected rom the ield. lor this
small pieces o plant parts were cleaned in household bleach oernight, washed in water,
and ragments o epidermis were then stripped o and mounted in glycerol ,de Jong et
al., 2004,. Photomicrographs o epidermal material on a set o these reerence slides were
used to identiy the ragments o cuticles obsered in samples o the animal aeces. At
Chapter 5
72
least 100 cuticle or epidermal ragments were identiied in each sample. 1o quantiy the
composition o the aecal material, the area o epidermal ragments was measured at a
magniication o 100-X using a grid o small squares ,each representing 0.01 mm
2
, in the
microscope eyepiece. 1he abundance o each species was calculated as a percentage o
the total area o the ragments measured ,Putman, 1984, Alipayo et al., 1992, lomolka &
leroldoa, 1992,.
Since, the dierence in plant species richness between the areas is likely to aect
the relationship between blue sheep niche width and ungulate species richness, we
accounted or this parameter by estimating it in the three areas. 1he inormation on plant
species richness or Kibber was taken rom Mishra ,2001,, while or Gya-Miru and
lemis, a transect was laid on an altitudinal gradient at eery 200 m alternately on either
side o the alley, starting at the alley-mouth. 1hese transects were laid in the main
alleys as well as in the side alleys. Lach transect was then diided into 50 m segments,
and a 2 x 2 m plot was sampled at eery 50 m intercept. 1he adequacy o the plot size
was ascertained by examining the species accumulation cures, which reached an
asymptote at 2 x 2 m. \e also estimated aboeground biomass rom these transects.
Plants in these plots were identiied in the ield using a plant ield guide ,Polunin &
Stainton, 1990,. 1he unidentiied ones were collected and later identiied at the \ildlie
Institute o India.

6WDWLVWLFDODQDO\VHV
Blue sheep`s niche width in terms o habitat and diet were determined using the
Shannon-\iener Index ,Magurran, 1988,. 1his index aries rom 0 or minimum
resource items to about 5 or niche spectrum with maximum resource items, taking into
account the number or abundance o each item. \e assigned dierent resource units
,e.g., 50 m in case o distance to cli, into discrete categories to determine the niche
,habitat, width, while or the diet width each plant species ormed a discrete category.
1he class interals or the physical ariables were as ollows: altitude ,interal 100 m,
range 4000 to 5300 m,, slope angle ,5 degrees, 0 to 65 degrees, and distance to cli ,50
m, 0 to 600 m,. Bootstrap resamplings were used to construct 95 conidence interals
to estimate the ariability in the measure associated with sampling errors. 1he dierences
in the niche width ,both diet and habitat, o blue sheep between the areas with diering
species richness were tested or signiicance with a special t-test with the Shannon-
\iener indices as
[ ]
2 / 1
2 1
2 1
) ' var( ) ' var(
' '
H H
H H
t
+
= , where l
1
` is the niche ,habitat or diet,
width o the species in one area and l
2
` is its niche width in another area ,Poole, 194,.
\e pooled the habitat data o 2005 and 2006 rom Kibber as there was no inter-
annual ariation in habitat use ,lotelling`s 1
2
~ 5.60, l ~ 1.81 p0.15 or summer & 1
2

~ 8.65, l ~ 2.58 p0.08 or winter,. In addition to the changes in niche width, there
might also be niche-shits, utilizing dierent resource units in dierent areas as well as
seasons. In order to assess this possible lexibility, we checked or signiicant dierences
in habitat use between areas as well as seasons with Analysis o Coariance ,ANCOVA,,
using aailability as coariate ,Zar, 1984,.

5HVXOWV
A total o 1 obserations on blue sheep habitat use during summer and 42 during winter
were made in Kibber. 1he mean group size o the animal in this area during summer was
18, while that during winter was 15 ,1able 1,. In Gya-Miru, 46 obserations were made
during summer and 86 were made during winter. A total o 4 obserations during
summer and 28 during winter were made in lemis. Median group size between seasons
73
,winter ~ 15, summer ~ 14, or between areas were not signiicantly dierent, and the
aerage group size o blue sheep was thus 13..

1able 1. Mean and range o group size o blue sheep 3VHXGRLVQD\DXUin the study sites in
the Indian 1rans-limalaya.

Place Season n Group size ,mean, Group size ,range,
lemis Summer 4 11. 1-34
\inter 28 8.0 1-23
Gya-Miru Summer 46 13.6 1-53
\inter 86 16.2 1-48
Kibber Summer 1 1.6 1-68
\inter 42 15.0 3-48

+DELWDW
During summer, blue sheep had the widest habitat width in Kibber ,l`~ 3.06,, where it
occurs allopatrically, and had the narrowest ,l` ~ 2.6, in lemis where it shared
resources with two sympatric species. 1his trend remained similar in winter ,lig. 1,. 1he
animal`s niche width, in terms o habitat, thus declined with increase in the number o
sympatric species in the assemblage ,lig. 1,. 1his decline was signiicant during summer
as well as during winter, adjudged by the dierences in the niche width o the animal
rom the three areas using t-tests ,1able 2,.

Summer
2.5
2.7
2.9
3.1
3.3
Kibber (1) Gya-Miru (2) Hemis (3)
Number of sympatric species
N
i
c
h
e

w
i
d
t
h

Winter
2.3
2.5
2.7
2.9
Kibber (1) Gya-Miru (2) Hemis (3)
N
i
c
h
e

w
i
d
t
h

lig. 1. 1he relationship between blue sheep`s habitat-niche width ,Shannon-\iener
indices with 95 conidence interals, and the number o sympatric species ,in
parentheses, in the Indian 1rans-limalaya.

1able 2. Dierences in habitat width o blue sheep 3VHXGRLVQD\DXUin Kibber ,allopatric,,
Gya-Miru ,with one sympatric species, and lemis ,with two sympatric species, in the
Indian 1rans-limalaya

Area pair Summer \inter
t-alue 3 t-alue 3
Kibber and Gya-Miru
Kibber and lemis
Gya-Miru and lemis
1.546
3.660
1.621
0.123
0.001
0.106
1.0
2.495
3.664
0.282
0.013
0.001

1he distance o blue sheep locations rom the nearest clis diered signiicantly between
areas with dierent species richness ,ANCOVA, l ~ 20.01, p 0.001, as well as seasons
Chapter 5
74
,l ~ 23.26, p 0.001, 1able 3,. lor instance, the mean distance to cli or blue sheep
sightings during summer in Kibber was 144 m whereas that in Gya-Miru and lemis were
114 m and 46 m, respectiely ,1able 4,. 1he species also diered in terms o the slope
angle o locations between the areas ,l ~ 8.4, p 0.001,, but not between seasons ,l ~
0.38, p ~ 0.539,. Neertheless, there was a signiicant interaction between species
richness and season, with the animal using steeper areas ,mean ~ 34
o
, during summer
and latter areas ,mean ~ 31
o
, during winter in Kibber and Gya-Miru and ice-ersa in
lemis ,l ~ 4.56, p 0.01, 1able 3 & 4,.

1able 3. Summary o ANCOVAs carried out on habitat use by blue sheep during two
seasons ,summer and winter, in three 1rans-limalayan sites with dierent species
richness with aailable habitat as coariate.

Variable

Lect l

d

P

Distance to cli ,m,

Species
Season
Species x Season
Aailable
20.01
23.26
1.5
0.02
2
1
2
1
0.001
0.001
0.14
0.885

Slope angle ,deg,
Species
Season
Species x Season
Aailable
8.4
0.38
4.56
2.1
2
1
2
1
0.001
0.539
0.011
0.141

Lleation ,m,
Species
Season
Species x Season
Aailable
36.15
18.55
6.33
3.93
2
1
2
1
0.001
0.001
0.001
0.048

Blue sheep also diered signiicantly in its use o the altitudinal gradient between the
areas ,l ~ 36.15, p 0.001, as well as between seasons ,l ~ 18.55, p 0.001,. 1here was
also an interaction eect ,l ~ 6.33, p 0.001,, as the animal used higher areas ,mean ~
4523 m, during summer and lower areas ,mean ~ 4385 m, during winter in Kibber where
it occurs allopatrically and in Gya-Miru with one sympatric species, while this seasonal
trend was opposite in lemis with two sympatric species ,1able 4,. But or this ariable,
there is an eect o the aailable habitat on these dierences ,l ~ 3.93, p ~ 0.048, 1able
3,.

'LHW
Blue sheep`s diet width had a hump-shaped relationship with the number o sympatric
species. 1he animal had a narrower diet width in areas o allopatry and areas o high
species richness, but wider niche width in areas with intermediate ungulate species
richness ,lig. 2, 1able 5,. lor instance, the diet width o blue sheep was narrower
,l`~1.86, in Kibber where the animal occurred allopatrically, and the widest in Gya-Miru
,l` ~ 2.81, with one sympatric species, and again narrower ,l`~2.36, in lemis with two
sympatric species.

75
1able 4. Mean ,- SL, o the seasonal habitat use in relation to aailability by blue sheep
in Kibber ,allopatric,, Gya-Miru ,one sympatric species, and lemis ,two sympatric
species, o the Indian 1rans-limalaya.
Distance ~ Distance to cli ,m,, Slope ~ Slope angle ,deg., and Alt ~ Altitude ,m,, Aail. ~Aailable

Its diet width diered between Kibber and Gya-Miru ,t ~ 2.948, p ~ 0.004, but not
between Kibber and lemis ,lig. 2,. 1he diet width o the animal urther narrowed down
,l` ~ 1.0, in \eniugou with six sympatric species, which was signiicantly dierent
rom blue sheep`s diet width in Gya-Miru ,t ~ 5.245, p 0.001, and lemis ,t ~ 4.499, p
0.001, 1able 5,.

1
1.5
2
2.5
3
3.5
Kibber (1) Gya-Miru
(2)
Hemis (3) Yeniugou
(6)
N
i
c
h
e

w
i
d
t
h

lig. 2. Relationship between blue sheep`s diet-niche width ,Shannon-\iener indices with
95 conidence interals, and the number o sympatric species ,in parentheses, in the
1rans-limalaya ,data source or \eniugou: larris and Miller, 1995,.

Variable Kibber Gya-Miru lemis
$YDLO 6XPPHU :LQWHU $YDLO 6XPPHU :LQWHU $YDLO 6XPPHU :LQWHU

Distance

112.4
-13.1

144.01
-15.39

8.42
- 13.3

238.5
-184.8

114.5
-16.09

60.29
-.03

243.8
-199.0

45.58
-5.09

25.35
-3.83
Slope 1.
- 9.4
2.59
-1.4
26.8
-2.08
2.0
- 8.12

34.13
-1.11

31.68
-1.11
26.02
- 8.85

30.8
-1.44
39.14
-2.5
Alt 4631.0
- 02.8
4523.10
-30.89
4384.2
-40.4
462.4
- 351.8
4530.8
-50.30
4158.6
-50.21
49.3
-21.
4082.09
-3.38
4104.5
-39.35
Chapter 5
76
1able 5. Dierences in diet width o blue sheep 3VHXGRLVQD\DXUduring summer in Kibber
,allopatric,, Gya-Miru ,with one sympatric species,, lemis ,with two, and \eniugou
,with six, in the 1rans-limalaya.

Area pair t-alue P
Gya-Miru and Kibber 2.95 0.01
Gya-Miru and lemis 2.58 0.01
Kibber and lemis 1.48 0.14
\eniugou and lemis 5.13 0.001
\eniugou and Kibber 5.22 0.001
\eniugou & Gya-Miru 8.38 0.001
3ODQWDYDLODELOLW\
1here were 21 plant species in Gya-Miru, 1 in lemis, 16 in Kibber, 20 in \eniugou that
are known to be important constituent o the diet o mountain ungulates in the 1rans-
limalaya. &DUH[, .REUHVLD, 6WLSD, )HVWXFD$VWUDJDOOXV2[\WURSLV/H\PXV3RWHQWLOOD1HSHWD and
$UWHPLVLD were some o the most abundant genra in the our sites. 1he plant species
richness did not dier between dierent areas ,p0.05 or all paired t-tests,. Data on
aboeground biomass aailable rom two sites show that the mean ,- SD, plant biomass
in Gya-Miru was 6.31 ,- 3., g,m
2
, while that in lemis was 4.15 ,- 2.6, g,m
2
, but the
dierence is statistically not signiicant ,t ~ 1.686, p ~ 0.10,.
'LVFXVVLRQ
1his study has shown that blue sheep`s niche width in terms o habitat declines as the
number o sympatric species increases in an area, which is in line with our irst
prediction. Such a relationship is in concordance with niche trends obsered in small
mammal ,lox, 1981, and ish communities ,1horman, 1982,. loweer, the animal`s diet
width, surprisingly, showed a hump-shaped relationship with the widest diet width at
intermediate species richness. lor instance, blue sheep had a narrower diet width in
Kibber where it occurred allopatrically, and in lemis with two sympatric species, but the
widest diet width in Gya-Miru with one sympatric species. 1his trend was urther
strengthened when we included the diet data o blue sheep in \eniugou with six
sympatric species ,larris & Miller, 1995,, where it had a diet width narrower than in any
o our study sites. 1his pattern is in contradiction to our expectation under competition
theory that as the number o sympatric species increases, the animal should widen its diet
width, incorporating less nutritious plants in its diet due to orage constraint imposed by
the sympatric species.
\e suggest that the narrow diet width o blue sheep in areas with greater number
o sympatric species is out o necessity, as the orage intake o herbiores in low
productie enironments such as the 1rans-limalaya is constrained by aailability o
plants that are sparsely distributed and are also ed on by sympatric species. 1hus the
animal narrows down its diet width, eeding on ewer but readily aailable plant species.
On the other hand, the narrower diet width in allopatry is presumably out o choice, as
the animal can choose the most nutritious plants rom an array o plants aailable. In any
case, it became apparent that niche o herbiores in terms o habitat and diet hae
dierent dynamics in areas with dierent number o sympatric species in high altitude
grazing ecosystems in the 1rans-limalaya. Although there are marmots in these areas,
gien their localised distribution in moist areas such as stream-banks with higher
egetation coer ,Alred et al., 2006,, they would not hae inluenced the oerall results
o this study.
77
A caeat o this study is the lack o replication. But this was judged to be a minor
disadantage, or the ield work in more areas in this obserational study would hae
ensued more enironmental ,natural as well as man-made, heterogeneity amongst the
study sites, thereby conounding the eect o sympatric species richness on blue sheep`s
niche width. \e howeer accounted or the dierences in habitat aailability that might
also hae an eect on blue sheep`s niche utilization, because changes in the aailability o
habitat ariables change the competitie balance o the co-occurring species.
lurthermore, the plant species richness was comparable in the three study areas.
Although Mishra ,2001, estimated plant species richness using a stratiied random
sampling method, gien that oer 2 years were spent in the area studying wildlie and
rangeland, it is less likely that species richness was underestimated. 1o our knowledge,
this is the irst study looking at the relationship between niche width and herbiore
species richness in wild ungulates at a regional scale, and has important implications or
their management in grazing ecosystems.
Blue sheep is the most widely distributed mountain ungulate in the Ladakh region
o the Indian 1rans-limalaya ,Namgail, 2006b,. 1his wide distribution o the animal
could be related to its ersatility in resource use according to aailability determined by
biotic actors as shown by this study as well as abiotic actors such as terrain. lor
instance, although the animal is known to use steep clis as anti-predator habitat
,Namgail et al., 2004,, it reportedly used boulders as escape terrain on the 1ibetan plateau
where aailability o clis was low ,larris & Miller, 1995,. Blue sheep`s wide distribution
thus may be the result o its dietary and habitat lexibility, suggesting that compared to
other Caprinae species, it may ace ewer constraints to recolonise an area ater local
extinction.
1o conclude, it became apparent that blue sheep`s niche aries across areas with
dierent number o sympatric species. 1hus, biotic interactions seem to play a role in
distribution o mountain ungulates, which should be addressed urther in other
mountainous grazing ecosystems. Species distribution models oten incorpoprate only
the enironmental ariables and tend to neglect the biotic interactions ,Arajo & Luoto,
200,, which perhaps leads to oerestimation o distributional range o species because
animals cannot occupy all the suitable habitats due to competition with other sympatric
species ,lutchinson, 1959,. 1hereore, our results underscores the role o interspeciic
interaction in species distributions and the importance o including this ariable in
species distribution models, perhaps by incorporating the diersity and abundance o
sympatric species in the model.
lurthermore, the conentional contention that a large herbiore widens its niche
width in areas with high species richness to aoid competition needs to be reexamined
especially in low productie enironments like the 1rans-limalaya, where aailability o
resources is minimal due to low plant diersity and biomass, which urther declines in
winter due to senescence and snow coer. lrom a conseration point o iew, it is
obious that gien the tendency o the wild ungulates to shit their niche in response to
sympatric species, the mountain ungulates in the Ladakh 1rans-limalaya are probably at
a disadentage in the ace o the recent increase in population o a ariety o liestock in
in the region ,Namgail et al., 200a,. Although studies hae been carried out at local
scales to address this issue ,Bagchi et al., 2004, Mishra et al., 2004, Namgail et al., 200b,,
there is no inormation at larger geographical scales, which is urgently needed due to the
relationship between mechanisms underlying local and regional species richness ,Rickles
& Schluter, 1993,.

