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Understanding habitat selection of the Vulnerable wild mutus on the

X UCHANG L IANG,AILI K ANG and N ATHALIE P ETTORELLI

Abstract We tested a series of hypotheses on drivers of change is also expected to affect the long-term availability habitat selection by the Vulnerable Bos mutus, of suitable habitats for the species (Schaller, ), although combining distribution-wide sighting data with species dis- there is little quantified and spatially explicit information tribution modelling approaches. The results indicate that available to inform discussions on potential management op- climatic conditions are of paramount importance in shaping tions. More broadly, quantitative information on the factors the wild yak’s distribution on the Tibetan Plateau. Habitat driving patterns in the seasonal distribution of wild is selection patterns were seasonal, with yaks appearing to se- still rare. Studies on Tibetan herbivore species rarely include lect areas closer to villages during the vegetation-growing data from the species’ entire distribution range (Sharma season. Unexpectedly, our index of forage quantity had a et al., ;Singhetal.,; St-Louis & Côté, ), which limited effect in determining the distribution of the species. prevents the identification of environmental management Overall, our results suggest that expected changes in climate actions to alleviate further pressures on populations at the for this region could have a significant impact on habitat necessary scale for the conservation of large species. availability for wild yaks, and we call for more attention to We aim to fill this knowledge gap by combining advances be focused on the unique wildlife in this ecosystem. in species distribution modelling with sighting data col- lected across most of the known distribution range of the Keywords Climate change, generalized additive model, wild yak. We predicted that () the species would exhibit highland, large herbivore, MaxEnt, random forest, seasonal distinct habitat selection patterns between seasons, which habitats, species distribution model has been suggested previously but has not been assessed To view supplementary material for this article, please visit in a quantitative manner (Harris & Miller, ; Schaller, http://dx.doi.org/./S ); in particular, we expected preferred habitats of wild yaks during the vegetation growing season to be found at higher altitudes, in more rugged terrain, and closer to gla- ciers (Schaller, ); () the species would select for quan- Introduction tity over quality of forage at the distribution-range scale, he wild yak Bos mutus is a rare yet iconic large herbi- given that wild yaks are non-selective grazers (Jarman,   Tvore inhabiting the Tibetan Plateau. One of the largest ); and ( ) predation risk, here captured by anthropogenic bovids, wild yaks are also the largest native species in their disturbances given the general lack of natural predators of  range, which formerly included China (Gansu, Sichuan, wild yaks in the area (Schaller, ), would be a significant , , Qinghai), northern India (), and factor shaping habitat selection patterns, and wild yaks (Schaller & Liu, ). The species declined in the would avoid areas near human communities (Leslie &  th century, mainly as a result of excessive hunting; the Schaller, ). The knowledge derived from this study total number of mature individuals was last estimated to will be used to predict seasonal habitat availability in the be c. ,,in (Schaller, ). The species is categor- context of climate change, which will help to highlight fu- ized as Vulnerable on the IUCN Red List (Harris & Leslie, ture global conservation challenges on the Tibetan Plateau. ) and most of the remaining individuals are found in isolated and fragmented populations in the central and nor- thern parts of the Tibetan Plateau. Remnant populations Study area face escalating threats from anthropogenic activities, such  as increasing competition with for good grazing The study area (Fig. ) covers c. . million km on the areas, and infrastructural development that causes degrad- Tibetan Plateau. It encompasses the entire Tibet Interior re- ation of their habitats (Leslie & Schaller, ). Climate gion, from Kunlun in the north to the Gangdise and Nyainqentanglha ranges in the south, with a slight eastward extension to incorporate part of the Sanjiangyuan region in XUCHANG LIANG and AILI KANG Wildlife Conservation Society, Beijing, China the Qinghai province of China. This region includes most of NATHALIE PETTORELLI (Corresponding author) Zoological Society of London, ’ the known current distribution range of the wild yak (Leslie Institute of Zoology, Regent s Park, London NW1 4RY, UK  E-mail [email protected] & Schaller, ). Mean annual precipitation follows a de- Received  July . Revision requested  August . creasing gradient from east to west and from south to north, Accepted  September . First published online  March . ranging from c.  mm in the south-east to ,  mm in the

Oryx, 2017, 51(2), 361–369 © 2016 Fauna & Flora International doi:10.1017/S0030605315001155 Downloaded from https://www.cambridge.org/core. IP address: 170.106.33.19, on 25 Sep 2021 at 14:53:02, subject to the Cambridge Core terms of use, available at https://www.cambridge.org/core/terms. https://doi.org/10.1017/S0030605315001155 362 X. Liang et al.

