Global Environmental Controls of Diversity in Large Herbivores

Nature. Feb, 2002  |  Pubmed ID: 11859367

Large mammalian herbivores occupy half of the earth's land surface and are important both ecologically and economically, but their diversity is threatened by human activities. We investigated how the diversity of large herbivores changes across gradients of global precipitation and soil fertility. Here we show that more plant-available moisture reduces the nutrient content of plants but increases productivity, whereas more plant-available nutrients increase both of these factors. Because larger herbivore species tolerate lower plant nutrient content but require greater plant abundance, the highest potential herbivore diversity should occur in locations with intermediate moisture and high nutrients. These areas are dry enough to yield high quality plants and support smaller herbivores, but productive enough to support larger herbivores. These predictions fit with observed patterns of body size and diversity for large mammalian herbivores in North America, Africa and Australia, and yield a global map of regions with potentially high herbivore diversity. Thus, gradients of precipitation, temperature and soil fertility might explain the global distribution of large herbivore diversity and help to identify crucial areas for conservation and restoration.

Spatial Heterogeneity and Irreversible Vegetation Change in Semiarid Grazing Systems

The American Naturalist. Feb, 2002  |  Pubmed ID: 18707414

Recent theoretical studies have shown that spatial redistribution of surface water may explain the occurrence of patterns of alternating vegetated and degraded patches in semiarid grasslands. These results implied, however, that spatial redistribution processes cannot explain the collapse of production on coarser scales observed in these systems. We present a spatially explicit vegetation model to investigate possible mechanisms explaining irreversible vegetation collapse on coarse spatial scales. The model results indicate that the dynamics of vegetation on coarse scales are determined by the interaction of two spatial feedback processes. Loss of plant cover in a certain area results in increased availability of water in remaining vegetated patches through run-on of surface water, promoting within-patch plant production. Hence, spatial redistribution of surface water creates negative feedback between reduced plant cover and increased plant growth in remaining vegetation. Reduced plant cover, however, results in focusing of herbivore grazing in the remaining vegetation. Hence, redistribution of herbivores creates positive feedback between reduced plant cover and increased losses due to grazing in remaining vegetated patches, leading to collapse of the entire vegetation. This may explain irreversible vegetation shifts in semiarid grasslands on coarse spatial scales.

Self-organization of Vegetation in Arid Ecosystems

The American Naturalist. Oct, 2002  |  Pubmed ID: 18707527

Determinants of Woody Cover in African Savannas

Nature. Dec, 2005  |  Pubmed ID: 16341012

Savannas are globally important ecosystems of great significance to human economies. In these biomes, which are characterized by the co-dominance of trees and grasses, woody cover is a chief determinant of ecosystem properties. The availability of resources (water, nutrients) and disturbance regimes (fire, herbivory) are thought to be important in regulating woody cover, but perceptions differ on which of these are the primary drivers of savanna structure. Here we show, using data from 854 sites across Africa, that maximum woody cover in savannas receiving a mean annual precipitation (MAP) of less than approximately 650 mm is constrained by, and increases linearly with, MAP. These arid and semi-arid savannas may be considered 'stable' systems in which water constrains woody cover and permits grasses to coexist, while fire, herbivory and soil properties interact to reduce woody cover below the MAP-controlled upper bound. Above a MAP of approximately 650 mm, savannas are 'unstable' systems in which MAP is sufficient for woody canopy closure, and disturbances (fire, herbivory) are required for the coexistence of trees and grass. These results provide insights into the nature of African savannas and suggest that future changes in precipitation may considerably affect their distribution and dynamics.

Perceived Conflicts Between Pastoralism and Conservation of the Kiang Equus Kiang in the Ladakh Trans-Himalaya, India