Chapter 5
78
Acknowledgements
1his study was carried out with inancial assistance rom the Ruord Small Grants
loundation, \ageningen Uniersity and the \ildlie Conseration Society. \e thank the
Department o \ildlie Protection, Jammu and Kashmir and the Department o lorest
larming and Conseration, limachal Pradesh, or granting permission to carry out
ieldwork in the protected areas. \e thank Dr. G.S. Rawat at the \ildlie Institute o
India or identiying the plants. \e thank Karma Sonam rom Ladakh and 1andup
Dorjey and Lobzang Gialson rom Spiti or their inaluable help during the ield work.

References
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contents analyses in studies o the ungulate diet. )ROLD=RRORJLFD 4J, 193-208.
lutchinson, G. L. ,195, Concluding remarks. &ROG 6SULQJ +DUERU 6\PSRVLXP RQ
4XDQWLWDWLYH%LRORJ\ 22, 415-42.
lutchinson, G. L. ,1959, lomage to Santa Rosalia or why are there so many kinds o
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Magurran, A. L. ,1988, (FRORJLFDO'LYHUVLW\DQG,WV0HDVXUHPHQWPrinceton Uniersity Press,
Princeton.
Mallon, D. P. ,1991, Status and conseration o Large Mammals in Ladakh. %LRORJLFDO
&RQVHUYDWLRQ S6, 101-119.
McDonald, R. A. ,2002, Resource partitioning among British and Irish mustelids. -RXUQDO
RI$QLPDO(FRORJ\ 7J 185-200.
79
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Rawat, G. S. & Adhikari, B. S. ,2005, lloristics and distribution o plant communities
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âtvrati.t, J22, 240-285.
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.vericav âtvrati.t, JJ6, 18-195.
1horman, S. ,1982, Niche Dynamics and Resource Partitioning in a lish Guild
Inhabiting a Shallow Lstuary on the Swedish \est Coast. Oi/o., 39, 32-39.
1oda, M. J., Kimura, M. 1. & 1uno, N. ,1999, Coexistence mechanisms o mycophagus
drosophilids on multispecies ungal hosts: aggregation and resource partitioning.
]ovrvat of .vivat cotog,, 68, 94-803.
1ot, C. A. ,1985, Resource Partitioning in Amphibians and Reptiles. Coeia, J, 1-21.
1onn, \. M., Magnuson, J. J., Rask, M. & 1oionen, J. ,1990, Intercontinental
comparison o small-lake ish assemblages: 1he balance between regional and
local processes. .vericav âtvrati.t, J36, 345-35.
\heeler, A. G. & Caler, M. C. ,1996, Resource partitioning in an island community o
insectiorous birds during winter. M|, 96, 23-31.
Chapter 5
80
Zar, J. l. ,1984, %LRVWDWLVWLFDODQDO\VLVPrentice-lall Inc., New Jersey, N.J., USA.

Co-distribution of urial and blue sheep
81
CHAPTER 6
&RH[LVWHQFHRI/DGDNKXULDODQGEOXHVKHHSDWPXOWLVSDWLDOVFDOHVLQ
WKH7UDQV+LPDOD\DQPRXQWDLQV

1sewang Namgail
1, 2
, Sipke L. an \ieren
2
, Charudutt Mishra
1
and lerbert
l.1. Prins
2

&RPSHWLWLRQ IRU UHVRXUFHV ZRXOG EH LQHYLWDEOH DQG FRH[LVWHQFH LPSRVVLEOH ZLWKRXW D JHRJUDSKLF
SDUWLWLRQLQJ RI UDQJHV 6\PSDWULF VXUYLYDO LV QRW SRVVLEOH LQ D VLPSOH KDELWDW IRU FDSULGV RI D
VLPLODUVL]H
George B. Schaller
1
Snow Leopard 1rust ,India Program,, Nature Conseration loundation, 306,5 IV-
Cross, Gokulam Park, Mysore-50002, Karnataka, India

2
Chapter 6
82
$EVWUDFW
Large wild herbiores are important ecologically and economically, and maintaining their
populations is an important management priority. loweer, herbiores might not be able
to occupy all suitable habitats due to the presence o other potentially competing wild or
domestic species. 1he Ladakh urial 2YLV YLJQHL YLJQHL is an endemic and endangered wild
sheep inhabiting the mountainous region o Ladakh, northern India, where its population
is restricted to narrow tracts along two rier-alleys. 1he causes o this restricted
distribution o the species are not understood. Lxcessie grazing by domestic liestock in
this landscape is considered as an important actor inluencing wild herbiore
populations, but whether a wild ungulate can inluence another wild ungulates` range is
less known. \e asked i competitie exclusion by the more abundant wild ungulate, the
blue sheep 3VHXGRLV QD\DXU, could explain the singular distributional pattern o Ladakh
urial. 1o explore this possibility, we studied the occurrence patterns o these two species
at multiple scales ,geographical, landscape and habitat,. \e ound that they occurred
independently at the geographical scale, but co-occurred at the landscape scale, acilitated
by diergence in seasonal resource use obsered at the habitat scale. Although the two
species segregated along diet as well as habitat dimensions in summer, enabling their
coexistence at the landscape leel, there remained a high potential or competition during
winter when they oerlapped in their habitat use. \e conclude that while human and
liestock impacts may currently aect its distribution, the presence o blue sheep appears
to be an important historical actor limiting the population growth and range expansion
o the Ladakh urial.

.H\ZRUGV 2YLV YLJQHL 3VHXGRLV QD\DXU niche relationship, habitat selection, resource
partitioning, 1rans-limalaya.

83
,QWURGXFWLRQ
Understanding spatial distributions o animals and the underlying mechanisms hae both
theoretical and practical alue ,Andrewartha & Birch, 1954,. Mechanistic understanding
o the geographical distribution o a species needs a multi-scale approach. lactors like
immigration and emigration play important roles in species distributions at a regional
scale ,Rickles & Schluter, 1993,, whereas species distributions at smaller scales are
goerned by the aailability o crucial resources or their growth, reproduction and
surial ,MacArthur, 192, Begon et al., 1996,. Neertheless, species oten cannot locally
occupy all the aailable habitats as other biological actors preent them rom using their
ull potential range. lor instance, utilization o all potential areas may be impeded by
interactions with other ecologically similar species ,lutchinson, 1959,.
It is well established that energetic constraints set a limit to the number o species
that an area can support ,MacArthur, 192, Ol & Ritchie, 1998,. Competitie
interactions may set an upper limit on local diersity, and the role o competition in
structuring ecological communities has become a central issue in community ecology
,Diamond, 195, Connor & Simberlo, 199,. Species that coexist in an area do so by
dierging in their resource use oer an eolutionary period, largely in response to
selection pressures generated by competition. Such adaptie diergence in resource use
by co-occurring species is called resource partitioning ,Schoener, 194, \alter, 1991,,
and has been documented in seeral taxa ,1ot, 1985 in reptiles and amphibians,
Albrecht & Gotelli, 2001 in birds, Namgail et al., 2004 in mammals,. lacilitation is also
thought to be an important process determining herbiore species richness in grazing
ecosystems ,Arsenault & Owen-Smith, 2002,. Although it may enhance species richness
in communities in high productie areas in the 1ropical systems ,luisman & Ol, 1998,,
it is less important in structuring herbiore assemblages in less productie ecosystems
where competition tends to be the dominant orm o interaction ,Mishra et al., 2002,
Mishra et al., 2004, Namgail et al., 200,.
1he Ladakh region ,32
o
to 36
o
N and 5
o
to 80
o
L, 80,000 km
2
, o the Indian
1rans-limalaya is a high altitude cold desert, but supports a rich assemblage o eight
wild ungulates, perhaps due to its location at the junction o two biomes: 1ibetan plateau
and lindukush-Pamir mountains. Amongst these ungulates, two are listed as
Lndangered`, two as Near 1hreatened` and one as Vulnerable` on the IUCN red-list o
threatened animals ,IUCN, 2008,. All o them except the blue sheep hae relatiely
patchy distributions, generating a spatial heterogeneity in herbiore diersity ,lox et al.,
1991, Chundawat & Qureshi, 1999,. Preious studies hae documented the importance
o liestock grazing as an important actor restricting wild ungulate populations in the
region ,Bagchi et al., 2004, Mishra et al., 2004, Namgail et al., 200, Namgail et al., 2008,.
loweer, the importance o competition amongst the wild ungulates themseles has not
been explored as a potential mechanism or species distributions ,but see Namgail, 2006,.
Unlike in tropical enironments, the plants in the 1rans-limalaya are stunted and
sparsely distributed, and the simple egetation structure does not appear to allow niche
partitioning amongst large herbiores, as seen in tropical systems ,Voeten & Prins, 1999,
Cromsigt & Ol, 2006,. 1hereore, instead o plants, the terrain eatures are more
important in niche separation amongst the herbiores in the region ,Namgail et al., 2004,
Namgail, 2006,.
1he Ladakh urial has a small population ,F2,000, in Ladakh, conined to narrow
tracts along the banks o the Indus and Shayok riers, while the blue sheep is the most
abundant ,F11,000, and widely distributed wild ungulate ,lox et al., 1991,. Amongst all
the wild ungulates, the Ladakh urial suered the most due to anthropogenic pressure
associated with modern deelopment as it inhabits the lowest slopes along the Srinagar-
Leh highway, the lieline o Ladakh ,Mallon, 1983,. 1he species was persecuted by
Chapter 6
84
hunters or trophy and meat due to its occurrence along the highway ,Mallon, 1983,.
Apart rom these the animal also suered due to pasture degradation associated with
excessie liestock grazing ,Chundawat & Qureshi, 1999,, and persecution by armers as
it reportedly damage their crops. Nonetheless, although these pressures might hae led to
decline in its density, the range o the species hae historically been restricted ,Mallon,
1983,. Currently, although urial`s distribution has penetrated the tributary alleys o the
Indus and Shayok, it has not progressed beyond F 15 km rom the alley-mouths
,Mallon, 1983,. 1his leads to the obious question o what restricts the distribution o
the species to this narrow belt.
Interspeciic competition is a possible actor, whereby more abundant species like
the blue sheep competitiely exclude the Ladakh urial, or preent its range expansion.
1hereore, we asked i the limited distribution o Ladakh urial can be explained by
possible competition with the blue sheep, gien that amongst all the wild large herbiores
o the region, the body masses o these two species ,mean ~ 52 kg, are most similar
,Mishra et al., 2002,, and thus they probably hae similar ecological requirements. 1heory
suggests that species that share ecological eatures may compete and coexist either by
geographical partitioning or resource partitioning along one or more resource axes
,Schoener, 194,. 1hus we hypothesised that blue sheep and Ladakh urial hae non-
oerlapping geographical distributions and,or dierging resource utilization patterns at a
local scale. \e aimed to understand the nature o interaction ,in terms o habitat and
diet, between the two species at smaller spatial scales and its implications or their
occurrence at larger spatial scales, which will shed light on the possible role o
competition with blue sheep in hindering the range expansion o urial. Understanding the
actors limiting urial distribution is crucial gien that the species is endemic to Ladakh
and is listed as an Lndangered` species on the IUCN red-list o threatened animals
,IUCN, 2008,.
0DWHULDOVDQG0HWKRGV
6WXG\DUHD
1he Ladakh Region ,F 80,000 km
2
, is a high altitude cold desert in the northern Indian
state o Jammu and Kashmir. \ithin this area, Ladakh urial occupy F 3000 km
2
.
Lleation in Ladakh ranges rom 2900 m to 000 m. 1he region is depried o the
monsoon clouds due to the rain-shadow eect o the Greater limalaya, and hence has
ery low primary productiity. 1he growth season is conined to a short period o 3-4
months during summer. 1rees like poplar 3RSXOXVspp. and willow 6DOL[spp. are conined
to oases along glacier-ed streams and riers.
1he Puyul alley ,33
o
43`N,
o
4`L ,, the habitat scale study site in the proposed
Gya-Miru \ildlie Sanctuary, encompasses F100 km
2
and is located at F15 km rom the
Indus Rier at Upshi. 1he alley marks the boundary o urial distribution along the
tributary stream o Gya. Lleation in the area ranges rom 3900 to 6500 m asl, and
proides dierse habitats ranging rom rugged tracts at higher altitudes and relatiely
open areas in lower areas. 1emperature ranges rom -30
o
C in winter to -30
o
C in
summer, and egetation is characterised by dry alpine steppe ,Rawat & Adhikari, 2005,.
1here are no other wild ungulates in the area except the study species, and a small
population o domestic sheep and goats ,F00, that grazes the area. 1here are howeer
seeral small mammals: 1ibetan woolly hare /HSXVRLRVWROXV mouse hare 2FKRWRQD sp. and
marmot 0DUPRWD EREDN that share the pastures with the study species. Mammalian
predators include the snow leopard 8QFLDXQFLD, wol &DQLVOXSXVlynx /\Q[OLVDEHOOLQD and
red ox 9XOSHVYPRQWDQD.

85
)LHOGPHWKRGV
1o determine the co-occurrence o blue sheep and urial at a geographical scale, a grid
,20x20 km, was oerlaid on a map o Ladakh, and presence-absence o the two species
was determined in 136 random grid-cells scattered all oer Ladakh. Co-occurrence at the
landscape leel was assessed by oerlaying a grid ,10x10 km, on a map o Ladakh urial`s
potential range, and then determining the presence-absence o blue sheep and urial in 22
randomly located grid-cells. Most o these grid-cells were located on the ground and the
presence-absence o the species in them was determined by direct obseration o the
animals as well as by indirect eidence such as the presence o horns etc. Inormation on
presence-absence o a species was also gleaned rom the literature as well as rom
knowledgeable local people and wildlie oicials.
lor the determination o habitat use at a local scale, data were collected between
June 2006 and March 200. lerds o blue sheep and urial were obsered rom trails
along alley bottoms and ridgelines. Scan sampling was the primary method or animal
obserations, which were aided by 8x40 binoculars and a 15-45X spotting scope. Records
were made o species type, group size and date.
labitat ariables YL] altitude, distance to cli, slope angle and aspect at the
animal locations were recorded. 1hese ariables hae been identiied as important in
determining habitat use and partitioning by large herbiores in the 1rans-limalaya
,Chundawat & Qureshi, 1999, Namgail et al., 2004,. Altitude was determined rom a
topographic map, while slope angle and distance to cli were isually estimated. A cli
was deined as a ery steep slope ,45
o
, on an area more than 20 m diameter with
ertical drops o more than 5 meters. Slope aspect was determined by using a compass.
lresh aecal pellets were collected to generate diet proiles o the species and to
assess the diet oerlap. 1o preent assigning pellets mistakenly to a dierent species than
the one intended to, we collected them rom bedding sites by waiting or the animals to
get up and moe away. A group o F 150 pellets was collected rom each herd o the
animals. Subsequently, ie pellets were randomly drawn rom each group to orm one
sample or the respectie herd. 1hus there were 11 samples or blue sheep and 10
samples or Ladakh urial, which were air-dried and stored in paper bags.