 FIG. 1 Distribution of the wild yak Bos mutus on the Tibetan Plateau, China. The study area covers c. . million km , encompassing the entire Tibet Interior region defined by the Kunlun mountains in the north and the Gangdise and Nyainqentanglha ranges in the south, with slight eastward extension to incorporate part of the Sanjiangyuan region in Qinghai Province.

north-west. Mean annual temperatures vary from  to ). The spatial resolution of the environmental variables  −°C, with winter extremes of , −°C. The Tibetan considered was set to  km . All the candidate variable layers Steppe is the main ecoregion in the study area. Vegetation were cropped to the extent of the study area and, if necessary,  is sparsely distributed in alpine meadows, alpine steppes, resampled to a  km spatial resolution using the nearest semi-arid steppes and cold deserts (Schaller, ). neighbour method in the raster library (Hijmans & van Etten, )inRv... (R Development Core Team, ). Methods  Presence data Presence data were collected by the Wildlife Climate The Bioclim variables (representative of the – Conservation Society (WCS) and partners in , , years ) from the WorldClim dataset (version    , ,  and . Most of the surveys were . ; Hijmans et al., ) were used to capture current conducted within areas known to hold wild yaks; however, climatic conditions in the study area. To predict future the surveys were not primarily designed to collect trends in habitat availability we downloaded the Bioclim  information on wild yaks, and sightings were layers for under two representative concentration   opportunistic. Sightings were georeferenced by trained pathways, RCP and RCP , which describe possible staff, following the field protocol established by WCS climate futures, depending on various trajectories of China. Ancillary data (e.g. whether sighting data were greenhouse gas concentration. The climate projections  collected while in vehicle or on foot, number of observers, used in this study were derived from the HadGEM -ES survey effort) were not systematically collected and climate model, an updated version of the HADGEM therefore could not be taken into account in subsequent model, which has been reported to adequately predict the  analyses. Vehicle surveys were not based on existing road Tibetan climate (Hao et al., ). systems but survey effort was shaped by the local topography as well as the distribution of seasonal rivers. Topography Topography is known to cause variation in the While conducting surveys the speed of the vehicle was quantity and quality of forage available for large herbivores, required to be ,  km per hour to avoid disturbing and to shape local predation risk (Brown, ;Illius& wildlife. The total number of independent records of wild O’Connor, ). The topographic ruggedness index, a yaks was ;  of these sightings were recorded during measurement developed by Riley et al. () to quantify the non-growing season (October–March; Yu et al., ) the total altitudinal change across a given area, was and the remainder (n = ) were during the vegetation calculated based on the downloaded digital elevation model growing season (April–September). layer GTOPO from the U.S. Geological Survey’sLong Term Archive (USGS, ). Calculations were performed Environmental variables We adopted a methodological in QGIS v. .. (Quantum GIS Development Team, ). framework that categorized relevant environmental variables according to limiting factors (i.e. climatic and topographical factors), disturbance (i.e. anthropogenic Anthropogenic influence Although natural predators of influence) and resource distribution (i.e. availability of wild yaks exist on the Tibetan Plateau (e.g. Schaller, ; forage and fresh water) (Guisan & Thuiller, ; Austin, Xu et al., ; Leslie & Schaller, ), predation risk for

