Table Of ContentJiří Doležal · Miroslav Dvorský · Annett Börner
Jan Wild · Fritz Hans Schweingruber
Anatomy, Age
and Ecology of High
Mountain Plants in
Ladakh, the Western
Himalaya
Anatomy, Age and Ecology of High Mountain
Plants in Ladakh, the Western Himalaya
Jiří Doležal • Miroslav Dvorský • Annett Börner
Jan Wild • Fritz Hans Schweingruber
Anatomy, Age and Ecology
of High Mountain Plants
in Ladakh, the Western
Himalaya
123
Ji(cid:284)í Doležal Miroslav Dvorský
Faculty of Science, Department of Botany Institute of Botany
University of South Bohemia The Czech Academy of Sciences
(cid:253)eské Bud(cid:268)jovice, Czech Republic Pr(cid:292)honice, Czech Republic
Annett Börner Jan Wild
Calyptra Pty Ltd Institute of Botany
Adelaide, Australia The Czech Academy of Sciences
Pr(cid:292)honice, Czech Republic
Fritz Hans Schweingruber
Swiss Federal Research Institute WSL
Birmensdorf, Switzerland
ISBN 978-3-319-78697-1 ISBN 978-3-319-78699-5 (eBook)
https://doi.org/10.1007/978-3-319-78699-5
Library of Congress Control Number: 2018947586
© Springer International Publishing AG, part of Springer Nature 2018
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Cover illustration: Haloxylon thomsonii, family of Amaranthaceae. Stem cross-section of a small shrub on rocks in
the desert of northern Ladakh. Single vascular bundles are embedded in aerenchymatic and sclerenchymatic tissues.
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This Springer imprint is published by the registered company Springer International Publishing AG part of Springer
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Contents
Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .1 Nitrariaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 280
Study region . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .3 Onagraceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 281
Climate and soil . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .4 Orobanchaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 283
Phytogeography . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .4 Papaveraceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 284
Species richness and diversity . . . . . . . . . . . . . . . . . . . . . . . . .5 Phrymaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 293
Vegetation and elevational belts. . . . . . . . . . . . . . . . . . . . . . . .5 Plantaginaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 294
The history of botanical exploration . . . . . . . . . . . . . . . . . . . .6 Plumbaginaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 297
Vegetation – basic types and zonation . . . . . . . . . . . . . . . . . .7 Polygonaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 299
Primulaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 320
Anatomical descriptions of horsetails and ferns . . . . . . . . . .35 Ranunculaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 324
Equisetaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .36 Rhamnaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 341
Athyriaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .37 Rosaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 342
Cystopteridaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .39 Rubiaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 359
Dryopteridaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .42 Salicaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 361
Woodsiaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .46 Santalaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 366
Saxifragaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 367
Anatomical descriptions of dicotyledons . . . . . . . . . . . . . . . .47 Scrophulariaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 374
Amaranthaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .48 Solanaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 385
Apiaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .69 Tamaricaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 388
Apocynaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .78 Urticaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 389
Asteraceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .79 Valerianaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 391
Betulaceae. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 141 Violaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 392
Biebersteiniaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 142 Zygophyllaceae. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 393
Boraginaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 143
Brassicaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 155 Anatomical descriptions of monocotyledons . . . . . . . . . . . .395
Campanulaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 183 De(cid:189) nition of culm features . . . . . . . . . . . . . . . . . . . . . . . . . 396
Capparidaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 185 Amaryllidaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 401
Caprifoliaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 186 Cyperaceae. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 403
Caryophyllaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 190 Juncaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 438
Celastraceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 205 Juncaginaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 443
Crassulaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 206 Liliaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 444
Cupressaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 211 Poaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 445
Elaeagnaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 212 Potamogetonaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 553
Ephedraceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 214
Euphorbiaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 217 Synthesis of anatomical, ecological and phylogenetical data . .555
Fabaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 218 Anatomy of dicotyledonous species . . . . . . . . . . . . . . . . . 555
Fumariaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 246 Culm anatomy in monocotyledonous species . . . . . . . . . 586
Gentianaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 247
Geraniaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 259 Material and methods . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .603
Grossulariaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 262
Hippuridaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 263 Acknowledgements . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 607
Lamiaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 264 References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 609
Morinaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 279 Species list . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 613
V
Abbreviations
ae aerenchym f fi ber phg phellogen
bpit bordered pit ft fi ber tracheid pit
ca cambium ge gelatinous fi bers pith
cal callus, parenchymatic cells gr growth ring r ray
clu cell lumen, cell lumina grb growth ring boundary rd resin duct
co cortex he helical thickenings sc sclereid
cork ivp intervessel pit sf septate fi bers
cry crystal la laticifers shc sheet cell
csi collapsed sieve tubes lcw lignifi ed cell wall si sieve tube, sieve element
ct conjunctive tissue lf libriform fi ber spit simple pit
cu cuticula lw latewood ta tannins
di (ray) dilatation lwt latewood tracheid te tension wood
ds dark-stained substances lwv latewood vessel tr tracheid
duct mu mucilage ty tylosis
en endodermis nu nucleus ulcw unlignifi ed cell wall
ep epidermis p perforation v vessel
ew earlywood pa parenchyma vab vascular bundle
ewt earlywood tracheid ph phloem vat vascular tracheid
ewv earlywood vessel phe phellem xy xylem
VII
JI(cid:908)Í DOLEŽAL has been working in the Himalayas for more than ten years and tack-
led the challenge of the (cid:189) eld plant collection together with many students from the
University of South Bohemia in (cid:859)eské Bud(cid:870)jovice. Determining and sorting of the col-
lected plant material from dry and cold Himalayan deserts were necessary steps before
laboratory sectioning and analysis of the rich anatomical structures. Ji(cid:909)í lives in (cid:859)eské
Bud(cid:870)jovice, Czech Republic, with three kids and wife Lucka, whom he thanks for her
constant support.