/DERUDWRU\PHWKRGV
1he dried aecal samples were boiled in water or about 1 h, soaked oernight, and then
crushed. 1he inner tissue was separated rom the epidermis and cuticle by mixing a 5 g
subsample with water or 1 min in a \aring blender, and was strained oer a plankton
siee ollowing de Jong et al. ,2004,. 1he residue was then washed again with tap-water,
transerred into a petri-dish and allowed to settle. Using a Pasteur pipette, ten random
grab samples o the residue were then taken, and each droplet was put on a glass slide,
spread out eenly and coered with a 2.4 cm coer slip.
\e prepared separate reerence slides or the plant parts such as lea, stem,
lower and seeds. lor this, small pieces o plant parts were cleaned in household bleach
oernight, washed in water, and then ragments o epidermis were stripped o and
mounted in glycerol ,de Jong et al., 2004,. Photomicrographs o epidermal material on a
set o these reerence slides were used to identiy the ragments o cuticles obsered in
samples o the animal aeces. At least 100 cuticle or epidermal ragments were identiied
in each sample. 1o quantiy the composition o the aecal material the area o epidermal
ragments was measured at a magniication o 100-X using a grid o small squares ,0.01
mm
2
, in the microscope eyepiece. 1he abundance o each species was calculated as a
percentage o the total area o the ragments measured ,Cid & Brizuela, 1990, Alipayo et
al., 1992,.

Chapter 6
86
'DWDDQDO\VLV
1o assess the co-occurrence at geographical and landscape leels, our null model
assumed that blue sheep and urial are randomly associated. 1he presence-absence data
were organised into a matrix ollowing Connor and Simberlo ,199,, where the row
represents a species and the column a grid-cell. 1he species` co-occurrence was
quantiied using the Cscore index ,Stone & Roberts, 1990,, calculated as Ci; ~ ;ri);r;),
where r
i
is the number o grid-cells with species i and r
;
the number o grid-cells with
species ; and being the number o shared grid-cells. 1his index quantiies the
checkerboard units` ,.ev.v Diamond, 195, or the species pair, and the larger the index
the less co-occurrence o the species. Signiicance o the obsered Cscores was assessed
through Monte Carlo simulations ,1000 iterations, using the co-occurrence module o
Lcosim .2 sotware ,Gotelli & Lntsminger, 2001,. Obsered C-score is signiicantly
smaller than expected at random, when P ,ObseredLxpected, 0.05.
lor assessing the dierential habitat use by the two species, we irst identiied the
most important ariables in habitat choice o each species at a local scale. lor this we
used Generalized Linear Model ,GLM, by taking used and unused ,but aailable, habitats
as a binary response ariable and distance to cli, slope angle, aspect and altitude as
predictor ariables. Subsequently, we used Akaike`s Inormation Criterion or small
samples ,AIC
c
, and their dierences ,A, to select the most parsimonious model with
ewest ariables ,lower the A, more parsimonious the model, that explain most o the
ariation in the data. All models with AIC
c
dierences ,, o less than two are useul in
explaining the ariability in the data.
Subsequently we perormed Discriminant Analysis to determine whether the
areas used by blue sheep, urial and the unused but aailable ones could be discriminated
on the basis o the most crucial ariables identiied. \e tested or signiicant dierences
between these areas on the canonical scores o the irst two unctions or axes with a one-
way ANOVA ollowed by o.t boc lisher`s LSD test. Signiicant dierences between blue
sheep and urial habitat use were also assessed by using t-tests or independent samples.
1he multiariate lotelling`s 1
2
test was used to check or dierences taking all the
ariables together and thus taking into account the relationship between them. 1he
habitat-niche ,all our ariables, and diet-niche oerlap between the species was
determined using Pianka`s Index ,Pianka, 193,.

=
2 2
.
.
Pik Pij
Pik Pij
Ojk Lqn. 1

where O
;/
is the measure o oerlap between species ; and /, and P
i;
and P
i/
are the
proportions o taxon i in the diet o species ; and / respectiely. Oerlap is complete
when O
;/
~ 1 and absent when O
;/
~ 0.

5HVXOWV
/DUJHVFDOHGLVWULEXWLRQV
At the geographical scale ,entire Ladakh, blue sheep occurred in 62 o the 132 grid-cells
sureyed, while Ladakh urial occurred in only 10 grid-cells. At the landscape scale ,in and
around urial`s range,, blue sheep occurred in 11 grid-cells while the Ladakh urial occurred
in 13 o the 22 grid-cells sureyed. 1he co-occurrence analyses showed that blue sheep
and urial are distributed independently at the geographical scale ,C-score ~ 110, P ~
0.82,, but their co-occurrence was signiicantly higher than expected by chance at the
landscape leel ,C-score ~ 48, P 0.05,.

87
+DELWDWVFDOH
Akaike Inormation Criterion or small sample size ,AIC
c
, indicated that altitude is the
most important ariable determining habitat choice by urial during summer as well as
winter ,1able 1,, while blue sheep habitat use is best modeled by using distance to cli,
slope angle and altitude as predictors during summer, and distance to cli and altitude
during winter ,1able 2,.

1able 1. Akaike`s inormation criterion scores ,AICc ,, their dierences ,, and number
o model parameters ,N, or habitat models deeloped or seasonal habitat use by the
Ladakh urial in Gya-Miru, Ladakh. 1he igures in bold are AIC dierences ,, o less
than two.

Summer Winter No. Model K
AICc A AICc A
1 Altitude 2 65.30 0.00 56.0 0.00
2 Distance - Altitude 3 65.1 0.4J 5.92 J.86
3 Distance - Slope - Altitude 4 6.02 J.72 59.92 3.86
4 Slope - Altitude 3 6.06 J.76 58.0 2.00
5 Altitude - Aspect 3 4.52 9.21 59.22 3.15
6 Distance - Altitude - Aspect 4 4.4 9.44 60.69 4.62
Distance -Slope - Altitude - Aspect 5 5.49 10.18 62.60 6.53
8 Slope - Altitude - Aspect 4 5.1 10.41 61.18 5.11
9 Slope 2 142.52 .21 100.24 44.1
10 Distance - Slope 3 144.46 9.16 90.35 34.29
11 Distance 2 145.09 9.9 88.46 32.39
12 Slope - Aspect 3 149.56 84.26 86.68 30.62
13 Aspect 2 150.0 85.39 84.9 28.90
14 Distance - Slope - Aspect 4 151.52 86.21 1.6 15.60
15 Distance - Aspect 3 152.09 86.8 69.69 13.62

Aspect was not important in the habitat choice o both species, and hence was dropped
rom urther analysis. Based on the AIC
c
dierences ,,, the models in 1able 1 are
ranked rom the best ,lowest alue, to the worst ,highest alue, or habitat use by
urial during summer, while or winter they are arranged haphazardly or the sake o
comparison with the summer data. 1he habitat models or blue sheep in 1able 2 are also
arranged in a similar ashion.

Habitat use and partitioning
6XPPHU
Discriminant Analysis showed that altitude had the highest loading on unction 1 in the
habitat use during summer ,1able 3,, indicating that it was the most important actor in
discriminating the locations used by urial rom those used by blue sheep and unused but
aailable locations. lig. 1 suggests that blue sheep during summer used higher areas
which were more aailable, whereas Ladakh urial occurred in the lower areas. 1hese
dierences were signiicant when these areas were contrasted on the basis o the
canonical scores o the irst two unctions or axes with a one-way ANOVA ,l~35.96, d
~ 4, P0.001,.

Chapter 6
88
1able 2. Akaike`s inormation criterion scores ,AICc ,, their dierences ,, and number
o model parameters ,N, or habitat models deeloped or seasonal habitat use by the
blue sheep in Gya-Miru, Ladakh. 1he igures in bold are AIC dierences ,, o less than
two.

Summer Winter No. Model K
AICc A AICc A
1 Distance - Slope - Altitude 4 130.22 0.00 118.18 J.96
2 Distance - Slope - Altitude -Aspect 5 132.66 2.44 122.12 5.90
3 Distance - Slope - Aspect 4 134.11 3.89 143.85 2.62
4 Slope - Altitude 3 134.39 4.1 139.25 23.02
5 Distance - Slope 3 134.48 4.26 15.54 41.31
6 Slope - Altitude - Aspect 4 136.35 6.13 143.04 26.81
Distance - Altitude 3 138.92 8.0 116.23 0.00
8 Slope - Aspect 3 139.42 9.20 192.23 6.00
9 Slope 3 140.54 10.32 210.22 93.99
10 Distance - Altitude - Aspect 4 143.4 13.25 120.16 3.93
11 Distance 2 144.2 14.05 155.5 39.34
12 Distance - Aspect 3 146.02 15.80 141.95 25.3
13 Altitude 2 150.08 19.86 13.93 21.0
14 Altitude - Aspect 3 153.0 22.85 141.5 25.52
15 Aspect 2 159.24 29.01 198.55 82.32
Comparing the habitat use o blue sheep and urial with a paired t-test also showed a
similar trend ,t ~ .92, P ~ 0.0001, 1able 4,, which was urther strengthened by the little
oerlap between the two species along an altitudinal gradient ,2
MN
~ 0.222,. Similarly, blue
sheep used habitats signiicantly closer to clis ,mean ~ 114 m,, while urial selected
habitats away rom clis ,mean ~ 209 m, t ~ 3.24, P ~ 0.001, 1able 4,. 1hese dierences
were signiicant taking all the ariables together ,l
3, 80
~ 23.028, P 0.001,.

1able 3. Standardized coeicients o discriminant unction coeicients o seasonal
habitat use by Ladakh urial and blue sheep in Gya-Miru, Ladakh.
Summer \inter Variable
Root 1 Root 2 Root 1 Root 2
Distance to cli ,m, 0.053 0.621 -0.55 -0.448
Slope angle ,deg, 0.093 -0.63 -0.093 0.85
Altitude ,m, -0.993 -0.153 -0.4 0.53

1able 4. Mean ,- SD, o the habitat eatures used by the blue sheep, Ladakh urial and
those aailable.
Var. Blue sheep Ladakh urial Aailable
Summer \inter Summer \inter
Mean SD Mean SD Mean SD Mean SD Mean SD
D1C 114
109.13
60
65.09
209
156.61
99
128.0
238
184.8
SA 34 .55 31 10.26 30 9.41 2 13.62 2 8.12
Alt 4530
341.15
4160
465.19
4051
164.94
4113
10.3
462
351.85
D1C ~ Distance to cli, SA ~ Slope angle, Alt ~ Altitude

89
Blue sheep
Ladakh urial
Available
-5 -4 -3 -2 -1 0 1 2 3 4
Decreasing Altitude
-3
-2
-1
0
1
2
3
4
5
D
e
c
r
e
a
s
i
n
g

S
l
o
p
e

A
n
g
l
e
lig. 1. Output o Linear Discriminant lunction Analysis to examine whether the areas
used by Ladakh urial, blue sheep and those aailable during summer could be
discriminated on the basis o habitat eatures in the areas.
Blue sheep
Ladakh urial
Available
-6 -5 -4 -3 -2 -1 0 1 2 3 4
Decreasing Altitude
-3
-2
-1
0
1
2
3
4
5
I
n
c
r
e
a
s
i
n
g

S
l
o
p
e

A
n
g
l
e
lig. 2. Output o Linear Discriminant lunction Analysis to examine whether the areas
used by Ladakh urial, blue sheep and those aailable during winter could be discriminated
on the basis o habitat eatures in the areas.
Chapter 6
90
:LQWHU
Altitude and slope angle had highest loadings on unction 1 and unction 2, respectiely,
during winter ,1able 3,, indicating that these ariables were important in discriminating
between the three locations: blue sheep, urial and aailable ,random,. lig. 2 suggests that
both blue sheep and urial used signiicantly lower areas than those aailable ,l~32.02, d
~4, P0.001,. 1hereore, they did not dier in habitat use on an altitudinal gradient
during winter ,t ~ 0.452, p ~ 0.652,, as also indicated by the greater habitat oerlap along
this axis ,O
MN
~ 0.885,. 1he animals were also similar in the use o slope angle ,t ~ 1.405,
p ~ 0.163, O
MN
~0.965,, although they diered only marginally in the use o distance to
cli ,t ~ 1.931, p ~ 0.056, O
MN
~0.890,.

'LHWSURILOHDQGRYHUODS
Diets o blue sheep and urial were dominated by non-graminoids ,1able 5,. Ladakh urial
consumed a high proportion o generatie parts ,lowers, ruits and seeds, o the plants.
1he diet spectrum o blue sheep encompassed six species o graminoids and 16 species
o non-graminoids, while that o urial encompassed 5 species o graminoids and 8 species
o non-graminoids. \ithin non-graminoids, 7KHUPRSVLVsp. ,20, and $UHQDULDsp. ,1.6,
were the most dominant species in blue sheep`s diet, while &DUDJDQDsp.,21, and 5XPH[
sp. ,11., were the most dominant in urial`s diet ,1able 5,. Although both species
consumed a greater proportion o non-graminoids, they ed on dierent plant species as
indicated by the less oerlap in diet ,2
MN
~ 0.293,.

'LVFXVVLRQ
Blue sheep and urial are distributed independently at the geographical scale, which
conorms to the disjoint potential distributions o these species modeled in a GIS
enironment ,Chundawat & Qureshi, 1999,. 1his distributional independence could be
related to their diering biogeographical ainities, urial haing adanced into Ladakh
rom the western lindukush mountains by penetrating the Indus and Shayok alleys
,Schaller, 19,, and blue sheep colonizing the region rom the eastern 1ibetan plateau
,Namgail et al., 2004,. But when we zoomed in at the landscape leel, the two species co-
occur, oten inhabiting the same catchments. \hen their distribution interace was
analyzed more closely, they were obsered to dierge in their resource use at the habitat
leel. Ladakh urial used lower areas whereas blue sheep occurred in higher reaches during
summer. Such habitat partitioning, which is known to preent both resource and
intererence competition, might hae allowed their co-occurrence at the landscape leel.
1hey howeer oerlapped in habitat use during winter when the blue sheep
descended to lower slopes due to snow accumulation in the higher areas. 1hus, urial
might be at a disadantage gien that blue sheep has a higher density ,lox et al., 1991,,
perhaps due to their ersatility in resource use ,Chapter 5 and ,. 1hus, since winter is
the season with a seere resource crunch in the 1rans-limalaya due to plant senescence
and heay snow coer ,Mishra, 2001, Namgail, 2006,, there is a high potential or
competition between them during this season. At least ie groups o urials were
obsered to leae the study area during this season, perhaps in response to the high
density o blue sheep in the lower areas.
1he altitudinal separation between the two species during summer could be
related to competitie exclusion o urial rom the higher areas, which are considered to
be more proitable in terms o energy gain as higher habitats hae more nutritious plants
during this season. 1his is tenable because large herbiores do take adantage o such
altitudinal dierences in the aailability o energy sources by moing to higher reaches
where resh plants sprout later in the summer ,lesta-Bianchet, 1988,. 1he higher reaches

91
are also cooler and ree rom insects, which are known to harass wild ungulates
,lagemoen & Reimers, 2002,.