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the species is considered to be shaped primarily by human To overcome uncertainties linked to the choice of species presence and activity (Leslie & Schaller, ). The distribution model we used three analytical approaches distribution of human communities within the study area that have been widely employed in species distribution is relatively dense south of °N, and sparse to the modelling exercises, namely, generalized additive models north. Livestock rearing is the predominant livelihood. (Yee & Mitchell, ), maximum entropy (MaxEnt; Elith Long-distance nomadism is now rare and pastoral activities et al., ) and random forests (Breiman, ). All models normally take place in designated grazing areas near villages were developed in R using the package biomod v. .– (Sheehy et al., ). The linear distance between the centre (Thuiller et al., ). of any given pixel and the nearest village was used as a Firstly, we explored the importance of the climatic and proxy for anthropogenic disturbance and was calculated for topographical variables in shaping the current distribution all pixels. Calculations were performed in QGIS using the of the wild yak. Because we expected habitat selection to Proximity function. The shapefile detailing the distribution be seasonal, models were run  times each for the growing of villages in the area was provided by WCS China. (G_I) and non-growing (NG_I) seasons (Table ). Secondly, we considered current yak distribution as a function of climat- ic conditions, topographical factors, availability of forage, gla- Availability of fresh water Glaciers have important effects cier distribution and anthropogenic influences. Again, these on the hydrological cycle of high-altitude regions models were run for the growing (G_II) and non-growing  (Nogués-Bravo et al., ). The melting ice and snowpack (NG_II) seasons (Table ). Multicollinearity was checked provide seasonal fresh water and soil moisture critical to using the variance inflation factor analysis (R library usdm;  local vegetation communities (Schaller, ). The linear O’Brien, ). We excluded some candidate variables to distance between the centre of any given pixel and the mitigate the effects of inflation caused by the high correlations nearest glacier was therefore estimated for all pixels, using amongst the predictor variables (Dormann et al., ). the Proximity function in QGIS.Theshapefileofglacier Yak presence data were split independently into % for distribution was acquired from the GLIMS Glacier Database training and % for testing (Araújo et al., ). Ten thou-  (GLIMS & National Snow and Ice Data Center, ). sand background points (representing pseudo-absence for generalized additive models) were selected at random Forage The normalized difference vegetation index throughout the study area. The generalized additive model (NDVI), one of the most intensely studied and widely was set with four degrees of freedom for smoothing (Austin,  used vegetation indices (Pettorelli, ), was considered a ). When performing MaxEnt, species prevalence was    proxy for forage availability. MODIS Terra NDVI products set to . (Elith et al., ). The maximum decision tree  (MODA, monthly data for –) were downloaded of the random forests model was set to (Cutler et al.,  using the MODIS Reprojection Tool Web Interface ). We assessed model performance using three evalu-  (USGS, ). As the reflected light waves captured by ation methods: Kappa (Cohen, ), the true skill statistic  satellite sensors can be influenced by a variety of natural (TSS; Allouche et al., ) and area under the curve (AUC;  phenomena (Achard & Estreguil, ), the downloaded Swets, ). Model predictions were categorized as excel- .    .   data layers were processed in R to convert all negative values lent for Kappa . (Fleiss, ), TSS . (Thuiller  .    to zeros and adjust the anomalous values, which were et al., ) or AUC . (Swets, ). assumed to reflect atmospheric noise in the MODA Predictions of species presence probability from the best- – dataset (see Garonna et al., , for full methdology). performing model were converted to presence absence pre- dictions using a transforming threshold selected as the one that maximized TSS scores (Allouche et al., ; Lobo et al.,  Modelling approach ). Variable importance was estimated using a variable permutation algorithm (Breiman, ). Information on alti- Species distribution models are numerical tools to assist tude, terrain ruggedness, and distance to the nearest village in quantifying species–environment relationships. and glacier was extracted from all predicted presence pixels Increasingly, they are used to gain ecological insights and for both seasons under the best model of current habitat suit- predict species distributions at large spatial scales (Guisan ability distribution for wild yaks; these values were then com- & Zimmermann, ). There are various types of species pared between seasons using Wilcoxon one-tailed sum rank distribution models that can be used in combination with tests (Hollander & Wolfe, ). presence data to assess habitat suitability; the predictive power of a given modelling approach may be context- Results specific and may vary depending on the study area, vari- ables and resolution considered, as well as the amount of Random forests models generally outperformed general presence data available (Guisan & Zimmermann, ). additive and MaxEnt models (Table ). In accordance

Oryx, 2017, 51(2), 361–369 © 2016 Fauna & Flora International doi:10.1017/S0030605315001155 Downloaded from https://www.cambridge.org/core. IP address: 170.106.33.19, on 25 Sep 2021 at 14:53:02, subject to the Cambridge Core terms of use, available at https://www.cambridge.org/core/terms. https://doi.org/10.1017/S0030605315001155 364 X. Liang et al.

TABLE 1 Predictor variables used in this study, with definition, group, and range of values.

