Partial Altitudinal Migration of the Near Threatened Satyr Tragopan Tragopan Satyra in the Bhutan Himalayas: Implications for Conservation in Mountainous Environments

Home , Altitudinal migration, Satyr tragopan

Partial altitudinal migration of the Near Threatened satyr tragopan Tragopan satyra in the Bhutan Himalayas: implications for conservation in mountainous environments

N AWANG N ORBU,UGYEN,MARTIN C. WIKELSKI and D AVID S. WILCOVE

Abstract Relative to long-distance migrants, altitudinal mi- To view supplementary material for this article, please visit grants have been understudied, perhaps because of a percep- http://dx.doi.org/./S tion that their migrations are less complex and therefore easier to protect. Nonetheless, altitudinal migrants may be at risk as they are subject to ongoing anthropogenic pressure from land use and climate change. We used global position- Introduction ing system/accelerometer telemetry to track the partial altitudinal migration of the satyr tragopan Tragopan satyra in elative to long-distance migrants, altitudinal migrants central Bhutan. The birds displayed a surprising diversity of Rhave been understudied, perhaps because of a percep- migratory strategies: some individuals did not migrate, tion that their migrations are less complex and easier to pro- others crossed multiple mountains to their winter ranges, tect. In montane regions many species migrate altitudinally others descended particular mountains, and others as- up and down mountain slopes (Stiles, ; Powell & Bjork, cended higher up into the mountains in winter. In all ; Burgess & Mlingwa, ; Chaves-Champos et al., cases migration between summer breeding and winter ; Faaborg et al., ). Although attempts have been non-breeding grounds was accomplished largely by walk- made (Laymon, ; Cade & Hoffman, ; Powell & ing, not by flying. Females migrated in a south-easterly dir- Bjork, ; Chaves-Champos et al., ; Hess et al., ection whereas males migrated in random directions. ), few studies have illustrated patterns of altitudinal mi- During winter, migrants occupied south-east facing slopes gration using telemetry (but see Norbu et al., ). whereas residents remained on south-west facing slopes. Montane regions, which cover an estimated .% of land Migratory and resident tragopans utilized a range of forest surface area (Kapos et al., ), are being exposed to cli- types, with migratory individuals preferring cool broad- mate change (Nogués-Bravo et al., ) and loss of forest leaved forests during winter. These complex patterns of mi- cover (Blyth et al., ; Pandit et al., ), both of gration suggest that conservation measures should extend which will put mountain species and their migrations at across multiple mountains, protect the full range of forest risk (Inouye et al., ). types and encompass multiple landscape configurations to In general, animal migrations are undergoing decline protect aspect diversity. Given the diversity of migratory (Wilcove & Wikelski, ; Harris et al., ) as a result strategies employed by this single species it seems clear of habitat loss and climate change (Both et al., ; that more research on altitudinal migrants is needed to Møller et al., ). These declines are of concern because understand what must be done to ensure their future in migration is important in maintaining ecological processes an era of widespread land-use and climate change. and shaping ecosystems (Holland et al., ; Wilcove & Wikelski, ; Bowlin et al., ). However, the conserva- Keywords Altitudinal migration, Bhutan, conservation, tion of migratory species remains a daunting task, given the mountains, partial migration, protected areas, satyr trago- complexity of the phenomenon and the geographical scale pan, Thrumshingla National Park at which it occurs (Moore et al., ; Wilcove, ; Faaborg et al., ).

NAWANG NORBU* (Corresponding author) and UGYEN Ugyen Wangchuck In comparison to long-distance migrants, the conserva- Institute for Conservation and Environment, Lamai Gompa Dzong, Bumtang, tion of altitudinal migrants may be perceived to be relatively Bhutan. E-mail [email protected] easier given their occurrence over a smaller geographical MARTIN C. WIKELSKI Max Planck Institute for Ornithology, Radolfzell, Germany area. This may not necessarily be the case, however. For ex- DAVID S. WILCOVE Woodrow Wilson School for Public and International Affairs ample, telemetry studies (Powell & Bjork, , , ) and Department of Ecology and Evolutionary Biology, Princeton University, USA have shown that habitat types required by two altitudinally migrating species, the resplendent quetzal Pharomachrus *Also at: The International Max Planck Research School for Organismal Biology, University of Konstanz, Germany mocinno and three-wattled bellbird Procnias tricarunculata, ’ Received  January . Revision requested  February . were not included within Costa Rica s protected area system. Accepted  June . First published online  March . To ensure the protection of altitudinal migrants, the