1able 5. Proportion o plant ragments in the diet o Ladakh urial and blue sheep during
summer in Gya-Miru, Ladakh.
3ODQWV %OXHVKHHS /DGDNKXULDO
*UDPLQRLGVYHJHWDWLYH
&DODPDJURVWLVsp. 1.6 12.0
'DFW\OLVsp. 2.3 0.0
(O\PXVsp. 2.3 1.1
)HVWXFDRYLQDsp. 4.9 9.5
6WLSDsp. 2.1 3.1
Unidentiied grass 2.3 6.5
*UDPLQRLGVJHQHUDWLYH
Glumes 0.5 1.6
lruits 0.0 0.9
1RQJUDPLQRLGYHJHWDWLYH
$FRJRQXPsp. 0. 0.0
$OOLXPsp. 0.5 0.0
$UHQDULD&HUDVWLXPsp. 1.6 0.0
$UWHPLVLDsp. 5.8 4.3
%LHEHUVWHLQLDsp. 0.5 0.0
,ULVsp. 0. 0.0
/DYDQGXODsp. 1.2 0.9
/RQLFHUDsp. 1.2 0.0
0DOYDsp. 0.2 0.0
1HSHWDsp. 0. 0.0
2[\WURSLVsp. 0.9 0.0
3RO\JRQXPsp. 2.8 0.0
7KHUPRSVLVsp. 20.0 0.0
&DUDJDQDsp. 3.2 21.2
(SKHGUDsp. 0.0 0.5
Dicot stems 5.3 0.
Unidentiied dicot 1 3.9 2.9
Unidentiied dicot 2 1.2 0.0
1RQJUDPLQRLGJHQHUDWLYH
$UWHPLVLD lower heads 0.0 4.
Asteraceae stems,lower head 1.4 2.9
5XPH[ lower and stem 3.9 11.
8UWLFD lower 0.0 0.4
9HURQLFD ruit 0.0 0.4
Unidentiied ruit 2.3 3.6
Unidentiied lower 0.0 0.5
Seeds 0.9 0.0
corky stem,scale,ruit 1.6 8.4
2WKHUV
Unidentiied cuticles .4 2.3

Although both animals included a higher proportion o non-graminoids in their diets
during summer, they ed on dierent species within this unctional group. 1hus, the two
species dierged in their dietary preerences during summer, perhaps as a consequence o
Chapter 6
92
the dierential habitat use and dierential aailability o orage plants on an altitudinal
gradient. Ladakh urial incorporated a remarkable proportion o generatie parts such as
lowers, ruits and seeds o non-graminoids, presumably to aoid competition with blue
sheep.
1hus, although the two species might co-occur in some alleys as a result o the
summer resource partitioning, competition with blue sheep during winter might be
hindering the growth o urial population, thereby making it diicult or the species to
expand its range urther. It is, howeer, to be noted that the urial range in Ladakh, being
located along two major riers, are also ertile and is used or cultiation and limited
grazing by domestic liestock ,Raghaan, 2003,. 1here is also a highway between Leh,
the capital city o Ladakh and Srinagar, the capital o Kashmir passing through urial
habitat along the Indus alley, which made their habitat more accessible to hunters
,Mallon, 1983,. 1hereore, although anthropogenic pressure might hae played a crucial
role in decimating its population more recently, urial`s range historically might had been
constrained by the presence o blue sheep.

&RQFOXVLRQV
Blue sheep and Ladakh urial associated independently at a geographical scale ,entire
Ladakh,, but they co-occurred at the landscape leel. An inestigation o the resource
utilisation pattern by the two species at a smaller ,habitat, scale showed that the animals
partition resources associated with their habitat segregation along an altitudinal gradient
during summer, as the blue sheep occurred at higher areas than urial during this season.
Such a separation at the habitat leel might hae enabled co-occurrence at the landscape
leel. Neertheless, the two species oerlapped in their habitat use during winter when
the blue sheep descended to lower slopes due to high snow coer in the higher reaches,
which appeared to displace urials rom their preerred sites. Such displacement o urial
during winter with a resource crunch suggests a competitie interaction, which might
ultimately be constraining the range o Ladakh urial despite the niche separation during
summer. Keeping these in iew, it is crucial to look or areas with less abundance or
absence o common species like blue sheep, i possible, when it comes to prioritising
areas or the conseration o endangered Ladakh urial in the 1rans-limalayan
mountains.
$FNQRZOHGJHPHQWV
1he study was inanced by \ildlie Conseration Society, Ruord loundation and
\ageningen Uniersity. \e thank Christine de Jong or help in laboratory and \ash
Veer Bhatnagar or support. \e thank the oicials o the Jammu and Kashmir \ildlie
Protection Department or granting research permission, and Karma Sonam, Mipham,
Gyalson and Namgyal or assistance.

5HIHUHQFHV
Albrecht, M. & Gotelli, N. J. ,2001, Spatial and temporal niche partitioning in grassland
ants. 2HFRORJLD 134-141.
Alipayo, D., Valdez, R., lolecheck, J. L. & Cardenas, M. ,1992, Laluation o
microhistological analysis or determining ruminant diet botanical composition.
-RXUQDORI5DQJH0DQDJHPHQW 148-152.
Andrewartha, l. G. & Birch, L. C. ,1954, 7KH GLVWULEXWLRQ DQG DEXQGDQFH RI DQLPDOV
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Arsenault, R. & Owen-Smith, N. ,2002, lacilitation ersus competition in grazing
herbiore assemblages. 2LNRV 313-318.
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Mediated by Local leterogeneity: An Lxperimental Approach. cotog,, 87, 1532-
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de Jong, C. B., an \ieren, S. L., Gill, R. M. A. & Munro, R. ,2004, Relationship
between diet and lier carcinomas in roe deer in Kielder lorest and Galloway
lorest. 1eterivar, Recora, JSS, 19-200.
Diamond, J. M. ,195, In cotog, ava erotvtiov of covvvvitie. ,eds M. L. Cody & J. M.
Diamond,, larard Uniersity Press, Massachusetts, USA.
lesta-Bianchet, M. ,1988, Seasonal range selection in bighorn sheep: conlicts between
orage quality, orage quantity, and predator aoidance. Oecotogia, 7S, 580-586.
Gotelli, N. J. & Lntsminger, G. L. ,2001, coiv: ^vtt voaet. .oftrare for ecotog,. 1er.iov
.2. .cqvirea vtettigevce vcororatiov ava Ke.e,ear
btt:,,rrr.vrv.eav,~biotog,,acvtt,,Cotetti,Cotetti.btvt>>.
lagemoen, R. I. M. & Reimers, L. ,2002, Reindeer Summer Actiity Pattern in Relation
to \eather and Insect larassment. ]ovrvat of .vivat cotog,, 7J, 883-892.
luisman, J. & Ol, l. ,1998, Competition and acilitation in multispecies plant-
herbiore systems o productie enironments. cotog, etter., J, 25-29.
animals .vericav âtvrati.t, 93, 145-159.
IUCN ,2008, IUCN, Gland, Switzerland.
Row.
Mallon, D. ,1983, 1he status o Ladakh urial Ori. orievtati. rigvei in Ladakh, India. iotogicat
Cov.erratiov, 27, 33-381.
Namgail, 1. ,2006, \inter labitat Partitioning between Asiatic Ibex and Blue Sheep in
the 1ibetan gazelle Procara icticavaata in Ladakh, northern India: towards a
Chapter 6
94
1ibetan argali Ori. avvov boag.ovi and blue sheep P.evaoi. va,avr in the Indian
1rans-limalaya. ]ovrvat of Zootog,, 262, 5-63.
Namgail, 1., lox, J. L. & Bhatnagar, \. V. ,200, labitat shit and time budget o the
Ol, l. & Ritchie, M. L. ,1998, Lects o herbiores on grassland plant diersity. 1reva.
iv cotog, ava rotvtiov, J3, 261-265.
Pianka, L. R. ,193, 1he structure o lizard communities. .vvvat Rerier of cotog, ava
,.tevatic., 4, 53-4.
Raghaan, B. ,2003, vteractiov. betreev tire.toc/ ava aaa/b vriat ;Ori. rigvei rigvei). Master`s
Dissertation, Saurashtra Uniersity.
Rawat, G. S. & Adhikari, B. S. ,2005, lloristics and distribution o plant communities
Schoener, 1. \. ,194, Resource partitioning in ecological communities. cievce, J8S, 2-
39.
Stone, L. & Roberts, A. ,1990, 1he checkerboard score and species distributions.
Oecotogia, 8S, 4-9.
1ot, C. A. ,1985, Resource Partitioning in Amphibians and Reptiles. Coeia, J, 1-21.
\alter, G. l. ,1991, \hat is resource partitioning ]ovrvat of 1beoreticat iotog,, JS0, 13-
143.

Synthesis
95
CHAP1LR 7

Mammalian herbivore species-richness: synthesis

1sewang Namgail

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Burton lillis

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Karnataka, India

Chapter 7
96
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Despite being one o the oldest patterns obsered in the ield o ecology, the spatial
heterogeneity in species diersity on earth and the processes that generate it hae
remained unresoled. Lcological literature is replete with accounts o species diersity
patterns and processes underlying them, listed in the beginning ,Chapter 1,, but hitherto
there is no consensus on the primary mechanisms. It is stressed in this thesis that the
issue is rather complex and its resolution will require a new synthesis drawing upon
disciplines like ecology, eolution, biogeography, systematics, phylogeography and
paleontology that deeloped during the last century but drited apart at the same time.
Because the phenomenon o species diersity spans such dierse ields, the
comprehensieness o the research eorts required to tackle the phenomenon must
equally be great. 1his thesis is written in the hope that the contents will generate more
light than heat on this general, but important discourse that transcends the ield o
ecology.
Since the distribution and diersity patterns o mammalian herbiores o the
limalaya and the regions beyond remain poorly understood, I ocused on the
mammalian herbiore distribution and diersity patterns in the Ladakh region o the
Indian 1rans-limalaya, about which ery little is known. Lcological inormation on the
processes underlying herbiore assembly patterns in the region hae started to come in
only recently ,Namgail et al., 2004, Namgail, 2006, Namgail et al., 2008,. 1he results o
this research project hae implications not only or understanding the ormation and
maintenance o herbiore assemblages in the region but also or deeloping conseration
strategies or the threatened mammalian herbiores.
1his section o the thesis takes a synthetic approach, drawing on the results
presented in the arious chapters as well as on results rom my preious inestigations on
the mammalian herbiores in the highlands o Ladakh. Unlike my preious approaches
o ocusing on single assemblages, the present inestigation stried to understand the
mechanisms o mammalian herbiore distribution and species-richness pattern at larger
geographical scales. 1o address arious biogeographical issues in the region, I capitalised
on both niche theory ,MacArthur & Leins, 196, and macroecological theory
,MacArthur & \ilson, 196, Brown & Maurer, 198,.
Perhaps the best way to start will be with a tribute to the our 19
th
century
scientists: Charles Darwin, Joseph looker, Alred Russel \allace and Philip Sclater, who
reolutionised our understanding o the spatial heterogeneity in biological diersity on
the planet. 1hey shared a common interest o understanding the origin, diersiication
and distribution o the world`s biota. Darwin deoted seeral years o his early carrier
inestigating geology and natural history around the world, which later led to ormulation
o the 1heory o Natural Selection. \allace traeled widely around the world collecting
specimens, and proposed a diision o the world in six biogeographical realms. Sclater
collected inormation on the distribution o birds, and proposed a diision o the world
on the basis o aiaunal distribution. looker was the irst botanist who attempted to
understand the plant diersity pattern on earth, and was the irst Luropean to collect
plants in the limalayas.
1he limalayan mountains represent a treasure-troe o lora due to the great
altitudinal gradient that proides aried habitats to plants. 1he loral diersity o the
limalayan range is increasingly being studied. One o the issues that are being
inestigated currently is the plant diersity along altitudinal gradients. But hitherto most
o the studies ocused on patterns at smaller scales with steep altitudinal gradients, and it
is not known as to what pattern emerges, i we inestigate this relationship at a regional
scale with a gradual altitudinal gradient. 1his is where Chapter 2 o this thesis contributes,
demonstrating that plant diersity has a hump-shaped relationship een at a regional
Synthesis
97
scale with moderate altitudinal gradient oer hundreds o kilometers. lurther, although
spatial distribution o aboeground plant biomass has been inestigated in the 1rans-
limalaya as well as other grazing ecosystems, we still need to understand how it aries
along an altitudinal gradient. It is generally presumed that it has a negatie relationship
with altitude, but it might not be the case in mountain desert ecosystems with simple
egetation structure and high topographical heterogeneity. Chapter 2 also addresses this
important issue, and shows that aboeground plant biomass in dry alpine areas aries
unimodally with altitude, which is in contradiction to the general belie that it declines
monotonically with altitude. 1he chapter discusses this unexpected trend, especially in
light o the biome idiosyncrasies and increasing liestock population in Ladakh.
Subsequently, I inestigated the herbiore distribution and diersity patterns in
Ladakh. Most o the biogeographical studies on herbiores were carried out in single
ecosystems or biomes, but the distribution and diersity pattern o herbiores are less
known in transition areas between discrete biomes. Recognising the act that Ladakh is
located at the intersection o two biomes, Chapter 3 stries to ill this gap in inormation.
1he work presented in the chapter inestigated as to how mammalian herbiores with
dierent biogeographic ainities interact, i.e., whether they mingle or orm separate
groups on the basis o their eolutionary histories. It became apparent that mammalian
herbiores in Ladakh orm geographical groups, which underscores the act that it is
important to examine the biogeographical backgrounds o species to understand
herbiore assembly patterns at local scales, presented in Chapter 4 and 6.
Ater studying these large scale biogeographic patterns, I shited my ocus on
niche-related diersity patterns. Studies addressing herbiore assembly patterns in the
past examined the niche relationships only between constituent species in a single
assemblage, but it is not known as to how the niche o a species can ary across
assemblages with arying number o sympatric species. Chapter 5 stries to understand
this issue, and reports the niche dynamics o blue sheep 3VHXGRLV QD\DXU in response to
herbiore species-richness. 1his chapter along with Chapter 4 and 6 also underline the
importance o interspeciic interactions in species distributions. It thus emphasises the
need to include interspeciic competition in distribution models, which has oten been
neglected, thereby oerestimating species` distributional ranges.
At a local scale, the essence o herbiore species assembly structure is the co-
existence o inading herbiores with the natie ones. \hen an inading species is
similar to the natie herbiores in its ecological requirements, either one natie species
that is less eicient than the inading species in extracting resources or the inading
species will be excluded rom the assemblage oer a period o time, unless they ind a
way to partition resources amicably, promoting long-term coexistence. In high altitude
enironments, interspeciic competition might become more intense in winter when the
resource leel is at its minimal due to plant senescence and snow coer. 1hus the
presence o one species might constrain the distributional range o another. Lmpirical
support or this is presented in Chapter 4.
Some species are more eicient in utilizing scarce resources and competitiely
exclude other species. Since the comprehension o negatie inluence o dominant
species on the distribution o subordinate species is crucial or understanding spatial
heterogeneity in herbiore species-richness, the issue is urther explored in Chapter 6,
which addresses the interaction between two similar-sized Caprinae species at dierent
spatial and temporal scales. 1he study was designed recognising the act that hitherto
most o the studies that inestigated species coexistence were carried out at a single
spatial scale. 1his study highlights the act that species co-existence is contingent upon
spatial scale o analysis, they co-occur at larger spatial scales, but segregate at smaller
Chapter 7
98
spatial scales. lurthermore, een at smaller spatial scales, species coexist in some seasons
when resources are abundant but not in others when there is resource scarcity.