FRITZ SCHWEINGRUBER sectioning a tiny stem of a herb with a „fossil“ of a mi-
crotome (Reic hert constructed approx. 1940) in his laboratory at the Research Institute
for Forest, Snow and Landscape at Birmensdorf, Switzerland. Anatomical work requires
primarily a steady hand, very sharp microtome blades and a lot of patience. To prepare
the micro-sections for comparative analysis they are stained, dehydrated and preserved
in a resin.
MIROSLAV DVORSKÝ took part in (cid:189) ve Himalayan expeditions and helped collecting
plant samples and vegetation data as a PhD student. He soon became increasingly keen
on taking photographs of all plant species encountered, and this was not long before the
idea of a (cid:189) eld guide to the local (cid:190) ora was conceived. Plant species identi(cid:189) cation in the
absence of any standard determination key was often a tricky part of the (cid:189) eldwork, and
required gathering of herbarium specimens and cooperation with taxonomy experts.
JAN WILD’s career is already more than twenty years related to geoinformatics or
broadly speaking to spatial data. Production of pretty maps is thus his hobby as well as a
way of communicating a scienti(cid:189) c work to the public. Jan lives with his wife Hanka and
two children on the southern border of Prague, Czech Republic, close to Pr(cid:925)honice,
where is the headquarter of the Institute of Botany, his long-term employer.
ANNETT BÖRNER has been working in the (cid:189) eld of scienti(cid:189) c publishing for more than
(cid:189) fteen years and took up the challenge of the book’s design and editorial process. Pre-
senting the material in a clear and structured way demands a high standard of technical
and botanical knowledge. Annett lives in Adelaide, Australia, and when she gets tired of
looking at plants on a computer screen she loves going on (cid:189) eld trips and looking at them
in real life.
IX
Tso Moriri Lake is at an
altitude of 4522 m a.s.l.
It is the largest of the
high altitude lakes
entirely within India.
INTRODUCTION
Despite widespread interest in alpine ecology, we have little “Leave the forest, leave the temperate climate and explore the
information on anatomy of high-mountain plants. Most studies anatomy of high-mountain plants in relation to their morphological
are based on data from European mountains (Schweingruber et and ecophysiological adaptations to frost and to drought in cold
al. 2011; Schweingruber and Büntgen 2013), while the largest and desert ecosystems” are the basic ideas of the present book. The
highest mountainous areas are located in Asia. On the Tibetan book introduces plants of very broad taxonomic spectrum and
Plateau and the neighboring Himalayan crests, which cover an distribution, ecological habitat preferences, morphology, longevity
area almost 13 times larger than the area of the European Alps, and evolutionary history, all growing naturally in Ladakh. Xylem
extensive areas of alpine climate and vegetation host a unique and and bark structures of selected representatives are described
species-rich (cid:190) ora (Miehe et al. 2011) and serve as grazing land for for Equisetophyta (1 species), Pteridophyta (10), Gnetophyta
yaks, wild sheep and other mammals, while many plant species are (3), Pinophyta (1), monocotyledons (155) and dicotyledons (345
utilised by local people for medicinal purposes. No comprehensive species) belonging to 63 plant families and 236 genera. A total
monography concerning both ecology and anatomy of alpine of about 1250 species of vascular plants are known to occur
plants is available from these areas. However, about 70% of all in the wild in Ladakh. The number seems to be surprisingly low
glaciers, representing the only reliable water reservoir for local as compared to, for example, the Czech Republic, which has a
people and their livelihoods, are presently retreating (Xu et al. slightly smaller surface area, but about four times more plant
2009; Schmidt and Nüsser 2017), indicating that a considerable species. While the low species richness obviously relates to a
environmental change in this high altitude region is underway. generally adverse, cold, arid, high altitude environment, the (cid:190) ora
encountered in Ladakh is still speciose, phylogenetically diverse
The general lack of studies on anatomy of high-mountain plants and ecologically impressive.