Oryx, 2017, 51(1), 166–173 © 2016 Fauna & Flora International doi:10.1017/S0030605315000757 Downloaded from https://www.cambridge.org/core. IP address: 170.106.202.226, on 03 Oct 2021 at 04:14:29, subject to the Cambridge Core terms of use, available at https://www.cambridge.org/core/terms. https://doi.org/10.1017/S0030605315000757 Satyr tragopan in the Bhutan Himalayas 167

FIG. 1 Location of the study area in Thrumshingla National Park, Bhutan.

incorporation of all habitat types within reserve systems has are believed to be extant in the wild (BirdLife International, been recommended as a principle of effective reserve design b). The tragopan has been shown to be a partial altitud- (Powell & Bjork, , , ). inal migrant (Norbu et al., ). Here, we assess the adequacy Establishment of protected areas (Terborgh et al., ; of a protected area in Bhutan to provide protection for an alti- Brooks et al., ; Cantú-Salazar et al., ) and biological tudinal migrant, by examining the migration mode, patterns corridors (Roever et al., ) has become one of the key and habitat requirements of the tragopan. strategies to protect biodiversity. However, the adequacy and effectiveness of protected areas and corridors have Study area been questioned (Hobbs, ; Beier & Noss, ; Ervin, ), particularly regarding the conservation of migratory We studied the tragopan in Thrumshingla National Park, a  species (Moore et al., ; Thirgood et al., ; Martin  km protected area in central Bhutan (Fig. ), at ,– et al., ); for example, wildlife movement data have , m elevation. Mean daily temperature is −–°C. The been used in only a few instances to influence corridor de- area has four distinct seasons, with most rainfall occurring sign (Zeller et al., ). during May–August as part of the Asian monsoons. The With  protected areas, the network of protected areas  study area is mostly covered by conifer forests dominated in the Himalayas covers an estimated , km (Chettri by Bhutan fir Abies densa, with rhododendron understorey et al., ). Despite this extensive coverage we are not at higher elevations (. , m) transitioning to mixed aware of any telemetry-based study of altitudinal migrants conifer forests (,–, m) comprising Sikkim spruce to assess the effectiveness of protected areas in the Picea spinulosa, Himalayan hemlock Tsuga dumosa and Himalayas. Given that the Himalayas are undergoing habi- Sikkim larch Larix griffithii. Below , m conifer forests tat loss (Pandit et al., ) and climate change (Shrestha give way to conifer–broadleaved mixed forests and to cool et al., ), and noting that many of the birds in the broadleaved forests of oak Quercus glauca and Quercus la- Himalayas are altitudinal migrants (Inskipp et al., ; mellosa. There are also a few patches of open grazing areas Grimmet et al., ; BirdLife International, ), data that are used by nomadic cattle herders. are needed to test the adequacy of conservation landscapes in protecting altitudinally migrating species. We used global positioning system (GPS)/accelerometer Methods telemetry to track the seasonal migration of the satyr trago- Trapping We trapped tragopans during June–October in pan Tragopan satyra, a pheasant endemic to the central and ,  and  using neck noose traps laid along ridges eastern Himalayas of Bhutan, India and Nepal. The species known to be used by tragopans, which we barricaded with (hereafter referred to as the tragopan) is categorized as Near bamboo and shrubs. We flushed tragopans towards traps Threatened on the IUCN Red List (BirdLife International, during early mornings and evenings. In , when the a) and listed on Appendix III of CITES (). Only an birds were trapped for the first time, all captured individuals estimated , individuals (c. ,–, mature adults) were released immediately after GPS tags were attached, to