*HRJUDSK\RIVSHFLHVULFKQHVV
Biogeography has a long history as a branch o science, striing to understand the spatial
patterns o biological diersity. Neertheless, it remained relatiely dormant or some
time because it was labeled as a descriptie science until Robert l. MacArthur and
Ldward O. \ilson breathed new lie into it with their amous Island Biogeography
1heory ,MacArthur & \ilson, 196,. Subsequently, some biogeographers also used
experimental approaches to address biogeographical issues ,Simberlo & \ilson, 1969,.
In the ollowing decades there was a renaissance o the ield due largely to parallel
deelopments in macroecology ,Brown, 1985,. Currently, biogeography is an emergent
and synthetic discipline, relying on ecology, eolutionary biology, population biology,
systematics and earth sciences.
1he relegation o biogeography in the past was also related to the diiculty
associated with testing the eects o historical actors on species distribution and
diersity patterns. loweer, recent deelopment in tools in the ield o genetics,
paleontology, geology, systematics, spatial ecology ,Remote Sensing and GIS, and robust
computer sotwares hae enabled analyses that can account or these actors. It is,
howeer, diicult to do controlled experiments in biogeography because o its
maniestation at large spatial and temporal scales. Nonetheless, gien the current scenario
o climate change that aects systems at large spatial scales, it is crucial to approach
systems at macroecological scales ,Chapter 2 and 3,.

lig. 1. Schematic representation o herbiore species-richness pattern and the underlying
processes in the Indian 1rans-limalaya. 1he oal-shaped boxes represent patterns, the
rectangular boxes are processes ,arranged rom top-global to bottom-local, and the
octagonal boxes are the major plausible causes. 1he patterns and processes in bold texts
are the ones dealt with in this thesis.
Local
berbivore species
ricbness
Regional Species
pool
Facilitation Competition
Immigiation
Species loss Species gain
Babitat gain Babitat loss
Resource
Pieuationpaiasitism

Poaching Livestock

Climate change
Biogeograpby

Extinction Speciation
Synthesis
99
Neertheless, it is important to note that local-scale processes ,Chapter 4 and 6, are also
important in understanding large-scale distribution and diersity patterns o mammalian
herbiores. lig. 1 proides a schematic representation o how these patterns and
processes are interdependent.
In the ollowing sections, I establish the releance and contribution o the
arious chapters o this thesis to the literature on herbiore distribution and diersity
patterns, especially in alpine enironments. lirst, I discuss the plant diersity pattern
along altitudinal gradients at dierent spatial scales, and then mammalian herbiore
distribution and species-richness patterns in the light o their eolutionary history and
current ecological actors.

%LRJHRJUDSK\RI/DGDNK
3K\WRJHRJUDSK\
Understanding distributional patterns o plants at dierent spatial scales is crucial or
understanding the biogeography o herbiores. Larlier, plant diersity was thought to
decrease monotonically with altitude ,Steens, 1992,, but numerous studies on the
subject showed that plant diersity peaks at the mid-altitude instead at lower altitudes.
1his led to a lurry o research in the mountainous regions all across the planet, and most
o the studies showed the same hump-shaped relationship, and any deiation rom this
has largely been attributed to discrepancies in sample size and eorts. Majority o the
studies were also carried out at smaller spatial scales with steep altitudinal gradients. I
inestigated this relationship both at local and regional scales to see i dierent patterns
emerge. It is to be noted that the altitudinal gradient in the larger scale inestigation is
ery gradual, spanning hundreds o kilometers. 1he results indicated that plant species-
richness peak at mid-altitudes at local as well as at large geographical scales ,Chapter 2,.
Although, spatial pattern o net primary productiity and its relation with climatic
actors was studied in 1rans-limalayan mountains ,\ang et al., 2009, and other
grassland ecosystems ,Sala et al., 1988, Lpstein et al., 199, Jobbagy et al., 2002,, the
pattern o aboeground biomass along an altitudinal gradient has not been explored
beore in a spatially explicit way, neither in the 1rans-limalaya nor, to my knowledge,
elsewhere. Altitude, as alluded to earlier, is an important gradient along which
aboeground biomass is expected to decline due to declining temperature and nutrients,
which get washed down ,Rastetter et al., 2004,. But there has been no apparent eort to
document this with empirical eidence, and it is less clear i such a trend can be expected
in drier enironments o deserts with little complexity in egetation structure. 1he
present study has shown that aboeground biomass o plants in drier alpine regions
aries unimodally with altitude ,Chapter 2,. \hen the geography o the species richness
o large herbiores in Ladakh was examined, I ound more species in central Ladakh,
which alls in the middle o the altitudinal gradient used in the study o plant diersity
pattern at the regional scale. 1hus, it is likely that there is a positie relationship between
plant species richness and,or productiity and herbiore species richness. Analysis on
this association is underway, which will shed light on the dynamics o phyto- and
zoogeography.

=RRJHRJUDSK\UROHRIHFRWRQH
Ladakh`s location at the junction o two biomes makes it an interesting system or
studying the distribution and diersity pattern o mammalian herbiores. 1hese two
biomes are the ast and open 1ibetan plateau in the east, encompassing F2.5 million km
2

and the rugged lindukush-Karakoram mountains ,including Ladakh and Zangskar
ranges, in the west. 1hese mountains in the latter biome orm a jumble o mountains,
also encompassing the ends o seeral other ranges: Kunlun Shan, Pamir, limalaya and
Chapter 7
100
1ian Shan. 1hese mountain ranges perhaps played a greater role in shaping the
biogeography o central Asia than biogeographers think. 1oday, many large herbiore
species are present on the Mongolian plateau north o the Kunlun Shan but are absent
rom the 1ibetan plateau. Similarly, seeral species with the cores o their ranges in the
limalaya and the Gangetic plains are absent rom the 1ibetan plateau ,see lig. 2,.
1hereore, it seems likely that the 1ibetan species deeloped as a result o icariance,
which needs to be explored. 1he Kunlun in the north and limalaya in the south
probably isolated the large herbiores o the 1ibetan plateau oer an eolutionary period.
Although the species expanded their ranges westwards, they are perhaps stopped by the
aorementioned jumble o mountains. Similarly, there are seeral large herbiores
including Moulon 2YLVURLHQWDOLVthat are distributed in the drier enironments o the west
Asian deserts, but are missing rom Ladakh. 1his could be due to the aorementioned
jumble o mountains hampering their eastward progress ,see urther discussion below,.

L
a
u
a
k
h
C
a
s
p
i
a
n

s
e
a
Aiabian sea
Bay of Bengal
L
ake
Ba
ikal
L
a
k e
B
a
lk
a
sh
Reu ueei
Nountain gazelle
Nouflon
Chiltan goat
Nusk ueei
Naikhoi
Black buck
Inuian gazelle
Nilgai
Bimalayan Tahi
uoial Takin
Pizewalskis gazelle
Nongolian gazelle
Wilu hoise
uoiteieu gazelle
uobi khulan
Cential Asian wilu boai Euiasian wilu boai
Saiga antelope
Aiabian sanu gazelle

lig. 2. Some Palaearctic and Oriental large herbiores ,positioned in the map in relation
to their approx. distributional ranges, that are missing rom the herbiore species-pool o
Ladakh.

1he geographical groupings o mammalian herbiores in Ladakh on the basis o
their biogeographic ainities ,Chapter 3, are telltale signs o the termination o their
range expansions due to high and ormidable mountain ranges. 1hus, hard boundaries do
seem to inluence herbiore species distribution and diersity patterns as predicted by
Colwell & Lees ,2000,. 1he two large groups o herbiores with similar distributions in
Ladakh might correspond to the two biomes that meet in Ladakh, as mentioned aboe,
which was presumed earlier, but was neer tested. 1he importance o topographical
actors such as slope angle and altitude in explaining the shared distributions o these
herbiores urther support this speculation. Riers are also likely to aect large herbiore
distributions, but they do not seem to inluence their distributions in Ladakh ,Chapter 3,,
perhaps because they reeze during winter allowing the animals to cross oer.
Synthesis
101
1ransition zones between biomes, or ecotones, are known to harbour unique and
endemic species and alleles, which may support the notion that ecotones sere as centers
o speciation ,\iens & Graham, 2005,. Lolutionary adanced species occur in
transitional ecotones in central Arica ,ljeldsa & Rahbek, 1998,. Moritz et al. ,2000,
urther proposed that areas such as the central Arican ecotones hold concentrations o
young species and eolutionary noelty, and are important or maintaining the
eolutionary processes. 1he Ladakh urial 2YLV YLJQHL YLJQHL, which is an endemic and
endangered species, is the most adanced wild sheep in the eolutionary history o the
genus 2YLV,Geist, 198,. 1his eolutionary noelty could be related to Ladakh`s location
at the interace between two biomes. Currently the species has a narrow distribution only
along two major riers and its range expansion is perhaps hindered by the abundant and
similar-sized blue sheep adancing into Ladakh rom the 1ibetan plateau in the east
,Chapter 6,. 1hus, the Ladakh urial is perhaps trying to ind a niche in an assemblage
with species co-eoled oer a long eolutionary period ,see urther discussion below,.
Because enironmental conditions generally closely correspond to a species`
ecological requirements at the center o its range ,Brown, 1984,, the enironmental
suitability or a species is expected to decline towards the periphery o its range. Most o
the mammalian herbiore populations in Ladakh represent species` populations at the
edge o their ranges ,Namgail, 2009,. 1hereore, the less population densities o large
herbiores in the region ,lox et al., 1991, could be attributed to their locations at the
edge o species` ranges. lurthermore, gien that the presence-absence o a species in an
area essentially relects the balance between colonization and extinction ,Rickles, 198,
luston, 1999,, the recolonisation potential o a species is also expected to decline
towards the edge o its range ,MacArthur, 192,. 1hus, the probability o occurrence o a
species ,i.e., density, at regional scales is expected to decline rom the center o its range
to the periphery due to a decline in the recolonisation potential. 1he ragmented
populations o mammalian herbiores in Ladakh engendering a spatial heterogeneity in
their species-richness could also be due to such edge related eects.

,QIOXHQFHRIHYROXWLRQDU\KLVWRU\
1here are myriad studies on the role o physical and biological actors in determining
herbiore distribution and species richness patterns, but the role o eolution and
historical distributions in the ormation and maintenance o herbiore assemblages has
not been addressed adequately. 1his is intriguing, gien that the taxonomic components
o a community organisation can be comprehended only by placing the local community
in its historical and biogeographical context ,Chapter 3,. One cannot understand the
geographical ariation in species diersity solely by studying dynamical interactions
among species within assemblages, as in Chapter 4. Geological, eolutionary and
historical actors play important roles in determining niche occupancy, assembly
structures and perhaps other attributes o herbiore communities, as alluded to earlier.
1he specialization o a species as a result o long-term eolution o its
morphology in a particular topographic setup limits its dispersal capability. Recent
examination o skeletal morphology o the Caprinae species in Ladakh has reealed that
they are dierent in terms o their appendicular structures associated with adaptations to
dierent topographical eatures ,Van den 1empel & De Vrij, 2006,. 1hus, distributions
o the mammalian herbiores o Ladakh are perhaps determined by niche conseratism,
and species ranges are restricted to areas with enironmental eatures to which they are
best adapted through long-term eolution. 1he ormation o two large geographical
groups o mammalian herbiores in Ladakh ,Chapter 3, could be related to this act.
1he small mammalian herbiores such as pikas 2FKRWRQD spp. and oles $OWLFROD
spp. in the central Asian rangelands orm taxa that remain little-known. It is, howeer,
Chapter 7
102
important to know their distributions and diersity pattern, because they enhance plant
species richness in the 1rans-limalayan rangelands ,Bagchi et al., 2006,, and hae been
identiied as indicator species o pasture degradation in the rangelands o central Asia
,lolzner & Kriechbaum, 2001,. Chapter 3 demonstrates that there is congruence in the
distributions o large herbiores and these smaller mammalian herbiores. Despite the
caeats mentioned in the Chapter, the biogeographical analysis o these little known taxa
in the context o oerall herbiore assemblage in Ladakh has important implications or
understanding their distribution pattern, which has not been addressed beore.

5ROHRILQWHUVSHFLILFFRPSHWLWLRQ
1he role o competition in structuring animal assemblages has been disputed or oer
three decades. It is, thereore, one o the most contested topics in ecology, controersy
raged oer its supremacy in structuring ecological communities, and acrimonious
discussions ensued amongst ecologists ,Diamond, 195, Connor & Simberlo, 199,
Gotelli & McCabe, 2002,. 1he competition drien assembly rules` suggested by
Diamond were downplayed by Connor and Simberlo, who suggested using null models
to test or non-random co-occurrence o species. lollowing this debate, myriad studies
were carried out to assess the role o competition in structuring species assemblages, but
most o them inestigated niche dynamics in single assemblages, and it is not known as
to how an indiidual species` niche can ary across assemblages responding to the
number o competing species in those assemblages.
1oday, there is consensus amongst ecologists that when two species rely on the
same resources that are in short supply, the subordinate species usually get competitiely
excluded ,Chapter 4 and 6,. A species cannot inhabit all the aailable habitats, and thus
occupies only a subset o the aailable habitat, oten reerred to as the 'realized niche'
,lutchinson, 1959,. 1hereore, assuming that all animals hae similar growth rates,
dispersal abilities and there are no geographical barriers, the structure o herbiore
assemblages is largely determined by species coexistence. loweer, whether two species
co-exist or not also depends on the spatial scale, as species do co-occur at large
geographical scales, but oten separate along one or more niche axes at the local scale
,Chapter 6,.
1he blue sheep is the most abundant and widespread wild ungulate in Ladakh
,Namgail, 2009,, perhaps as a result o the lexibility in terms o resource use compared
to rare and endangered species ,Chapter 6,. \hen the diet width o this species and that
o Ladakh urial were studied in two locations with dierent number o herbiore
sympatric species, it had a narrower diet width in the area with high number o species,
whereas urial`s diet width did not dier between the two areas ,see lig. 3,, suggesting
that blue sheep is lexible in its resource use. lurther, blue sheep specialized by
consuming both egetatie and generatie parts o plant species in areas with higher
number o sympatric species ,lig. 5,. 1hereore, owing to its ersatility in resource
utilisation, blue sheep is likely to outcompete other similar-sized species with dierent
eolutionary histories. 1hereore, I studied its interaction with other Caprinae species:
Asiatic ibex ,Chapter 4, and Ladakh urial ,Chapter 6, which are similar to blue sheep in
their body sizes but hae dierent biogeographic ainities ,Chapter 3,. Blue sheep has
high potential or competition with both these species especially in winter, when the
resources are at their minimal leels due to plant senescence and snow coer. 1he study
on blue sheep and Ladakh urial was carried out at multiple spatial scales ,Chapter 6,, and
the role o spatial scale in species co-existence became apparent as the two species
associated randomly at the geographical scale but they co-occurred at the landscape leel
as a result o resource partitioning at the local-leel habitat.
Synthesis
103
2.2
2.3
2.4
2.5
2.6
2.7
2.8
2.9
2 3
N
i
c
h
e

w
i
d
t
h
Blue sheep
Ladakh urial

lig. 3. 1he diet-niche width o blue sheep and Ladakh urial in relation to the number o
sympatric large herbiore species in the 1rans-limalayan mountains ,Note the sharp
decline in blue sheep`s niche width,.