may re(cid:190) ect working habits of biologists. While ecologists tend to
use outdoors as a laboratory, for the plant anatomist, laboratory This book illustrates and describes more than 500 species of
work is done indoors, and the habit of working both in the (cid:189) eld and Ladakh’s vascular (cid:190) ora. We have tried to select a set of common,
in the laboratory apparently does not come easily to most plant widespread species as well as some rarer ones, which are
anatomists, who commonly do comparative work on material they considered endemic, phytogeographically interesting or potentially
seldom collect. High-mountain areas in the Himalayas and Tibet endangered. All described plants grow in high, largely treeless
are often dif(cid:189) cult to access, and so plant anatomists are less likely altitudes in a cold arid region with short vegetation periods. Most
to visit these areas. Perhaps more signi(cid:189) cantly, plant anatomists species are shown through photos in their natural habitats in
tend not to be interested in plant ecology and physiology (and Ladakh. Where appropriate photos were not available (mostly of
vice versa for plant ecologists), such that those kinds of data grasses and sedges) we provided images of scanned herbarium
needed for structure/function correlations are often unavailable specimens. Annotated for each species are the accepted Latin
in tandem. Collaboration among researchers from different (cid:189) elds name and its synonyms, key identi(cid:189) cation features, geographical
seems a desirable way of proceeding with studies on anatomy, distribution and habitat. The range of each species in Ladakh is
ecology and evolutionary adaptations of high-mountain plants. shown as a distribution map, and its frequency along the altitudinal
The present book is a result of such collaboration among plant gradient is depicted as a diagram.
ecologists, anatomist and graphic and GIS experts in the high and
sacred realm of Ladakh, located in a mountain landscape in the Anatomical descriptions are based on root collars (transition
Transhimalaya of Northwest India. between roots and shoots) or rhizomes of several individuals
© Springer International Publishing AG, part of Springer Nature 2018
J. Doležal et al., Anatomy, Age and Ecology of High Mountain Plants in Ladakh, the Western Himalaya, https://doi.org/10.1007/978-3-319-78699-5_1 1
collected for each species in the (cid:189) eld. These parts of plants identical genotype to develop differently — based on speci(cid:189)c
provide the maximum number of annual rings (Schweingruber conditions during their ontogeny (Sultan 2000), and evolutionary
and Poschlod 2005). Most of the material was preserved in constraints (phylogenetic inertia) in which taxa that share part
40% ethanol and sectioned without embedding in transverse, of their evolutionary history possess similar ‘blue-prints’ (Ackerly
tangential and radial directions with a sliding microtome, stained 2000). The present book compares variation in more than 120
with Astrablue/Safranin and preserved in Canada Balsam. anatomical features of Dicots and 42 features of Monocots
Herbarium material was used for all Monocots and a few annual among (1) dominant habitat types in Ladakh such as cold
Amaranthaceae. Preparation methods are described in detail by deserts, alpine steppes, saline wetlands, alpine grasslands, ruderal
Gärtner and Schweingruber (2013). vegetation, alpine screes and subnival zones, (2) with respect to
species optima on the broad elevation gradient in Ladakh from
Anatomical features of xylem of dicotyledonous trees and 2650 to 6150 m, (3) among different growth and life forms such
shrubs are principally de(cid:189) ned by the IAWA List (Wheeler et al. as short-lived plants, perennial plants with main tap roots, various
1989). After sectioning and inspecting herbaceous species from clonal rhizomatous plants, compact alpine cushion plants and rare
non-forest regions the number of features in the xylem had to arborescent species, (4) and among plant height classes. Patterns
be expanded and bark features newly de(cid:189) ned (Crivellaro and of species altitudinal and geographical distributions were derived
Schweingruber 2016). The given anatomical feature codes in from more than 122,000 records of occurrence of vascular plant
this book are based on Schweingruber et al. (2011). Anatomical species in Ladakh collected by Dr. Leoš Klimeš in the period
characteristics of Monocots are mostly based on Metcalfe (1960) 1997–2007.
and Schweingruber and Berger (2017).