Environmental Management. Dec, 2006  |  Pubmed ID: 16955231

An emerging conflict with Trans-Himalayan pastoral communities in Ladakh's Changthang Plateau threatens the conservation prospects of the kiang (Equus kiang) in India. It is locally believed that Changthang's rangelands are overstocked with kiang, resulting in forage competition with livestock. Here, we provide a review and preliminary data on the causes of this conflict. Erosion of people's tolerance of the kiang can be attributed to factors such as the loss of traditional pastures during an Indo-Chinese war fought in 1962, immigration of refugees from Tibet, doubling of the livestock population in about 20 years, and increasing commercialization of cashmere (pashmina) production. The perception of kiang overstocking appears misplaced, because our range-wide density estimate of 0.24 kiang km(-2) (+/- 0.44, 95% CL) is comparable to kiang densities reported from Tibet. A catastrophic decline during the war and subsequent recovery of the kiang population apparently led to the overstocking perception in Ladakh. In the Hanle Valley, an important area for the kiang, its density was higher (0.56 km(-2)) although even here, we estimated the total forage consumed by kiang to be only 3-4% compared to 96-97% consumed by the large livestock population (78 km(-2)). Our analysis nevertheless suggests that at a localized scale, some herders do face serious forage competition from kiang in key areas such as moist sedge meadows, and thus management strategies also need to be devised at this scale. In-depth socioeconomic surveys are needed to understand the full extent of the conflicts, and herder-centered participatory resolution needs to be facilitated to ensure that a sustainable solution for livelihoods and kiang conservation is achieved.

Concurrent Monitoring of Vessels and Water Turbidity Enhances the Strength of Evidence in Remotely Sensed Dredging Impact Assessment

Water Research. Aug, 2007  |  Pubmed ID: 17583768

Remotely sensed assessment of dredging impacts on water turbidity is straightforward when turbidity plumes show up in clear water. However, it is more complicated in turbid waters as the spatial or temporal changes in turbidity might be of natural origin. The plausibility of attributing turbidity patterns to dredging activities would be greatly enhanced when demonstrating association between dredging infrastructure and water turbidity. This study investigated the possibility to strengthen the inference of dredging impact while simultaneously monitoring vessels and water turbidity in the northern Poyang Lake, China, where dredging was first introduced in 2001 and rapidly extended onwards. Time-series of Landsat TM and MODIS images of 2000-2005 were used to estimate the distribution and number of vessels as well as water turbidity. MODIS images revealed a significant increase in water turbidity from 2001 onwards. Landsat TM image analysis indicated a simultaneous increase in the number of vessels. Regression analysis further showed a highly significant positive relationship (R2=0.92) between water turbidity and vessel number. Visual interpretation of ship locations led to the conclusion that clear upstream waters developed turbidity plumes while passing the first cluster of vessels. We concluded that dredging caused the increase in water turbidity, and simultaneously monitoring the water turbidity and vessels enhanced the strength of evidence in remotely sensed dredging impact assessment.

Selection at the Y Chromosome of the African Buffalo Driven by Rainfall

PloS One. 2007  |  Pubmed ID: 17971851

Selection coefficients at the mammalian Y chromosome typically do not deviate strongly from neutrality. Here we show that strong balancing selection, maintaining intermediate frequencies of DNA sequence variants, acts on the Y chromosome in two populations of African buffalo (Syncerus caffer). Significant correlations exist between sequence variant frequencies and annual rainfall in the years before conception, with five- to eightfold frequency changes over short time periods. Annual rainfall variation drives the balancing of sequence variant frequencies, probably by affecting parental condition. We conclude that sequence variants confer improved male reproductive success after either dry or wet years, making the population composition and dynamics very sensitive to climate change. The mammalian Y chromosome, interacting with ecological processes, may affect male reproductive success much more strongly than previously thought.

Impacts of Savanna Trees on Forage Quality for a Large African Herbivore

Oecologia. Mar, 2008  |  Pubmed ID: 18309522

Recently, cover of large trees in African savannas has rapidly declined due to elephant pressure, frequent fires and charcoal production. The reduction in large trees could have consequences for large herbivores through a change in forage quality. In Tarangire National Park, in Northern Tanzania, we studied the impact of large savanna trees on forage quality for wildebeest by collecting samples of dominant grass species in open grassland and under and around large Acacia tortilis trees. Grasses growing under trees had a much higher forage quality than grasses from the open field indicated by a more favourable leaf/stem ratio and higher protein and lower fibre concentrations. Analysing the grass leaf data with a linear programming model indicated that large savanna trees could be essential for the survival of wildebeest, the dominant herbivore in Tarangire. Due to the high fibre content and low nutrient and protein concentrations of grasses from the open field, maximum fibre intake is reached before nutrient requirements are satisfied. All requirements can only be satisfied by combining forage from open grassland with either forage from under or around tree canopies. Forage quality was also higher around dead trees than in the open field. So forage quality does not reduce immediately after trees die which explains why negative effects of reduced tree numbers probably go initially unnoticed. In conclusion our results suggest that continued destruction of large trees could affect future numbers of large herbivores in African savannas and better protection of large trees is probably necessary to sustain high animal densities in these ecosystems.