Variable Definition (unit) Group* Range (mean) Alt Altitude (m) G_I NG_I 242–7,423 (4775) G_II NG_II TRI Topographic ruggedness index G_I NG_I 0–1,080 (76) (m) G_II NG_II Bio3 Isothermality (mean diurnal G_I NG_I 28–46 (38) range divided by annual tem- G_II NG_II perature range *100) Bio15 Precipitation seasonality (coeffi- G_I NG_I 35–154 (105) cient of variation*100) G_II NG_II Bio8 Mean temperature of wettest G_I −84–283 (65) quarter (°C * 10) G_II Bio13 Precipitation of wettest month G_I 6–618 (69) (mm) G_II Bio11 Mean temperature of coldest NG_I −282–160 (−136) quarter (°C * 10) NG_II Bio14 Precipitation of driest month NG_I 0–38 (1.7) (mm) NG_II V_distance Distance to nearest village (km) G_II NG_II 0–412 (57) G_distance Distance to nearest glacier (km) G_II NG_II 0–259 (53) Change_AM Changes in NDVI values between G_II −1,503–3,975 (100) April & May Change_MJ Changes in NDVI values between G_II −2,122–5,164 (442) May & June (* 10,000) Change_JA Changes in NDVI values between G_II −3,156–2,549 (95) July and August (* 10,000) Change_AS Changes in NDVI values between G_II −3,232–3,835 (−367) August & September (* 10,000) Ave_allmon Averaged NDVI values across G_II 0–8,521 (1,237) years (* 10,000)

*The groups G_I (growing season) and NG_I (non-growing season) included only topographical and climatic variables; G_II and NG_II also included variables capturing information on anthropogenic influence, glacier distribution, and forage availability.

TABLE 2 Performance of random forests, general additive, and MaxEnt models in terms of area under the curve (AUC), true skill statistic (TSS) and Kappa, for the growing and non-growing seasons. Each model was run  times for each season, and model performance was evaluated independently for each run.

Growing season Non-growing season Mean ± SD Mean ± SD AUC TSS Kappa AUC TSS Kappa Random forests 0.985 ± 0.01 0.91 ± 0.04 0.87 ± 0.03 0.95 ± 0.007 0.77 ± 0.02 0.62 ± 0.03 Generalized additive model 0.98 ± 0.007 0.90 ± 0.02 0.68 ± 0.04 0.92 ± 0.005 0.73 ± 0.01 0.39 ± 0.02 MaxEnt 0.97 ± 0.01 0.82 ± 0.03 0.63 ± 0.05 0.92 ± 0.006 0.74 ± 0.02 0.38 ± 0.02

with our first prediction, wild yaks showed distinct seasonal temperatures were more likely to be preferred (Fig. ). patterns of habitat selection; climatic conditions were strong Preferred habitats during the growing season were found at determinants of these patterns at the spatial scale considered higher altitudes (W = ,P, .), closer to gla- (Fig. ). During the growing season wild yaks appeared to ciers (W = ,P, .) and in more rugged terrain select areas with low levels of fluctuations in monthly pre- (W = ,P, .) than those used during the cipitation; they also appeared to favour areas with relatively non-growing season (Supplementary Table S). Contrary to abundant precipitation in the peak summer month (i.e. July). our third hypothesis, however, wild yaks tended to be During the non-growing season drier areas with greater fluc- found closer to villages during the growing season than dur- tuation in monthly precipitation and less extreme winter ing the non-growing season (W = ,P, .). All