The present book has four major goals: a) Describing the stem-
The detailed knowledge of plant anatomical characters and anatomical structures of vascular plants from Ladakh, b) Providing
their variation among different species and habitats is key to basic information about species morphology, geographical and
understanding plant evolution and function (Niklas 1985). elevational distribution and habitat preferences, c) Constructing
Variation in anatomical structure is a result of several forces such a phylogenetic tree for target species from nucleotide sequences,
as: the adaptation of species to the prevailing conditions in their d) Relating stem anatomical features to phylogeny, ecology,
habitats (Schweingruber et al. 2014, Schweingruber and Börner morphology and ecophysiology.
2018), phenotypic plasticity as an ability of individuals with an
2
Study region
Ladakh is a mountainous region located in NW India (Jammu & Geomorphologically, western and southern parts of Ladakh
Kashmir State), ranging between 2600 and 7672 m a.s.l. Its nearly differ considerably from Eastern Ladakh. They are dominated by
87,000 km2 include a considerable part of the transitional zone deep, narrow valleys, and glaciated peaks. Precipitation is much
between outer Himalayan ranges and Western Tibet, forming a higher, especially at higher elevations, and the snow line is situated
naturally delimited area of Transhimalaya. The interface between by 1000 m lower than in the east. Consequently, the (cid:190) ora of
the humid outer Himalaya and dry cold plains of central and Western Ladakh differs considerably, and true Himalayan species
western Tibet is generally quite sharp. For example, in Nepal, are more prominent. Eastern Ladakh represents an area which is
this transitional zone is usually not more than a few kilometers well delimited climatically, biogeographically and culturally, and its
broad. In such conditions (cid:190) ora of the transitional zone is not (cid:190) ora is mostly related to that of the Tibetan Plateau. Northern
particularly interesting, as it is formed by depauperate (cid:190) oras of Ladakh, encompassing a part of Eastern Karakorum Range, has
outer Himalayas and Tibet. Only in the Western Himalayas (and dry climate and rugged terrain, and the highest summits of Ladakh
to some extent in the easternmost Himalayas) the transitional are located there.
zone is broad enough to host a speci(cid:189) c (cid:190) ora, relatively rich in
species restricted to this zone. In spite of the high elevation
and severity of the environment, local (cid:190) ora shows a remarkable
phylogenetical and ecological diversity. Nevertheless, in terms
of species richness per area the whole region belongs among
biodiversity coldspots. Plants in Ladakh are generally subjected
to multiple stresses, such as low precipitation, extreme diurnal
temperature (cid:190) uctuation, strong winds, soli(cid:190) uction at the higher
and salinity at lower elevations (Stewart 1916–1917).
(cid:26)(cid:25)(cid:131)(cid:40) (cid:26)(cid:26)(cid:131)(cid:40) (cid:26)(cid:27)(cid:131)(cid:40) (cid:26)(cid:28)(cid:131)(cid:40) (cid:27)(cid:19)(cid:131)(cid:40)
(cid:42)(cid:85)(cid:82)(cid:90)(cid:76)(cid:81)(cid:74)(cid:3)(cid:86)(cid:72)(cid:68)(cid:86)(cid:82)(cid:81)(cid:3)(cid:70)(cid:79)(cid:76)(cid:80)(cid:68)(cid:87)(cid:72)
(cid:22)(cid:23)(cid:131)(cid:49)(cid:22)(cid:22)(cid:131)(cid:49) !!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!(cid:40)!!!!!!!!!!!!!!!!!!!!!(cid:79)!!!!!!!!!(cid:72)!!!!!!!!!!!!!!!!!!(cid:89)!!!!!!!(cid:22)(cid:23)(cid:24)(cid:25)(cid:68)!!!!(cid:19)(cid:19)(cid:19)(cid:19)(cid:87)!!!!!!!!!!(cid:76)!!!!!!!!(cid:19)(cid:19)(cid:19)(cid:19)!!!(cid:82)!!!!!!!!!!!!!(cid:19)(cid:19)(cid:19)(cid:19)!!!(cid:81)!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!G!!!!!!!!!!!!!!!!!!!!!")(cid:46)!!!!!!!!!!!!!!!!!!!!!!r!!!!!!(cid:68)!!!!e!!!!!!!!!!(cid:85)!!!!!!(cid:74)!a!!!!!!!!!!(cid:76)!!!!!!!!t!!(cid:79)!!!!!!!!!!!!!!!!!!!!!!!!!!!H!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!i!!!!!!!!!m!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!a!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!l!!!!!!!!!!!!!!!!!!!!!!!!!a!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!y!