Spatio-temporal Dynamics of Global H5N1 Outbreaks Match Bird Migration Patterns

Geospatial Health. Nov, 2009  |  Pubmed ID: 19908191

The global spread of highly pathogenic avian influenza H5N1 in poultry, wild birds and humans, poses a significant pandemic threat and a serious public health risk. An efficient surveillance and disease control system relies on the understanding of the dispersion patterns and spreading mechanisms of the virus. A space-time cluster analysis of H5N1 outbreaks was used to identify spatio-temporal patterns at a global scale and over an extended period of time. Potential mechanisms explaining the spread of the H5N1 virus, and the role of wild birds, were analyzed. Between December 2003 and December 2006, three global epidemic phases of H5N1 influenza were identified. These H5N1 outbreaks showed a clear seasonal pattern, with a high density of outbreaks in winter and early spring (i.e., October to March). In phase I and II only the East Asia Australian flyway was affected. During phase III, the H5N1 viruses started to appear in four other flyways: the Central Asian flyway, the Black Sea Mediterranean flyway, the East Atlantic flyway and the East Africa West Asian flyway. Six disease cluster patterns along these flyways were found to be associated with the seasonal migration of wild birds. The spread of the H5N1 virus, as demonstrated by the space-time clusters, was associated with the patterns of migration of wild birds. Wild birds may therefore play an important role in the spread of H5N1 over long distances. Disease clusters were also detected at sites where wild birds are known to overwinter and at times when migratory birds were present. This leads to the suggestion that wild birds may also be involved in spreading the H5N1 virus over short distances.

Resprouting As a Persistence Strategy of Tropical Forest Trees: Relations with Carbohydrate Storage and Shade Tolerance

Ecology. Sep, 2010  |  Pubmed ID: 20957956

Resprouting is an important persistence strategy for woody species and represents a dominant pathway of regeneration in many plant communities, with potentially large consequences for vegetation dynamics, community composition, and species coexistence. Most of our knowledge of resprouting strategies comes from fire-prone systems, but this cannot be readily applied to other systems where disturbances are less intense. In this study we evaluated sapling responses to stem snapping for 49 moist-forest species and 36 dry-forest species from two Bolivian tropical forests. To this end we compared in a field experiment the survival and height growth of clipped and control saplings for a two-year period, and related this to the shade tolerance, carbohydrate reserves, and the morphological traits (wood density, leaf size) of the species. Nearly all saplings resprouted readily after stem damage, although dry-forest species realized, on average, a better survival and growth after stem damage compared to moist-forest species. Shade-tolerant species were better at resprouting than light-demanding species in moist forest. This resprouting ability is an important prerequisite for successful regeneration in the shaded understory, where saplings frequently suffer damage from falling debris. Survival after stem damage was, surprisingly, only modestly related to stem reserves, and much more strongly related to wood density, possibly because a high wood density enables plants to resist fungi and pathogens and to reduce stem decay. Correlations between sampling performance and functional traits were similar for the two forest types, and for phylogenetically independent contrasts and for cross-species analyses. The consistency of these results suggests that tropical forest species face similar trade-offs in different sites and converge on similar sets of solutions. A high resprouting ability, as well as investments in stem defense and storage reserves, form part of a suite of co-evolved traits that underlies the growth-survival trade-off, and contributes to light gradient partitioning and species coexistence. These links with shade tolerance are important in the moist evergreen forest, which casts a deep, more persistent shade, but tend to diminish in dry deciduous forest where light is a less limiting resource.

Rainfall-driven Sex-ratio Genes in African Buffalo Suggested by Correlations Between Y-chromosomal Haplotype Frequencies and Foetal Sex Ratio

BMC Evolutionary Biology. 2010  |  Pubmed ID: 20416038

The Y-chromosomal diversity in the African buffalo (Syncerus caffer) population of Kruger National Park (KNP) is characterized by rainfall-driven haplotype frequency shifts between year cohorts. Stable Y-chromosomal polymorphism is difficult to reconcile with haplotype frequency variations without assuming frequency-dependent selection or specific interactions in the population dynamics of X- and Y-chromosomal genes, since otherwise the fittest haplotype would inevitably sweep to fixation. Stable Y-chromosomal polymorphism due one of these factors only seems possible when there are Y-chromosomal distorters of an equal sex ratio, which act by negatively affecting X-gametes, or Y-chromosomal suppressors of a female-biased sex ratio. These sex-ratio (SR) genes modify (suppress) gamete transmission in their own favour at a fitness cost, allowing for stable polymorphism.