Urial has a ery limited distribution in Ladakh, and is endemic to the region. It is
probably an eolutionary noel species, trying` to it in a large herbiore assemblage that
eoled oer eons as speculated earlier. I inestigated this by studying the species packing
in the large herbiore assemblage in Ladakh, including hare and marmots. I related the
natural log-transormed body weight with species rank on the basis o weight ratio, irst
without urial and then with urial in the assemblage. I ound a slightly stronger
relationship in the imaginary assemblage without urial than that in the assemblage with
urial ,lig. 4,, suggesting that the species perhaps did not eole with the rest o the large
herbiore species in Ladakh.
y = 0.5026x + 1.4736
R
2
= 0.957
y = 0.5949x + 1.3359
R
2
= 0.9768
0
1
2
3
4
5
6
7
0 2 4 6 8 10
Species rank
I
n

b
o
d
y

m
a
s
s

lig. 4. Relationship between species rank ,increasing weight by a constant proportion,
and log-transormed body masses o large herbiores in Ladakh. 1he open circles
represent the assemblage without Ladakh urial, and the illed circles represent the
assemblage with Ladakh urial.

1he potential or competition in the 1rans-limalayan large herbiores is urther
indicated by the expansion o blue sheep`s niche width with decrease and contraction
Chapter 7
104
with increase in the number o sympatric species in an assemblage ,Chapter 5,. 1his
relationship, howeer, diered between the habitat-niche width and diet-niche width.
1he habitat width declined monotonically with the number o sympatric species, whereas
the diet width had a hump-shaped relationship with the number o symptaric species.
1he ormer pattern is understandable that as the number o sympatric species increases
in an assemblage there would be less habitat-space aailable or a particular species, but
the hump-shaped relationship between diet-niche width and herbiore species richness is
intriguing. I suggest that the narrow diet width in allopatry is out o choice as the animal
can eat the most nutritious plants aailable to itsel, and the narrow diet width in species-
rich areas is out o necessity as it needs to narrow down its diet spectrum, eeding on
ewer plant species that are spared by the competitors ,see Chapter 5,. 1hus, the animal
seems to specialize on some plant species in species-rich assemblages, perhaps eeding on
both egetatie and generatie parts such as lowers and seeds ,see lig. 5,.

0
10
20
30
40
50
60
70
Graminoid
vegetative
Non-graminoid
vegetative
Non-graminoid
generative
Others
Diet composition
P
e
r
c
e
n
t
a
g
e
Gya-Miru
Hemis

lig. 5. Diet proile o blue sheep in Gya-Miru with one sympatric species and lemis with
two sympatric species ,Note that in lemis it is consuming a higher proportion o
generatie parts such as lowers and seeds, perhaps to aoid competition with other
species,.

Chapters 4, 5 and 6, thus, indicate that competition might play a role in structuring
herbiore assemblages at local scales ,alpha diersity, in Ladakh. 1hese results also hae
wider implications or understanding large herbiore assemblages in other alpine
ecosystems, and warrants inclusion o interspeciic interactions in species distribution
models, which until now were constructed largely on the basis o the aailability o the
physical enironment. Consequently, many species distribution models tend to
oerestimate the range o a species, as the animals actually do not occur in some parts o
the predicted range because potentially competing species exclude them rom those
potential areas. lurthermore, it became apparent that topography is more important in
resource partitioning by large herbiores in the high altitude drier enironments o
Ladakh with less complexity in egetation structure ,Namgail et al., 2004, Chapter 3 to 6,.
1he discourse in Chapter 5 takes us back to the concept o undamental and realized
niches propounded by Lelyn lutchinson ie decades ago ,lutchinson, 1959,: species
cannot occupy their ull potential ranges ,undamental niche, because they are not able to
surie in areas with strong competitors, and thus they need to conine themseles to
Synthesis
105
smaller habitats ,realized niche,. Most o the species distribution models till now
predicted the distribution in terms o undamental niche, but what is needed is the
distribution in terms o the realized niche to delineate more accurate ranges o species.
Although blue sheep might interact negatiely with similar-sized species rom
other biomes ,Chapter 4 and 6,, it does not compete with similarly-sized species rom the
same biome, perhaps by irtue o their shared eolutionary history. lor instance, a
resource partitioning study between the 1ibetan argali 2YLVDPPRQKRGJVRQLand blue sheep
in the past showed that these species are dierent in terms o their use o terrain eature
with the ormer using more open areas and the later using rugged terrain ,Namgail et al.,
2004,, which resulted in dierntial use o plant communities. 1his is perhaps determined
by their skeletal morphology eoled oer a long eolutionary period ,Van den 1empel
& De Vrij, 2006,. 1hus, the 1ibetan argali`s range is perhaps constrained due to niche
conseratism. loweer, they might hae competed in the past and eoled dierent
morphological eatures, but this cannot be proen as we cannot in retrospect see i
species competed in the past or not.
'LGFRPSHWLWLRQVKDSHWKHUHJLRQDOVSHFLHVSRRO"
1he current regional herbiore species-pool in Ladakh is perhaps a culmination o
competition amongst wild herbiores oer eons. Otherwise, why are there only 12 large
herbiores , 2 kg, in Ladakh Gien that the region is located at the junction between
the Oriental and Palaearctic biogeographic realms, one would expect the region to hae
at least one large herbiore rom the Oriental, but there is none. All are Palaearctic
except one: Cape hare rom the Lthiopian realm. lurther, why not more rom the
Palaearctic, which has scores o large herbiores in areas with almost similar
enironmental conditions as those o Ladakh. lor instance, the Przewalski`s horse (TXXV
FDEDOOXV SU]HZDOVNLL and Mongolian gazelle 3URFDSUD JXWWXURVD that occur in south-western
Mongolia ,F 2000 km rom Ladakh, are absent rom Ladakh ,see lig. 2,. It is possible
that the Przewalski`s horse was competitiely excluded by the almost same-sized kiang
(TXXV NLDQJ, and the Mongolian gazelle was outcompeted by the similar-sized 1ibetan
antelope 3DQWKRORSVKRGJVRQL.
Similarly, species such as the Indian gazelle or Chinkara *D]HOODEHQQHWLL that occur
in the Gangetic plains south o the limalaya do not occur in Ladakh ,lig. 2,. Many
species like the musk deer 0RVFKXV FKU\VRJDVWHU and limalayan tahr +HPLWUDJXV MHPODKLFXV
that are distributed in the limalaya are also missing rom the regional species pool o
Ladakh ,lig. 2,. Probably, the 1ibetan gazelle 3URFDSUDSLFWLFDXGDWD and Asiatic ibex &DSUD
LEH[VLEHULFDrespectiely,out-competed these species rom Ladakh. A small population o
musk deer was present in western Ladakh until ery recently ,Pister, 2004,. It is possible
that the adenturers` could not establish iable populations due to lack o
metapopulation dynamics as moement o the animals were restricted by the high
mountain passes. Another possibility is that the animals occurring at lower latitudes and
altitudes in the Indian Peninsula are physiologically constrained, as they cannot store at
to surie in the extremely low temperatures o the 1rans-limalaya. One might also ask
why megaherbiores like the elephants (OHSKDV PD[LPXV LQGLFXV and rhinoceros 5KLQRFHURV
XQLFRUQLV are not present in Ladakh. 1he apparent answer to that would be inadequate
resources or these animals to surie.
Another interesting act about the regional species pool o Ladakh is that there
are two antelopes rom the east: 1ibetan gazelle and 1ibetan antelope, but not a single
one rom the west. Seeral antelopes occur in the Arabian Peninsula including the
Mountain gazelle *D]HOOD JD]HOOD FRUD, Arabian sand gazelle *D]HOOD VXEJXWWXURVD PDULFD and
Goitered gazelle *D]HOOD VXEJXWWXURVD, but are missing rom the herbiore species pool o
Ladakh. 1his might relect the possibility that high mountains in lindukush and Pamir
Chapter 7
106
ranges hindered their eastward progress, urther suggesting the role o high mountains as
geographical barriers. 1hese need to be explored to understand the herbiore diersity
patterns as well as eolution and dispersal routes o mountain ungulates in central Asia.

7KLVWKHVLVDQGELRJHRJUDSKLFPDFKLQHU\
larking back to the objects o this research, i.e., understanding ,1, species richness
pattern, and ,2, the plausible causes operating at local and regional scales, I now examine
with the help o a conceptual model whether these hae been achieed and where do the
chapters contribute towards understanding the oerall machinery o biogeography ,see
lig. 6,. Chapter 2 explores the plant species-richness and aboeground biomass pattern
at local as well as geographical scale ,objects 1 and 2,. Chapter 3 contributes towards
understanding the herbiore species-richness pattern ,object 1,. Chapter 4 helps in
understanding the cause o species richness at a local scale ,object 2,. Chapter 5
augments our understanding o the plausible causes o herbiore species-richness at a
regional scale ,object 2,. Chapter 6 explores co-existence o herbiores at both local and
regional scales ,objects 1 and 2,.

lig. 6. Map showing the contributions o arious chapters to the broad scheme o
biogeography ,see lig. 1 or the meaning o the shapes o dierent compartments,.
0DQDJHPHQWLPSOLFDWLRQV
1he loss o herbiores rom the ace o the planet has become a major concern and
eorts to stem such loss are stymied by lack o inormation on proximate and ultimate
actors inluencing the spatial pattern o herbiore diersity at dierent spatial and
temporal scales. Large herbiores are especially threatened due to a plethora o human
endeaours, and gien that large herbiores play a unique role in ecosystem unctioning,
are economically and aesthetically important ,Gordon et al., 2004,, there is an urgent
need to understand the mechanisms goerning their species-richness pattern, so that
Local heibivoie
species iichness
Facilitation Competition
Immigiation
Species loss Species gain
Babitat gain Babitat loss
Pieuationpaiasitism

Poaching Livestock

Climate change
Biogeogiaphy

Extinction Speciation
Ch
Ch
Ch
Resouice
Ch
Ch
Ch
Regional Species
pool
Synthesis
107
their loss rom grazing ecosystems can be stemmed. 1his thesis contributes towards
deeloping conseration strategies or the large herbiores in the 1rans-limalayan
region o Ladakh, which are highly threatened, and some are on the erge o local
extinction ,Namgail, 2009,.
Amongst the eight wild ungulates o Ladakh, two are listed as Lndangered`, two
as Near 1hreatened` and one as Vulnerable` on the Red List o 1hreatened Species by
the International Union or Conseration o Nature ,IUCN, 2008,. Seeral smaller
mammalian herbiores are also threatened due to human endeaours associated with
liestock grazing ,Bagchi et al., 2006,, yet inormation on their distributions and diersity
patterns is ery scanty ,Chapter 3,. 1he population o seeral wild ungulates, especially
those o the 1ibetan gazelle and 1ibetan argali declined precipitously in the last century
due to habitat degradation associated with liestock grazing and many other natural and
manmade actors ,Bhatnagar et al., 2006, Namgail et al., 200b, Namgail et al., 2008,.
Currently, these and seeral other species hae highly ragmented populations in Ladakh
as well as globally ,Schaller, 19, Schaller, 1998, Namgail, 2009,. 1he results o this
project contribute towards understanding the distributional pattern and plausible causes
o the ragmented nature o their distribution, which will help in prioritising areas or
their conseration.
Chapter 2 underscores the importance o mid-altitude areas or the conseration
o loral wealth o Ladakh, and perhaps herbiore species. lence, mid-altitude areas are
needed to be considered irst while prioritising areas or protection o lora in Ladakh.
Chapter 3 emphasises the importance o the topography more than the climatic actors in
determining the distribution and diersity patterns o mammalian herbiores in Ladakh.
lurther, it shows that when research and monitoring are constrained by time and money,
the large and prominent herbiores such as ungulates can be used as proxy species to
prioritise areas or the conseration o smaller herbiores such as pikas and oles
,diicult to locate in the ield,, because there is a congruence in the distribution o these
herbiorous taxa.
Chapters 4 through 6 emphasise the importance o interspeciic competition in
the present distributional patterns o mammalian herbiores in Ladakh. 1hey showed
that abundant and wide-spread species like the blue sheep are lexible in their resource
use, and can competitiely exclude other species such as the Asiatic ibex and Ladakh urial
that are perhaps more specialized in their habitat and diet use. Such competitions are
more likely between species that are similar in body-size but are dierent in their
biogeographic ainities. lor instance, both blue sheep and 1ibetan argali that hae the
core o their ranges on the 1ibetan plateau, suggesting similar eolutionary histories, do
not compete, and partition resources on the basis o the physical habitat ,Namgail et al.,
2004,. 1hus, aboe acts need to be considered while deeloping conseration strategies
or endangered species like the Ladakh urial.
1he niche shit o wild herbiores ,Chapter 5, in response to the number o
sympatric species is o concern to wildlie conserationists largely in the light o
increasing liestock population in the region ,Namgail et al., 200a,. Domestic liestock
deplete rangeland resources ,Namgail et al., 2008, and competitiely exclude wild
herbiores rom some rangelands ,Mishra et al., 2002, Mishra et al., 2004,. Liestock
grazing also relegates the wild herbiores to sub-optimal habitats, thereby aecting the
ital actiities important or their growth and reproduction ,Namgail et al., 200b,.
Almost all the protected areas in the 1rans-limalaya are grazed perasiely by domestic
liestock, which thwarts conseration eorts. Gien these and the results o chapter 5,
liestock grazing should be curtailed or stopped in ecologically sensitie areas with
endangered wild species.
Chapter 7
108
)XWXUH'LUHFWLRQV
Despite the arious caeats, the arious chapters in this thesis contribute towards
understanding distribution and community dynamics o large herbiores in Ladakh, and
more importantly sere as baseline inormation or urther studies on the herbiore
assembly rules and biogeography o the 1rans-limalaya and perhaps other alpine
regions. 1he thesis also seres as a precursor to studies exploring the origin o limalayan
ungulates, their eolution and routes o dispersal, which hitherto remain unclear. 1he
limalayan region is thought to be the cradle o eolution o the sub-amily Caprinae
,Schaller, 19,. Unortunately the Caprinae arose and diersiied during a turbulent
period when climates changed and the earth heaed and olded. Being essentially adapted
to hills and clis, the distributional patterns o the animals were thus altered, and since
bones are seldom presered in mountains the Caprini hae reealed little o their ossil
past ,Schaller, 19,.
Not much has been done on the geographical distribution o Caprinae and the
associated taxa, because o the remoteness and inaccessibility o the mountainous areas
where they inhabit. Understanding the geography o the diersity pattern o the 1rans-
limalayan mammalian herbiores including the caprids, their eolution and dispersal
routes is also constrained by lack o inormation on their phylogeny. 1hereore, this
particular ield needs to be researched urther by taking adantage o the recent
deelopments in the ields o genetics and molecular biology.
1o understand the community dynamics o mammalian herbiores, it is crucial to
know whether herbiore species richness also aects the secondary productiity, i.e.,
whether it aects the population dynamics o the constituent species o an assemblage.
\e know that plant species richness enhances aboeground biomass ,Spehn et al., 2000,
and consumer species richness ,\right & Samways, 1998,, but it is not known i
consumer species-richness leads to a greater consumer biomass. 1here are two possible
mechanisms that can lead to this process: ,a, higher herbiore species-richness leads to
greater eiciency in utilizing the primary productiity, and ,b, acilitation o one
herbiore species by another or complete exploitation o the primary productiity leads
to higher consumer biomass irrespectie o the energy transer eiciency ,Prins & lritz,
2008,. 1he apparent assumption is that all the members o the assemblage are similar in
body size.
Although there are seeral other ields that need attention, it is not possible to list
them all here. 1hereore, inally, the metapopulation dynamics o large herbiores in the
1rans-limalaya needs to be explored, which remains elusie not only in this assemblage
but also in assemblages in other ecosystems ,see Llmhagen & Angerbjorn, 2001,. 1his
lack o inormation is largely related to the long generations o large herbiores, and the
diiculty associated with dealing with large spatial scales. But with the deelopment o
macroecology, landscape genetics and spatial statistics, it is becoming more easible to
study population dynamics o large herbiores at regional and continental scales.
Metapopulation studies are especially releant in the herbiore assemblage o the 1rans-
limalaya, where species hae highly ragmented distributions. Such studies will help not
only in understanding their phylogeny and dispersal routes, but also in establishing
protected areas and corridors linking them.