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!a!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!R!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!a!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!n!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!g!!!!!!!!!!!!!!(cid:51)!!!!!!!!e!")!!!!!!!!!(cid:68)!!!!!!!!!!!!(cid:71)!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!(cid:88)!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!(cid:80)!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!(cid:39)!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!(cid:76)!!!!!!!!!!!!!!!!!!!!!!!!!!!!(cid:86)!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!(cid:78)!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!(cid:76)!!!!!!!!!!!!!!!!!!!!(cid:87)!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!")!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!")!!!!!!!!(cid:54)!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!")!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!(cid:68)!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!(cid:85)!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!(cid:70)!!!!!!!!!!!!!!!!!!!!!!!!!(cid:47)!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!(cid:75)!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!(cid:72)!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!(cid:88)!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!(cid:75)!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!")!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!(cid:56)!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!(cid:83)!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!(cid:86)!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!(cid:75)!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!(cid:76)!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!(cid:47)!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!(cid:374)!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!")!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!(cid:282)!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!(cid:48)!!!!!(cid:349)!!!!!!!!!(cid:258)!!!!!!!!!!!!!!(cid:68)!!!!!!(cid:18)!!(cid:75)!!!!(cid:346)(cid:72)!(cid:349)(cid:374)(cid:258) (cid:48)(cid:72)(cid:68)(cid:81)(cid:3)(cid:68)(cid:76)(cid:85)(cid:3)(cid:87)(cid:72)(cid:80)(cid:83)(cid:72)(cid:85)(cid:68)(cid:87)(cid:88)(cid:85)(cid:72)(cid:3)(cid:11)(cid:131)(cid:38)(cid:12)(cid:48)(cid:72)(cid:68)(cid:81)(cid:3)(cid:86)(cid:82)(cid:76)(cid:79)(cid:3)(cid:80)(cid:82)(cid:76)(cid:86)(cid:87)(cid:88)(cid:85)(cid:72)(cid:3)(cid:11)(cid:8)(cid:12)(cid:20)(cid:20)(cid:20)(cid:20)(cid:19)(cid:24)(cid:19)(cid:24)(cid:19)(cid:24)(cid:19)(cid:24)(cid:22)(cid:22)(cid:19)(cid:19)(cid:19)(cid:19)(cid:19)(cid:19) (cid:23)(cid:23)(cid:19)(cid:19)(cid:19)(cid:19)(cid:19)(cid:19) (cid:24)(cid:24)(cid:19)(cid:19)(cid:19)(cid:19)(cid:19)(cid:19) (cid:25)(cid:25)(cid:19)(cid:19)(cid:19)(cid:19)(cid:19)(cid:19) (cid:22)(cid:23)(cid:131)(cid:49)(cid:22)(cid:22)(cid:131)(cid:49)
(cid:19) (cid:21)(cid:24) (cid:24)(cid:19) (cid:26)(cid:24) (cid:20)(cid:19)(cid:19)
(cid:11)(cid:80)(cid:3)(cid:68)(cid:17)(cid:86)(cid:17)(cid:79)(cid:17)(cid:12) (cid:78)(cid:80) (cid:40)(cid:79)(cid:72)(cid:89)(cid:68)(cid:87)(cid:76)(cid:82)(cid:81)(cid:3)(cid:11)(cid:80)(cid:3)(cid:68)(cid:17)(cid:86)(cid:17)(cid:79)(cid:17)(cid:12)
(cid:26)(cid:25)(cid:131)(cid:40) (cid:26)(cid:26)(cid:131)(cid:40) (cid:26)(cid:27)(cid:131)(cid:40) (cid:26)(cid:28)(cid:131)(cid:40) (cid:27)(cid:19)(cid:131)(cid:40)
Distribution of vegetation plots and in situ measured climate in the elevation (Pearson’s r = -0.96), while mean soil moisture increases
study region in Ladakh, NW Himalayas. Floristic data were collected (Pearson’s r = 0.49; graphs on the right side). The trend lines were
in 4,150 plots (100×100 m, black dots) and climate measured at 36 (cid:189) tted with GAM and shaded regions represent 95% con(cid:189) dence in-
sites (red dots) between 2013 and 2014. Histogram on the left side tervals. Background map is based on SRTM (http://www2.jpl.nasa.gov/
shows the frequency of vegetation plots within respective elevational srtm/) and OpenStreetMap data.
bands. Mean air temperature of the growing season decreases with
3