Personality Predicts the Use of Social Information

Ecology Letters. Jul, 2010  |  Pubmed ID: 20482585

The use of social information is known to affect various important aspects of an individual's ecology, such as foraging, dispersal and space use and is generally assumed to be entirely flexible and context dependent. However, the potential link between personality differences and social information use has received little attention. In this study, we studied whether use of social information was related to personality, using barnacle geese, Branta leucopsis, where boldness is a personality trait known to be consistent over time. We found that the use of social information decreased with increasing boldness score of the individuals. Individuals had lower feeding times when they did not follow the social information and this effect was unrelated to boldness score. When manipulating social information, thereby making it incorrect, individuals irrespective of their boldness score, learned that it was incorrect and ignored it. Our results show that social information use depends on the personality type of an individual, which calls for incorporation of these personality-related differences in studies of spatial distribution of animals in which social information use plays a role.

The Effect of Personality on Social Foraging: Shy Barnacle Geese Scrounge More

Proceedings. Biological Sciences / The Royal Society. Feb, 2010  |  Pubmed ID: 19864281

Animals foraging in groups can either search for food themselves (producing) or search for the food discoveries of other individuals (scrounging). Tactic use in producer-scrounger games is partly flexible but individuals tend to show consistency in tactic use under different conditions suggesting that personality might play a role in tactic use in producer-scrounger games. Here we studied the use of producing and scrounging tactics by bold and shy barnacle geese (Branta leucopsis), where boldness is a personality trait known to be repeatable over time in this species. We defined individuals as bold, shy or intermediate based on two novel object tests. We scored the frequency of finding food patches (the outcome of investing in producing) and joining patches (the outcome of investing in scrounging) by bold and shy individuals and their feeding time. Shy individuals had a higher frequency of joining than bold individuals, demonstrating for the first time that personality is associated with tactic use in a producer-scrounger game. Bold individuals tended to spend more time feeding than shy individuals. Our results highlight the importance of including individual behavioural variation in models of producer-scrounger games.

The Effect of Boldness on Decision-making in Barnacle Geese is Group-size-dependent

Proceedings. Biological Sciences / The Royal Society. Jul, 2011  |  Pubmed ID: 21123271

In group-living species, decisions made by individuals may result in collective behaviours. A central question in understanding collective behaviours is how individual variation in phenotype affects collective behaviours. However, how the personality of individuals affects collective decisions in groups remains poorly understood. Here, we investigated the role of boldness on the decision-making process in different-sized groups of barnacle geese. Naive barnacle geese, differing in boldness score, were introduced in a labyrinth in groups with either one or three informed demonstrators. The demonstrators possessed information about the route through the labyrinth. In pairs, the probability of choosing a route prior to the informed demonstrator increased with increasing boldness score: bolder individuals decided more often for themselves where to go compared with shyer individuals, whereas shyer individuals waited more often for the demonstrators to decide and followed this information. In groups of four individuals, however, there was no effect of boldness on decision-making, suggesting that individual differences were less important with increasing group size. Our experimental results show that personality is important in collective decisions in pairs of barnacle geese, and suggest that bolder individuals have a greater influence over the outcome of decisions in groups.

Increased Searching and Handling Effort in Tall Swards Lead to a Type IV Functional Response in Small Grazing Herbivores