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Bhatnagar, \. V., \angchuk, R. & Mishra, C. ,2006, Decline o the 1ibetan gazelle
Procapra picticaudata in Ladakh, India. 2U\[ 40, 229-232.
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Brown, J. l. ,1985, Macroecotog,, Uniersity o Chicago Press, Chicago.
Brown, J. l. & Maurer, B. A. ,198, Lolution o species assemblages: eects o
energetic constraints and species dynamics on the diersiication o North
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Colwell, R. K. & Lees, D. C. ,2000, 1he mid-domain eect: geometric constraints on the
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Connor, L. l. & Simberlo, D. ,199, 1he assembly o species communities: chance or
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Diamond, J. M. ,195, In cotog, ava erotvtiov of covvvvitie. ,eds M. L. Cody & J. M.
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Llmhagen, B. & Angerbjorn, A. ,2001, 1he applicability o metapopulation theory to
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Lpstein, l. L., Lauenroth, \. K. & Burke, I. C. ,199, Lects o temperature and soil
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Gordon, I. J., lester, A. J. & lesta-Bianchet, M. ,2004, 1he management o wild large
herbiores to meet economic, conseration and enironmental objecties. ]ovrvat
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111
6XPPDU\
1he spatial heterogeneity in biological diersity on earth is one o the most ascinating
natural phenomena. Deciphering the underlying mechanisms o this phenomenon is a
major challenge as it is a complex issue that requires insights rom arious biological and
non-biological ields like geology and geography. One o the most discernible
biodiersity patterns is the declining biological diersity as one moe rom the tropics to
the temperate and arctic regions. Len at a continental scale some biogeographic zones
are richer in biodiersity than others. 1hus the pattern repeats itsel at dierent scales in
a hierarchical ashion. Relying on the niche theory as well as on macroecological theory, I
inestigated the spatial heterogeneity in mammalian herbiore species-richness in a little
known herbiore assemblage o the Ladakh 1rans-limalaya. 1his 1rans-limalayan
region in north Indian state o Jammu and Kashmir is located at the junction o two
biogeographic realms: Palaearctic and Oriental. At a smaller spatial scale, it orms an
ecotone between the 1ibetan plateau in the east and the lindukush-Karakoram
mountains ,including the Ladakh and Zangskar ranges, in the west.
1he study took a multi-scale approach by irst understanding the broad
phytogeographical and zoogeographical patterns and subsequently studying the
mechanisms operating at smaller spatial scales. At the outset, I inestigated the diersity
and aboeground biomass patterns o ascular plants in relation to altitude. 1he diersity
and biomass were spatially structured at both local and regional scales. At the local scale
both graminoid and non-graminoid species-richness had a hump-shaped relationship
with altitude. 1he phytodiersity also aried unimodally along an altitudinal gradient at a
regional scale. lurther, the aboeground biomass also had a non-linear relationship with
altitude, which is contradictory to the popular belie that biomass decreases
monotonically with altitude. Since the sampling was done mostly in high pastures with
liestock grazing, such a non-linear trend could be related to depletion o plants by
liestock at the lower altitudes, which were grazed more intensiely. But the pattern
could also be related to low precipitation at lower altitudes due to higher temperature.
Secondly, I asked i the mammalian herbiores in Ladakh orm chorotypes or
groups that share distributions. I ound that they ormed two large geographical groups
that perhaps correspond to the two biomes: 1ibetan plateau and lindukush-Karakoram
mountains. One o them occurred in the ast eastern plains o Changthang, while the
other occurred in the rugged mountains o north and south-western Ladakh. Such
groupings could be related to niche conseratism, as the species in each group had
similar biogeographic ainities. 1opographical eatures were more important than the
climatic ones in explaining the geographical groupings o mammalian herbiores. Riers
did not inluence the geography o herbiore species-richness. I suggest that the relatiely
high diersity o mammalian herbiores in Ladakh is because o its location at the
intersection o two biogeographical realms and two biomes.
Subsequently, I shited my ocus on patterns and processes operating at local
scales, which might inluence spatial heterogeneity in large herbiore species-richness at
meso- and mega scales. More speciically I studied the nature o interaction between large
herbiores that hae comparable body sizes and perhaps similar ecological requirements.
It became apparent that the ormation and maintenance o large herbiore assemblages
could also be inluenced by competitie interaction between species that are similar in
body sizes but dierent biogeographic ainities. lor instance, the distribution o the
Asiatic ibex &DSUDLEH[VLEHULFDseemed to be constrained by similarly-sized species like the
blue sheep 3VHXGRLVQD\DXU. It is suggested that interspeciic competition can inluence the
assembly patterns o mammalian herbiores in the dry alpine grazing ecosystems.
Recognising the act that studies on niche relationship amongst large herbiores
were carried out only in single assemblages, I also examined the niche dynamics o blue
112
sheep in response to the number o sympatric species in an assemblage. 1he results
showed that a species` habitat-niche width and diet-niche width dier in their
relationships with herbiore species-richness, as the ormer declines monotonically
whereas the latter has a hump-shaped relationship with the number o sympatric species
in assemblages. 1he ormer association is understandable, but it is more diicult to
explain the latter. It is suggested that narrow diet width in allopatry is due to a species`
eeding on the most nutritious plants aailable to itsel, whereas the narrow diet width in
areas with high number o sympatric species is due to a species` specialisation on ewer
plant species that are more proitable as the sympatric species do not preer them.
1he inluence o interspeciic competition on assembly patterns o large
herbiores is urther indicated by the aderse niche relationship between Ladakh urial
2YLVYLJQHLYLJQHL and blue sheep. lirst, the results suggested that the coexistence o the two
species depends on the spatial scale o analysis. lor instance, their distributions were not
related at large geographical scales as they associated randomly, but they co-occurred at
the landscape leel. \hen their distributions and interactions were urther ealuated at
the local habitat leel, I ound that they separate along an altitudinal gradient. Blue sheep
used the higher pastures but it descended to lower slopes during winter when the higher
pastures got coered with thick snow, thereby increasing the potential or competition
with the Ladakh urial that used the lower slopes. 1hus, although they might co-occur at
the landscape leel as a result o the niche separation during summer, there is a high
potential or competition during winter, the pinch period, which perhaps negatiely
inluence the reproductie perormance o Ladakh urial, thereby hampering its range
expansion.

113
6DPHQYDWWLQJ
De ruimtelijke heterogeniteit an de biologische diersiteit op aarde is een an de meest
ascinerende natuurlijke enomenen. let ontcijeren an de onderliggende mechanismen
an dit enomeen ormt een grote uitdaging, omdat het een complex onderwerp is wat
inzichten ergt an erschillende biologische en niet-biologische disciplines zoals
geologie en geograie. Len an de makkelijkst waarneembare patronen is de aname an
biologische diersiteit an de tropen naar de gematigde en Arctische zones. Zels op een
continentale schaal zijn sommige biogeograische zones rijker dan andere. Zo herhaalt
het patroon zich op hirarchische wijze op erschillende schalen. Vanuit de niche theorie
en macro-ecologische theorie heb ik de ruimtelijke heterogeniteit an de soortenrijkdom
an plantenetende zoogdieren onderzocht in een weinig bekende assemblage an
herbioren in Ladakh 1rans-limalaya. Deze 1rans-limalaya regio in de Noord-Indische
staten Jammu en Kashmir beind zich op het grenslak an twee biogeograische rijken:
het Palearctische en het Orientaalse. Op een kleinere schaal ormt het een ecologische
gradint tussen de 1ibetaanse hooglakte in het oosten en het lindukush-Kardakoram
gebergte ,inclusie de Ladakh en Zangskar bergketens, in het westen.
Deze studie olgt een benadering op meerdere schaalnieaus door eerst de
algemene phytogeograische en zoogeograische patronen te proberen te begrijpen en
daarna de mechanismen die op kleinere schaal opereren te bestuderen. In eerste instantie
heb ik de patronen in diersiteit en boengrondse biomassa an aatplanten onderzocht
in relatie tot hoogte. De diersiteit en biomassa waren ruimtelijk gestructureerd op zowel
een lokale als een regionale schaal. Lokaal ertoonden zowel de soortenrijkdom an
grasachtigen als die an niet-grasachtigen een optimum cure in relatie tot de hoogte. De
phytodiersiteit arieerde ook unimodaal langs een hoogtegradint op regionale schaal.
De boengrondse biomassa had een niet-lineaire relatie tot hoogte, in tegenstelling tot de
algemene aanname dat biomassa eenredig aneemt met hoogte. Omdat de bemonstering
oornamelijk plaatsond op hooggelegen weidegronden die door ee worden begraasd,
kan deze non-lineaire trend gerelateerd zijn aan de uitputting an planten op relatie lage
hoogten, waar intensieer begraasd wordt. let patroon kan echter ook gerelateerd zijn
aan de lagere hoeeelheid neerslag op lage hoogten, anwege hogere temperaturen.
Daarnaast heb ik de raag gesteld o de plantenetende zoogdieren an Ladakh
chorotypen o groepen ormen die hun erspreidingsgebied delen. Ik heb geonden dat
zij twee grote geograische groepen ormen, die misschien oereenkomen met de twee
biomen: de 1ibetaanse hooglakte en het lindukush-Kardakoram gebergte. Len an hen
bestrijkt de oostelijke steppen an Changthang, terwijl de andere de ruige bergen an
noord en zuidwest Ladakh beslaat. Len dergelijke groepering zou gerelateerd kunnen
worden aan niche conseratisme, omdat de soorten an beide groepen een soortgelijke
biogeograische ainiteit hebben. 1opograische eigenschappen hadden een grotere
erklarende waarde dan klimatologische eigenschappen oor de geograische groepering
an de plantenetende zoogdieren. Riieren hadden geen inloed op de geograie an de
soortenrijkdom an herbioren. Ik stel oor dat de relatie hoge soortenrijkdom an
plantenetende zoogdieren in Ladakh is te danken aan zijn locatie op het grenslak an
twee biogeograische rijken en twee biomen.
Verolgens heb ik mijn ocus erlegd naar patronen en processen op lokale
schaal, welke de ruimtelijke heterogeniteit in de soortenrijkdom an grote herbioren op
meso- en megaschaal kan benloeden. Ik heb speciiek de aard an de interactie tussen
grote herbioren met ergelijkbare lichaamsgrootte en mogelijk ergelijkbare ecologische
behoeten bestudeerd. let werd duidelijk dat de orming en handhaing an
soortenassemblages ook benloed kon worden door competitiee interacties tussen
soorten met ergelijkbare lichaamsgrootte maar erschillende biogeograische ainiteiten.
De erspreiding an de Aziatische Ibex &DSUD LEH[ VLEHULFD, lijkt bijoorbeeld te worden
114
beperkt door de een grote Blauwschaap 3VHXGRLV QD\DXU. Dit suggereert dat inter-
speciieke competitie de assemblagepatronen an plantenetende zoogdieren in droge
alpine ecosystemen kan benloeden.
Rekening houdend met het eit dat de studies naar niche relaties tussen grote
herbioren slechts in enkele assemblages zijn uitgeoerd, heb ik de nichedynamiek an
Bharal onderzocht in relatie tot het aantal sympatrische soorten in een assemblage. De
resultaten laten zien dat de habitat-nichebreedte en de dieet-nichebreedte erschillen in
hun relatie met de soortenrijkdom an herbioren, omdat de eerste eenredig aneemt
met het aantal sympatrische soorten in een assemblage, terwijl de laatste een unimodale
optimumcure laat zien. De eerste relatie is makkelijk te begrijpen, maar het is moeilijker
om de laatste te erklaren. let suggereert dat een smalle dieet-nichebreedte in allopatrie
eroorzaakt wordt doordat een soort zich oedt met de meest oedselrijke plant die
beschikbaar is, terwijl een smalle dieet-nichebreedte in gebieden met een groot aantal
sympatrische soorten eroorzaakt wordt doordat een soort zich specialiseert in een
kleiner aantal plantensoorten die beschikbaar zijn omdat sympatrische soorten ze minder
graag eten.
De inloed an interspeciieke competitie op assemblage patronen wordt erder
gellustreerd door de tegenoergestelde niche-erhouding tussen Ladakh Oerial 2YLVYLJQHL
YLJQHL, en de Bharal. Aanankelijk leken de resultaten a te hangen an de ruimtelijke
schaal an de analyse. lun erspreiding was bijoorbeeld niet geassocieerd op grote
geograische schaal, waar deze een willekeurig patroon lieten zien, maar op
landschapsnieau kwamen ze op dezelde plekken oor. Na erdere ealuatie an hun
erspreiding en interactie op lokaal habitatnieau, heb ik geonden dat zij zich
onderscheidden langs een hoogtegradint. De Bharal gebruikten de hogere weiden, maar
daalde a in de winter naar lagergelegen hellingen wanneer de hoge weiden bedekt zijn
met een dikke laag sneeuw, waarbij het potentieel oor competitie met de Ladakh Oerial
die met name de lagergelegen hellingen gebruiken, toeneemt. lieruit olgt dat, hoewel ze
samen oorkomen op landschapsnieau als geolg an niche specialisatie gedurende de
zomer, er een groot potentieel is oor competitie gedurende de winter, de kritieke
periode, wat mogelijk een negatie eect heet op de oortplantingsprestaties an de
Ladakh Oerial en een beperking is oor de uitbreiding an het areaal an de Ladakh
Oerial.
115
$IWHUZRUG
1he core o this thesis touches upon an old issue in ecology on which controersy raged
or a long time, namely whether ecological communities are well-deined unctional units
o species that are interdependent or are loose assemblages o independent species
interacting randomly. 1his thesis supports the latter school o thought. Species come
across noel enironmental conditions and either eole morphologically, physiologically
and,or behaiourally, engendering new species oer a period o eolutionary time, or go
extinct. At a shorter temporal scale, indiiduals o a species emigrate rom one
area,population and join new assemblages, and either coexist or compete with the natie
species and consequently exclude one o the natie species or get excluded oer a period
o time.
Such capabilities o species and the spatial heterogeneity in the enironment leae
no room or doubt that ecological communities are open and dynamic assemblages.
1here hae also been propositions that communities act like super organisms, eoling as
a whole, i.e., consistently maintaining the number o species in a particular unctional
group ,e.g., grazers, across ecosystems, and adding or eliminating species depending on
the enironmental scenario. Len i such processes occur in nature, it would be diicult
to detect in the contemporary world because there is no place let on earth where
humans hae not meddled with the system. Neertheless, there would still be remnant
processes that would be worth a look.
One o the implicit objects o this research project was to understand why some
species hae restricted distributions in Ladakh Although this is a parochial question, it
has uniersal releance, and applies to the entire planet earth. lor instance, why
Kangaroos are ound only in Australia \hy Lemurs are conined to Madagascar \hy
snow leopards roam only in central Asian mountains and not in the Luropean Alps
\hy Guanacos did not moe out o South America \hy Giraes are ound only in
Arica 1hese questions lie at the heart o biogeography. In a letter to John Dalton
looker in 1845, Charles Darwin reerred to the study o geographic distribution as
.that almost keystone subject o the laws o creation`. 1his opinion implies that these
questions stimulated the 1heory o Natural Selection.
So what are the plausible answers to these questions 1here are more answers
than one would think exist, o which ew were addressed in this thesis. 1hese include: ,1,
the areas where a species does not occur were historically unsuitable, but became suitable
recently, and species hae not colonized them yet, ,2, Diuse competition preents the
species rom persisting in the gap areas, ,3, 1he acant areas are not habitable, only our
ignorance makes us think otherwise, ,4, 1hese areas had once supported the species,
which went extinct locally, ,5, 1he species in the regional pool are poor dispersers, ,6,
Anthropogenic pressures preented species rom colonizing acant areas. Len i species
did colonize, the adenturers` were drien to extinction beore they could establish a
population, ,, Geographical barriers such as mountains and riers hindered species
moement.
1he aboe answers and numerous others suggest that the issue is complex. 1his
thesis only seres as the irst step o a long journey. Although the questions deal with
undamental issues in ecology, they hae conseration implications. 1hus, it is hoped that
the scientiic knowledge generated through this research would help not only in
understanding the ormation and maintenance o herbiore assemblages but also in
protecting the large herbiores o Ladakh or posterity.