reduce the risk of any handling-related fatalities. Tragopans period (December–February) we extracted the aspect (in captured in  and  were weighed (to the nearest g) degrees) of the given location from a digital elevation  and their tarsus length (mm) and beak size (mm) were model at a spatial resolution of  x  m . For each measured. All trapping was approved by the Ministry of individual, using the Rayleigh statistic at a significance Agriculture and Forests in Bhutan. level of ., we assessed whether the aspect data were distributed uniformly or whether the individual showed a preference for a particular aspect. For individuals with a GPS/accelerometer tags and data acquisition We used significant preference for a certain aspect, we calculated GPS/accelerometer tags (e-obs, Munich, Germany) to the mean aspect for each individual and calculated record the location and activity of tagged tragopans. The summary statistics separately for two groups: migrants recorded data (i.e. location, elevation, date, time and and residents. We used Oriana v. . (Kovach acceleration) were downloaded remotely with a hand-held Computing Services, Anglesey, UK) for all circular statistics. base station after each individual was relocated via its tag’s VHF radio pulse. To help locate tagged birds the tags were   programmed to ping every seconds for hours every day. Quantifying bouts of flying/running, walking and resting  Tags with harnesses weighed g. Except for the smallest We used accelerometer data (Wilson et al., ; Brown  individual, which weighed g, all tagged tragopans et al., ) to quantify bouts of flying/running, walking .    weighed kg, with the largest weighing . kg; the mean and resting. Acceleration data were obtained for the    weight of individuals was . kg. At the mean weight, vertical z-axis (up-and-down axis) continuously for   tags constituted % of the body weight; this is within the hours every day from tag initiation to the last date of tag   acceptable limit of % (Hawkins, ). Tags deployed in download. Acceleration data consisted of  raw   were programmed to record a GPS reading every acceleration values within every -minute interval (bout).  –  hours during . . . Given battery power constraints, We calculated differences between successive raw  in tags were programmed to record only two GPS observations and obtained the variances of these      readings per day, at . and . ,andin they differences for each -minute bout. We then used the       recorded three readings per day, at . , . and . . variances to categorize each -minute bout into one of To optimize battery performance, tags were programmed three categories of activity: flight/running, walking/  with a give-up time of minutes, after which the tag would foraging or resting. We categorized all variances ,  as cease trying to obtain a GPS reading for a particular location. resting, –, as walking/foraging, and . , as flying/running. We could not separate flying and running because the activity signatures for the two were difficult to Distance, duration and direction of migration We distinguish. We checked the accuracy of our categorization categorized all individuals that showed distinct by cross checking with categorizations obtained using non-overlapping summer and winter ranges (i.e. Acceleration Viewer (Movebank, Max Planck Institute of individuals that stayed .  months at winter ranges) as Ornithology, Munich, Germany). To calibrate activity migrants and the remainder as residents. Using the Show signatures for the software we observed a tagged domestic Elevation Profile tool in Google Earth v. . (Google Inc., chicken Gallus gallus for  days and recorded the times Mountain View, USA) we measured migration distance as when it walked, ran, flew and rested. We then inspected the distance between the location on the day when its activity patterns in Acceleration Viewer. migration was initiated and the first location on the day when migration was terminated. We ascertained the direction of migration between summer and winter Data analysis and archiving All geographical information habitat using the Ruler tool in Google Earth. system operations were carried out with ArcGIS v. .. (ESRI, Redlands, USA) and statistical tests with Rv... (R Development Core Team, ). All data have been Habitat use patterns For all individuals we overlaid GPS archived at Movebank (). fixes on a land-cover map produced by the Ministry of Agriculture and Forests (). We counted total fixes for all individuals in various land-cover types across months. Results To test whether habitat preferences changed over seasons,    we fitted generalized linear models in Rv. . . (R We obtained movement data for  individuals over  years  Development Core Team, ) to predict the proportion (–). Fourteen (nine females and five males) mi- of habitats used (as indicated by the total GPS fixes within grated and  (two females and eight males) did not. Of – a habitat in each month) across months during September the  migrants, we obtained return migration data for April. For every GPS point recorded during the winter only  females and  male (Table ).

TABLE 1 Numbers of satyr tragopans Tragopan satyra for which movement data were recorded during autumn and spring migrations in Thrumshingla National Park, Bhutan (Fig. ), during  –, including sedentary individuals.