Oecologia. Jul, 2011  |  Pubmed ID: 21221651

Understanding the functional response of species is important in comprehending the species' population dynamics and the functioning of multi-species assemblages. A Type II functional response, where instantaneous intake rate increases asymptotically with sward biomass, is thought to be common in grazers. However, at tall, dense swards, food intake might decline due to mechanical limitations or if animals selectively forage on the most nutritious parts of a sward, leading to a Type IV functional response, especially for smaller herbivores. We tested the predictions that bite mass, cropping time, swallowing time and searching time increase, and bite rate decreases with increasing grass biomass for different-sized Canada geese (Branta canadensis) foraging on grass swards. Bite mass indeed showed an increasing asymptotic relationship with grass biomass. At high biomass, difficulties in handling long leaves and in locating bites were responsible for increasing cropping, swallowing, and searching times. Constant bite mass and decreasing bite rate caused the intake rate to decrease at high sward biomass after reaching an optimum, leading to a Type IV functional response. Grazer body mass affected maximum bite mass and intake rate, but did not change the shape of the functional response. As grass nutrient contents are usually highest in short swards, this Type IV functional response in geese leads to an intake rate that is maximised in these swards. The lower grass biomass at which intake rate was maximised allows resource partitioning between different-sized grazers. We argue that this Type IV functional response is of more importance than previously thought.

Large Herbivores May Alter Vegetation Structure of Semi-arid Savannas Through Soil Nutrient Mediation

Oecologia. Apr, 2011  |  Pubmed ID: 21225433

In savannas, the tree-grass balance is governed by water, nutrients, fire and herbivory, and their interactions. We studied the hypothesis that herbivores indirectly affect vegetation structure by changing the availability of soil nutrients, which, in turn, alters the competition between trees and grasses. Nine abandoned livestock holding-pen areas (kraals), enriched by dung and urine, were contrasted with nearby control sites in a semi-arid savanna. About 40 years after abandonment, kraal sites still showed high soil concentrations of inorganic N, extractable P, K, Ca and Mg compared to controls. Kraals also had a high plant production potential and offered high quality forage. The intense grazing and high herbivore dung and urine deposition rates in kraals fit the accelerated nutrient cycling model described for fertile systems elsewhere. Data of a concurrent experiment also showed that bush-cleared patches resulted in an increase in impala dung deposition, probably because impala preferred open sites to avoid predation. Kraal sites had very low tree densities compared to control sites, thus the high impala dung deposition rates here may be in part driven by the open structure of kraal sites, which may explain the persistence of nutrients in kraals. Experiments indicated that tree seedlings were increasingly constrained when competing with grasses under fertile conditions, which might explain the low tree recruitment observed in kraals. In conclusion, large herbivores may indirectly keep existing nutrient hotspots such as abandoned kraals structurally open by maintaining a high local soil fertility, which, in turn, constrains woody recruitment in a negative feedback loop. The maintenance of nutrient hotspots such as abandoned kraals by herbivores contributes to the structural heterogeneity of nutrient-poor savanna vegetation.

The Spatial Scaling of Habitat Selection by African Elephants

The Journal of Animal Ecology. Jan, 2011  |  Pubmed ID: 21054380

1. Understanding and accurately predicting the spatial patterns of habitat use by organisms is important for ecological research, biodiversity conservation and ecosystem management. However, this understanding is complicated by the effects of spatial scale, because the scale of analysis affects the quantification of species-environment relationships. 2. We therefore assessed the influence of environmental context (i.e. the characteristics of the landscape surrounding a site), varied over a large range of scales (i.e. ambit radii around focal sites), on the analysis and prediction of habitat selection by African elephants in Kruger National Park, South Africa. 3. We focused on the spatial scaling of the elephants' response to their main resources, forage and water, and found that the quantification of habitat selection strongly depended on the scales at which environmental context was considered. Moreover, the inclusion of environmental context at characteristic scales (i.e. those at which habitat selectivity was maximized) increased the predictive capacity of habitat suitability models. 4. The elephants responded to their environment in a scale-dependent and perhaps hierarchical manner, with forage characteristics driving habitat selection at coarse spatial scales, and surface water at fine spatial scales. 5. Furthermore, the elephants exhibited sexual habitat segregation, mainly in relation to vegetation characteristics. Male elephants preferred areas with high tree cover and low herbaceous biomass, whereas this pattern was reversed for female elephants. 6. We show that the spatial distribution of elephants can be better understood and predicted when scale-dependent species-environment relationships are explicitly considered. This demonstrates the importance of considering the influence of spatial scale on the analysis of spatial patterning in ecological phenomena.