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Plateau, showing that the species’ habitat selection patterns are seasonally distinct and largely driven by climatic factors. Yet two of our predictions were not well supported by our findings. The first pertains to the importance of forage quantity in driving habitat selection. Wild yaks are non- selective grazers (Schaller, ) and are therefore not ex- pected to select forage on the basis of quality over quantity (Jarman, ). Although we expected forage biomass to be a key factor in determining wild yak occurrence, our results show that most variables based on NDVI play little if any role in shaping wild yak distribution. Unlike previously re- ported cases where NDVI could be linked to the distribution of large herbivores (see Pettorelli, , for a review), vari- ables based on this index may not have captured vegetation dynamics correctly in our study area as a result of high soil reflectance (Pettorelli et al., ). However, our results may also suggest that wild yaks select for quality over quantity of forage to an extent beyond our initial expectation. The nu- tritious -dominant moist meadows, favoured by wild yaks in summer according to empirical observations (Harris & Miller, ), are not as productive in terms of vegetation biomass as other vegetation types, such as grasslands, which are more widely distributed in our study FIG. 2 The importance of individual variables (Table )in area (Schaller, ). Pixels with higher NDVI values would predicting the distribution of wild yaks under the best model, for thus fail to capture the distribution of these favoured, yet less (a) the vegetation growing season and (b) the non-growing productive, meadows. The low level of fluctuation in  season. The best model was run times for each season, and monthly precipitation, and the abundant precipitation in variable importance was evaluated independently for each run. July (the two conditions identified as being key to capturing wild yak distribution during the growing period) are also the variables based on NDVI were comparatively of much key factors determining the biomass and nutrient value of lower importance in defining habitat selection patterns Kobresia-dominant moist meadows (Yu et al., ).  than the top climatic variables (Fig. ). These meadows are associated with high levels of vapour  Based on these results it is likely that under the RCP loss (Körner, ) and are therefore dependent on water scenario the distribution of suitable habitats for wild availability to prevent desiccation. In July, in particular,    yaks would expand by and %by in the growing vegetation in the Kobresia-dominant moist meadows is nor-  and non-growing seasons, respectively. Under the RCP mally at its early phenological stages (Schaller, ); timely scenario, however, the distribution of suitable habitats and abundant precipitation could thus be particularly bene-  during the growing season would expand by %, whereas ficial to plant development in these meadows. Studies from the availability of suitable habitats during the non-growing other parts of the plateau on the Tibetan ammon   season is expected to decrease by %(Fig. ). Shifts in the hodgsoni (Singh et al., ) and kiang Equus kiang distribution of suitable habitats are also expected to occur. (St-Louis & Côté, ) similarly suggest that forage quality Based on our analyses the distribution of suitable habitats can be a key factor shaping habitat selection patterns for  during the growing season could shrink by % under these large herbivores. At this stage it is difficult to be con-    RCP , and % under RCP . Likewise, the distribution clusive about the role of forage quantity and quality in driv- of suitable habitats during the non-growing season could ing wild yak habitat selection; further research is needed.     shrink by % under RCP , and % under RCP The second prediction that our results failed to support is  (Supplementary Table S ). For information on topographical that wild yaks avoid human settlements, especially during features of potential suitable habitats under both RCPs, the period when forage is relatively abundant and when  see Supplementary Table S . there is thus no need to take greater risks associated with proximity to humans (Frid & Dill, ; Creel et al., Discussion ). The low influence of anthropogenic disturbance on the distribution of wild yaks may suggest that individuals Our results largely support current expectations about the in the area are essentially unaffected by human distribution factors shaping wild yak distribution on the Tibetan during the growing season, but this result may also be

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FIG. 3 Distribution of suitable habitat for wild yaks in (a) the vegetation growing season and (b) the non-growing season, and the predicted distributions under the RCP scenario (a &b) and under the RCP scenario for both seasons (a &b).

underpinned by the spatial proximity of villages to resulting from human presence from disturbance resulting Kobresia-dominant moist meadows. Another potential ex- from livestock. This lack of differentiation is a result of a planation is based on the distribution of domestic yaks, lack of information on the spatial distribution of people found near villages. Habitat selection patterns of polygyn- and livestock in the area. As efforts to gather such data ous male herbivores are likely to be dependent on the spa- increase it would be interesting to contrast the influence tio–temporal distribution of females during the mating of humans and livestock on the distribution of wild yaks. season (Jarman, ; Clutton-Brock, ). One could ex- Thirdly, the dataset may have been biased by the survey pect wild males to be attracted by the presence of a large methods. In the growing season, in particular, limited number of domestic females, with no competitors present. accessibility to various areas can limit survey efforts to This hypothesis is supported by the increasingly reported regions closer to villages, and therefore our dataset may mingling and hybridization of wild and domestic yaks in not capture the full range of environmental conditions Tibet (Leslie & Schaller, ). where yaks can be found during this period. This sampling There are a number of caveats associated with our data bias could lead to underestimation of the distribution and and modelling work. Firstly, apart from geographical coor- size of suitable habitats during the growing season, as well dinates and group size the survey teams recorded no other as misidentification of the ecological forces shaping the dis- observations (e.g. topography, climatic conditions, primary tribution of the species (Syfert et al., ). Moreover, this productivity). All the environmental information used in study, based on a series of correlative modelling approaches, the analyses was therefore derived from global products, intrinsically assumes that wild yaks are living in equilibrium which have not been validated locally. Future research with their environment (Pearson & Dawson, ); how- should ground-truth these products to ascertain the ever, the observed distribution may not reflect the optimal robustness of our conclusions. Secondly, our proxy of an- patterns of habitat selection but rather habitat use as con- thropogenic disturbance does not differentiate disturbance strained by a number of factors, including those associated