116
117
9,7$
1sewang Namgail was born in Skurbuchan, Ladakh, on the 3
rd
o May 194. le
completed his secondary school education at Goernment lari Singh ligher Secondary
School in 1991, and then B.Sc. at Goernment Gandhi Memorial Science College,
Jammu in 1995. 1hereater, he studied Zoology ,lonours, at the Panjab Uniersity,
Chandigarh, and graduated in the year 1998. le worked on insect pests o poplar trees in
Chandigarh or his M.Sc. dissertation. le then went to the Uniersity o 1romso,
Norway, to pursue an M.Phil. Degree in \ildlie Science. lor the dissertation, he worked
on the 1ibetan argali under the superision o Dr. Joe lox. Subsequently he acquired
inancial support rom seeral national and international organisations to work on
coexistence o mountain ungulates, wildlie-liestock interactions, herbiore behaiour,
herbiore-plant interactions, human-carniore conlicts, breeding biology and assembly
patterns o birds in Ladakh.
le joined the Ph.D. program at the Resource Lcology Group, \ageningen
Uniersity, in September 2005. In addition to scientiic research, he helps local people in
managing natural resources in a sustainable way. le, in collaboration with colleagues, has
started arious conseration programs in Ladakh. le also had a short stint at the \ildlie
Institute o India, continuing the work on 1ibetan argali. le was conerred PL & RC
Publication Award 2008 or the work on argali. Apart rom these, he conducted seeral
sureys in Ladakh to know the status and distribution o mammalian herbiores. le was
interiewed on the local radio and teleision seeral times, and his work was eatured in
seeral Indian newspapers. le is ond o music and plays seeral instruments including
lute and harmonica, and won a bronze medal in a national orchestra competition in New
Delhi. Currently he works with the Nature Conseration loundation, India.
PUBLICATIONS
-RXUQDODUWLFOHV
Namgail, 1., Mishra, C., de Jong, C. B., an \ieren, S.L. & Prins, l.l.1. ,2009,. Lects o
herbiore species richness on blue sheep`s niche dynamics and distribution in the 1rans-
limalaya. 'LYHUVLW\DQG'LVWULEXWLRQV 15: 940-94.
Namgail, 1., Mudappa, D. & Raman, 1.R.S. ,2009,. \aterbird numbers at ligh Altitude
Lakes in Lastern Ladakh, India. :LOGIRZO59: 13-144.

Namgail, 1. & \om-1o, \. ,2009,. Lleational range and timing o breeding in the birds
o Ladakh: the eects o body mass, status and diet. -RXUQDORI2UQLWKRORJ\ 150: 505-
510.

Namgail, 1., lox, J.L. & Bhatnagar, \.V. ,2009,. Status and distribution o the Near
1hreatened 1ibetan argali 2YLV DPPRQ KRGJVRQL in Ladakh, India: eect o a hunting ban.
2U\[43: 288-291.

Namgail, 1. ,2009,. Mountain ungulates o the 1rans-limalayan region o Ladakh, India.
,QWHUQDWLRQDO-RXUQDORI:LOGHUQHVV, 15: 35-40.

Namgail, 1., Bagchi, S., Mishra, C. & Bhatnagar, \.V. ,2008,. Distributional correlates o
the 1ibetan gazelle in northern India: 1owards a recoery programme. 2U\[, 42: 10-
112.
118
Namgail, 1., lox, J.L. & Bhatnagar, \.V. ,200,. labitat shit and time budget o the
1ibetan argali: the inluence o liestock grazing. (FRORJLFDO5HVHDUFK, 22: 25-31.
Namgail, 1., Bhatnagar, \.V., Mishra, C. & Bagchi, S. ,200,. Pastoral nomads o the
Indian Changthang: production system, landuse and socio-economic changes. +XPDQ
(FRORJ\, 35: 49-504.

Namgail, 1., lox, J.L. & Bhatnagar, \.V. ,200,. Carniore-caused liestock mortality in
1rans-limalaya. (QYLURQPHQWDO0DQDJHPHQW, 39: 490-496.

Namgail, 1. ,200,. Vigilance behaiour o the 1ibetan argali 2YLV DPPRQ KRGJVRQL in the
Indian 1rans-limalaya. Acta Zoologica Sinica, 53: 195-200.

Bhatnagar, \.V., Seth, C.M., 1akpa, J., Ul-haq, S., Namgail, 1., Bagchi, S. & Mishra, C.
,200,. A strategy or conseration o the 1ibetan gazelle Procara icticavaata in Ladakh.
Conservation and Society, 5: 262-26.

Bagchi, S., Namgail, 1. & Ritchie, M.L. ,2006,. Small mammalian herbiores as mediators
o plant community dynamics in the high-altitude arid rangelands o 1rans-limalayas.
Biological Conservation, 12: 438-442.

Namgail, 1. ,2006,. \inter labitat Partitioning between Asiatic Ibex and Blue Sheep in
Ladakh, Northern India. Journal of Mountain Lcology, 8: -13.

Namgail, 1., Bagchi, S., Bhatnagar, \.V. & \angchuk, R. ,2005,. Occurrence o the
1ibetan sand ox 1vte. ferritata lodgson in Ladakh: A new record or the Indian sub-
Continent. Journal of Bombay Natural History Society, 102: 21-219.

Namgail, 1. ,2005,. \inter birds o the Gya-Miru \ildlie Sanctuary, Ladakh, Jammu and
Kashmir, India. Indian Birds, 1: 26-28.

Namgail, 1., lox, J.L. & Bhatnagar, \.V. ,2004,. labitat segregation between sympatric
1ibetan argali Ori. avvov boag.ovi and blue sheep P.evaoi. va,avr in the Indian 1rans-
limalaya. Journal of Zoology (London), 262: 5-63

1ewari, P.K. & Namgail, 1. ,1999,. Biology o 1rabala ishnou ,Leeere, ,Lepidoptera:
Lasiocampidae, on Povtv. Linn. Indian Journal of Iorestry, 22: 348-350.

Book Chapter
Mishra, C., Bagchi, S., Namgail, 1. & Bhatnagar, \.V. ,2010,. Multiple use o 1rans-
limalayan rangelands: reconciling human lielihoods with wildlie conseration. In: !ita
Ravgetava.: Cov.errivg !itatife !bite Maivtaivivg ire.toc/ iv evi.ria co.,.tev.. du 1oit,
J.1., Kock, R. & Deutsch, J. ,eds,.\iley-Blackwell, London

Articles in press or review
Namgail, 1., an \ieren, S.L. & Prins, l.l.1. ,In reiew,. Biogeography o mammalian
herbiores in Ladakh: distribution in relation to enironmental and geographical barriers.

Namgail, 1., an \ieren, S.L., Mishra, C., & Prins, l.l.1. ,In reiew,. Is the distributional
range o Ladakh urial constrained by blue sheep in the 1rans-limalayan mountains

119
Namgail, 1., Bhatnagar, \.V. & lox, J.L. ,in press,. Pastoral Production and \ildlie
Conseration in a 1rans-limalayan \ildlie Resere in Ladakh. In: ,VVXHVRI3DVWRUDOLVPLQ
WKH+LPDOD\DQ5HJLRQ. G.B. Pant Institute o limalayan Lnironment and Deelopment.

Morup, 1., Namgail, 1. & 1sering, 1. ,in press,. Liestock s. wildlie: Conlict o
interests on the rangelands o Changthang. Proceedings o the 13th IALS ,International
Association or Ladakh Studies, Colloquium, Rome, Italy, -11 September 200.
3RSXODUDUWLFOHV
Namgail, 1. ,accepted,. Dainty but scanty: 1ibetan gazelle near extinction in India,
+RUQELOO

Namgail, 1. ,accepted,. Conseration woes in the Indian Changthang. :LOGOLIH
&RQVHUYDWLRQ.

Namgail, 1. ,2006,. 1ibetan sand ox in Ladakh. /DGDJV0HORQJ, Aug. 2006

Namgail, 1. ,2006,. \ildlie at lemis. +RUQELOO, June, 2006

Namgail, 1. ,2004,. Zangskar: mystic land. 6DQFWXDU\$VLD. 24: 44-4.

Namgail, 1. ,2003,. Gya-Miru: last reuge o the 1ibetan argali. 6DQFWXDU\$VLD. 23: 16-
21.

0LVFHOODQHRXVSXEOLFDWLRQV
Namgail, 1. ,2008,. Signs o Climate Change on Roo-top o the world. Mountain lorum
Bulletin, 8 ,2,: 21-22.

Namgail, 1. ,200,. Pastoralism and wildlie conseration: assessing the coexistence o wild and
domestic ungulates or multiple rangeland use in the 1rans-limalaya. Ruord Small Grants
loundation.

Namgail, 1. ,200,. Ladakh`s nightmare rom climate change. 1he Magpie, 16 July 200.
Namgail, 1. ,200,. Changing ace o Ladakh. 1he Magpie, 8 January 200

Namgail, 1. ,2006,. 1rans-limalayan large herbiores: status, conseration and niche
relationships. Report submitted to the \ildlie Conseration Society, Bronx Zoo, New
\ork.

Bhatnagar, \.V., Namgail, 1., Bagchi, S. & Mishra, C. ,2006,. &RQVHUYLQJWKH7LEHWDQ*D]HOOH.
CLRC 1echnical Report No. 13. Nature Conseration loundation., Mysore, India.

Namgail, 1. ,2006,. 1he Great 1ibetan Sheep. 1he Magpie, June 12 2006.

Namgail, 1. ,2006,. In Search o the Great 1ibetan Sheep. 1he Magpie, August 29 2006.

Namgail, 1. ,2006,. 1he Grey Ghost in Danger. 1he Magpie, June 5 2006.

Namgail, 1. ,2006,. 1ibetan gazelle: antelope o the northern plains. 1he Magpie, June 26
2006.
120

Namgail, 1. ,2006,. 1ibetan antelope or Chiru. 1he Magpie, July 31 2006.

Namgail, 1. ,2006,. Ladakh urial: Great Ladakhi Sheep. 1he Magpie, July 1 2006.

Namgail, 1. ,2006,. Asiatic ibex: God`s Own Goat. 1he Magpie, July 3 2006.

Namgail, 1. ,2004,. Lurasian lynx in Ladakh. Cat News, 40: 21-22.

Namgail, 1., lox, J.L. & Bhatnagar, \.V. ,2003,. Argali and liestock! lriends or oes
Interactions between 1ibetan argali and domestic sheep,goats in a prospectie 1rans-
limalyan resere. Report submitted to the International Snow Leopard 1rust.

Namgail, 1. ,2001,. labitat selection and ecological separation between sympatric
1ibetan argali and blue sheep in northern India. M.Phil. thesis, Uniersity o 1rosmo,
Norway.

Namgail. 1. ,1998,. Insect pests o Populus Linn., along with notes on the Biology o
1rabala ishnou, Le. ,Lasiocampiadae: Lepidoptera, & Diacrisia sp. M.Sc. thesis, Panjab
Uniersity, Chandigarh, India

352)(66,21$/$&7,9,7,(6
5HIHUHH Animal Conseration, Oryx, Mammalia, Lnironmental Management, Journal
o Natural listory

0HPEHU International Society o Zoological Sciences, International Association or
Lcology, \orld Association o \oung Scientists, Snow Leopard Network, Mountain
lorum, Indian Crane and \etland \orking Group
121
PL&RC PhD Lducation Certificate

\ith the educational actiities listed below the PhD candidate has
complied with the educational requirements set by the C.1. de \it
Graduate School or Production Lcology and Resource
Conseration ,PL&RC, which comprises o a minimum total o
32 LC1S ,~ 22 weeks o actiities,

Review of Literature (4.2 LC1S)
- lerbiore species diersity at multi-spatial and temporal scales ,2005,

Writing of Project Proposal (7 LC1S)
- lerbiore species richness in the 1rans-limalaya: understanding the causes o spatial
ariation ,2006,

Post-Graduate Courses (4.4 LC1S)
- Community ecology, SLNSL ,2005,
- Adanced statistics, PL&RC ,200,
- Multiariate analysis, PL&RC ,2009,

Deficiency, Refresh, Brush-up Courses (J.7 LC1S)
- Basic statistics, PL&RC ,2006,
- Use o geo-inormation and remote sensing, I1C ,200,
-
Competence Strengthening / Skills Courses (4.2 LC1S)
- Inormation literacy, PL&RC ,2006,
- Scientiic publishing, SLNSL ,200,
- 1echniques or writing and presenting a scientiic paper, SLNSL ,200,
- Project and time management, SLNSL ,200,
- Science, media and general public, SLNSL ,2009,
Discussion Groups / Local Seminars and Other Scientific Meetings (4 LC1S)
- lorest and conseration ecology ,2005-200,
- Spatial methods ,200-2009,
PL&RC Annual Meetings, Seminars and the PL&RC Weekend (J.2 LC1S)
- PL&RC Annual meeting: 1ruth o Science` ,2005,
- PL&RC \eekend ,200,
International Symposia, Workshops and Conferences (JJ.S LC1S)
- 12
th
Benelux Congress o Zoology, \ageningen, the Netherlands ,2005,
- IV \orld Congress on Mountain Ungulates, Munnar, India ,2006,
- 11
th
Meeting o the Goose Specialist Group, Ladakh, India ,2008,
- V \orld Congress on Mountain Ungulates, Granada, Spain ,2009,
- 12
th
Meeting o the Goose Specialist Group, Sweden ,2009,
Supervision of MSc Students (Carola van den 1empel & Mark de Vrij)
- Lcomorphology o mountain ungulates: linking resource selection to skeletal
measurements ,30 days,
122
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8QLYHUVLW\XQGHUWKH6DQGZLFK3K'SURJUDP

Namgail 2009

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Chapter 1. lerbiore species richness: Introduction 1

Chapter 2. Aboeground biomass and phytodiersity along altitudinal 19

lig. 1. A simple model

.H\ZRUGVVascular plants, species richness, aboeground biomass, altitude, Rapoport`s

Species lamily Abbr. in CCA

2UGHUlamily 6FLHQWLILFQDPH &RPPRQQDPH $EEU

~ 23.49, p 0.05, 1able 1,. 1hey also used areas with

Distance to cli ,m,

Species richness and niche dynamics

Co-distribution of urial and blue sheep

Namgail, 1. ,200,. Changing ace o Ladakh. 1he Magpie, 8 January 200

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