Comparing the Community Composition of European and Eastern Chinese Waterbirds and the Influence of Human Factors on the China Waterbird Community

Ambio. Feb, 2011  |  Pubmed ID: 21404825

We compared the European and eastern Chinese waterbird assemblages and checked whether the effects of human disturbance could be detected in the assemblages' composition. For the different Chinese provinces, we expected to find a negative effect of economic development on the mean bird species mass and on the proportion of bentivorous, piscivorous and insectivorous bird species. We also expected to find relatively fewer large species in the Chinese assemblage. Species rank-abundance curves were relatively similar, but China had significantly more species with smaller body masses. The China assemblage was characterized by relatively higher abundance of heavy-bodied species, contrary to our expectations. Mean bird body mass decreased in China with increasing disturbance and increasing gross domestic product (GDP). For coastal provinces in China the percentage of bentivorous, piscivorous and insectivorous bird species declined with increasing GDP, maybe through the increased use of pesticides or fertilizer.

Genome Wide SNP Discovery, Analysis and Evaluation in Mallard (Anas Platyrhynchos)

BMC Genomics. 2011  |  Pubmed ID: 21410945

Next generation sequencing technologies allow to obtain at low cost the genomic sequence information that currently lacks for most economically and ecologically important organisms. For the mallard duck genomic data is limited. The mallard is, besides a species of large agricultural and societal importance, also the focal species when it comes to long distance dispersal of Avian Influenza. For large scale identification of SNPs we performed Illumina sequencing of wild mallard DNA and compared our data with ongoing genome and EST sequencing of domesticated conspecifics. This is the first study of its kind for waterfowl.

Soil Nutrient Status Determines How Elephant Utilize Trees and Shape Environments

The Journal of Animal Ecology. Jul, 2011  |  Pubmed ID: 21418208

1. Elucidation of the mechanism determining the spatial scale of patch selection by herbivores has been complicated by the way in which resource availability at a specific scale is measured and by vigilance behaviour of the herbivores themselves. To reduce these complications, we studied patch selection by an animal with negligible predation risk, the African elephant. 2. We introduce the concept of nutrient load as the product of patch size, number of patches and local patch nutrient concentration. Nutrient load provides a novel spatially explicit expression of the total available nutrients a herbivore can select from. 3. We hypothesized that elephant would select nutrient-rich patches, based on the nutrient load per 2500 m(2) down to the individual plant scale, and that this selection will depend on the nitrogen and phosphorous contents of plants. 4. We predicted that elephant would cause more adverse impact to trees of lower value to them in order to reach plant parts with higher nutrient concentrations such as bark and root. However, elephant should maintain nutrient-rich trees by inducing coppicing of trees through re-utilization of leaves. 5. Elephant patch selection was measured in a homogenous tree species stand by manipulating the spatial distribution of soil nutrients in a large field experiment using NPK fertilizer. 6. Elephant were able to select nutrient-rich patches and utilized Colophospermum mopane trees inside these patches more than outside, at scales ranging from 2500 down to 100 m(2) . 7. Although both nitrogen and phosphorus contents of leaves from C. mopane trees were higher in fertilized and selected patches, patch choice correlated most strongly with nitrogen content. As predicted, stripping of leaves occurred more in nutrient-rich patches, while adverse impact such as uprooting of trees occurred more in nutrient-poor areas. 8. Our results emphasize the necessity of including scale-dependent selectivity in foraging studies and how elephant foraging behaviour can be used as indicators of change in the availability of nutrients.

Biomass and Diversity of Dry Alpine Plant Communities Along Altitudinal Gradients in the Himalayas

Journal of Plant Research. Jan, 2012  |  Pubmed ID: 21638006

A non-linear relationship between phytodiversity and altitude has widely been reported, but the relationship between phytomass and altitude remains little understood. We examined the phytomass and diversity of vascular plants along altitudinal gradients on the dry alpine rangelands of Ladakh, western Himalaya. We used generalized linear and generalized additive models to assess the relationship between these vegetation parameters and altitude. We found a hump-shaped relationship between aboveground phytomass and altitude. We suspect that this is engendered by low rainfall and trampling/excessive grazing at lower slopes by domestic livestock, and low temperature and low nutrient levels at higher slopes. We also found a unimodal relationship between plant species-richness and altitude at a single mountain as well as at the scale of entire Ladakh. The species-richness at the single mountain peaked between 5,000 and 5,200 m, while it peaked between 3,500 and 4,000 m at entire Ladakh level. Perhaps biotic factors such as grazing and precipitation are, respectively, important in generating this pattern at the single mountain and entire Ladakh.