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with the presence of livestock. To address this, future of any conservation scheme aiming to mitigate the impact large-scale studies should attempt to incorporate informa- of climatic change, helping them to track shifting climatic tion on the distribution of domestic yaks when modelling zones and colonize new territories. Our findings also suggest the distribution of wild yaks. Various biotic interactions, that the number of domestic yak holdings should be more as well as the dispersal ability of wild yaks, need to be strictly controlled in communities adjacent to known wild taken into account to identify scale-dependent limiting fac- yak populations. Livestock grazing activities should be lim- tors and consequent patterns in habitat selection (Pearson & ited to designated areas to minimize competition for the Dawson, ). Another limitation to this study is that we winter resources of wild yaks. These two points are especial- did not consider the influence of sex. Dimorphic ly relevant for two regions that include parts of the Ali and can be substantially divergent in their niche requirements Naqu prefectures, where there are high densities of wild (Kie & Bowyer, ). We were unable to explore differences yaks. These regions are likely to remain suitable for the spe- in habitat selection patterns between males and females cies under both RCP scenarios during the growing season; because the sex of the individuals sighted was not recorded however, they may not remain so during the non-growing reliably. The seasonal patterns identified should thus be season. They are not within the borders of any protected understood as averaged results based on the unknown sex area and are subject to high levels of human activity. ratio. Finally, uncertainties associated with the modelling ap- Conservation interventions may be necessary in these re- proaches considered should be acknowledged (Araújo et al., gions and we recommend the establishment of monitoring ). Predictions derived from these models vary substan- systems as soon as possible, to assess any direct threats, such tially; for example, if we adopted the predictions of the gener- as illegal hunting. Current patterns of land use (e.g. grazing alized additive model (which is also acceptable in terms of sites for domestic yaks) should be evaluated and possibly AUC and TSS) the importance of factors such as altitude rearranged to take into consideration the habitat needs of and mean temperature in determining suitable habitats for wild yaks. Lastly, we recommend the rapid definition wild yaks in summer would be much higher than suggested and implementation of a plan to connect these regions to by the random forests model (Supplementary Fig. S), and the nearest protected areas that contain other wild yak suitable habitats would be more extensive in both seasons populations. (Supplementary Table S). Such method-induced variability highlights the importance of interpreting model outputs with caution. Acknowledgements An important contribution of this study is its quantifica- tion of the possible impact of climate change on the avail- Special thanks are due to George Schaller, Madhu Rao, ability of suitable habitats for wild yaks. According to our Manuel Mejia, Marcus Rowcliffe, Joel Berger, Clare current knowledge wild yaks are mostly found at –°N, Duncan, Xueyan Li, Josephine Arthur and two anonymous in regions that are likely to be severely affected by climate reviewers for their generous help in developing this study, change. In terms of conservation priorities for the species, and valuable comments on the manuscript. its autumn and winter habitats appear to be more vulnerable to the impacts of climate change than its spring and summer habitats, and a lower winter to summer habitat ratio may re- References present a high risk to population stability. In a seasonal graz- ing system, ungulate population size can be more sensitive ACHARD,F.&ESTREGUIL,C.() Forest classification of Southeast  to limited availability of resources during the non-growing Asia using NOAA AVHRR data. Remote Sensing of Environment, , –. season than to abundance of resources during the growing  ’  ALLOUCHE, O., TSOAR,A.&KADMON,R.( ) Assessing the season (Illius & O Connor, ). In the future the distribu- accuracy of species distribution models: prevalence, kappa and the tion of suitable habitat during the growing season is more true skill statistic (TSS). Journal of Applied Ecology, , –. likely to be threatened by anthropogenic activities than by ARAÚJO, M.B., PEARSON, R.G., THUILLER,W.&ERHARD,M.() climate change. Any increase in the distribution of suitable Validation of species–climate impact models under climate change.  – habitats will create economic opportunities for domestic yak Global Change Biology, , . AUSTIN,M.() Species distribution models and ecological theory: herders, which could result in resource competition between a critical assessment and some possible new approaches. 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XU, A., JIANG, Z., LI, C., GUO, J., WU,G.&CAI,P.() Summer Biographical sketches food habits of brown bears in Kekexili Nature Reserve, Qinghai: Tibetan plateau, China. Ursus, , –. X UCHANG L IANG and A ILI K ANG are conservation biologists. YEE, T.W. & MITCHELL, N.D. () Generalized additive models in They have spent years monitoring yaks and studying their ecology on plant ecology. Journal of Vegetation Science, , –. the Tibetan plateau. N ATHALIE P ETTORELLI heads the Environmental YU, H., XU, J., OKUTO,E.&LUEDELING,E.() Seasonal response Monitoring and Conservation Modelling team at the Institute of Zoology, of grasslands to climate change on the Tibetan Plateau. PLoS ONE,  UK, which is interested in developing tools and methodologies to support (), e. the sustainable management of natural resources.

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