No. of individuals for which data were recorded No. of tagged birds for which no data were Year Autumn migration Spring/return migration Sedentary individuals recorded 2009 2 (1 female, 1 male) 2 (1 female, 1 male) 6 2010 7 (5 females, 2 males) 0 1 (1 male) 6 2011 5 (3 females, 2 males) 3 (2 females, 1 male) 7 (1 female, 6 males) 2 2012 2 (2 females) Total 14 (9 females, 5 males) 5 10 (2 females, 8 males) 14

FIG. 2 (a) Autumn migratory routes of  satyr tragopans Tragopan satyra, including the individual with tag ID  (Supplementary Fig. S). Grey lines indicate the routes of two individuals that migrated to higher elevations in winter. The start and end points of the migrations are indicated by filled rectangles and circles, respectively. (b) Year-round locations of resident individuals, each represented by a unique symbol.

Migration patterns, distance and duration Three distinct birds traversed a mean distance of . ± %CI. km patterns of migration were recorded: seven individuals between summer and wintering grounds, in . ± %CI (five females and two males) crossed multiple mountains  days. Individuals crossing multiple mountains migrated between summer breeding and winter non-breeding further than individuals descending longitudinally (n = , grounds; five individuals (three females and two males) t = .,P=.). However, there were no significant descended longitudinally (i.e. parallel to mountain slopes) differences in the number of migration days between to winter grounds; and two individuals (one female and those two categories (n = , t = .,P=.). Given one male) climbed to higher elevations during winter. The that only two individuals climbed to higher elevations in

winter, we excluded them from the analyses of migratory distances and days travelled. Females traversed a mean distance of . ± %CI. km, migrating preferentially in a south-easterly direction, with a mean bearing of .°(n=; Rayleigh test, P = .), whereas males migrated in random directions (n = ; Rayleigh test, P=.), covering a mean distance of . ± %CI. km. All migrants remained within the boundaries of Thrumshingla National Park (Fig. a). Of the  migrants, seven crossed the highway once and two crossed it at two locations during their migration.

Mode of migration Tragopan migration is not a single-flight event. The birds covered the migratory distance by walking, interspersed with short bouts of running/flight. When the migration spread over multiple days and involved crossing multiple mountains, the birds stopped to rest on mountain slopes (Supplementary Fig. S). For all migrants throughout the migratory period FIG. 3 Habitat use by satyr tragopans in Thrumshingla National bouts of running/flight (median = .) were significantly Park (Fig. ) in each month, as indicated by the proportion of less frequent than bouts of walking (median = .). All GPS fixes recorded in each habitat type. Numbers above the bars bouts of running/flight were singular events lasting ,  indicate total number of individuals for which GPS fixes were obtained in a given month. minutes. Resting bouts (median = .) were in turn significantly more frequent than both walking and flight/ running bouts (Kruskal–Wallis ANOVA: H = ., μ   – P , .; Tukey test: rest vs flight, q = .,P, .; rest (mean vector, )of . ° during winter (December vs walk, q = .,P, .; walk vs flight, q = .,P, .). February). During the same period migrants preferred south-east facing slopes (n = ; Rayleigh test, P = .)at a mean aspect of .°(Fig. ). During summer (May–  Habitats used For individuals we obtained complete July) both migrants and residents preferred to stay on data for the winter months of December, January and south-west facing slopes (n =  ( sedentary and  February, and for eight individuals we obtained complete migrants); Rayleigh test, P = .; Fig. ). data for May, June and July. Migratory individuals used all forest types (fir, mixed conifer and cool broadleaved; Figs a & ) in the study area (Fig. ). Over an -month period (September–April) we identified a significant Discussion change in the pattern of habitat use (generalized linear  In general, altitudinal migrants have been poorly studied, model; for cool broadleaved forest: adjusted R = .,  and this is especially the case in the Himalayas. Our study P=.; for mixed conifer forests: adjusted R = ., of the satyr tragopan in Bhutan yielded new information P=.), with migrants using cool broadleaved forests that will inform the conservation of this species and possibly more frequently, switching from high-elevation fir and other terrestrial, forest-dependent altitudinal migrants, par- mixed conifer forests during winter (November– ticularly Galliformes. Whereas some of the individuals in February). Nevertheless, the tragopans did not completely  our study population simply moved down a mountain for abandon mixed-conifer forests during winter (χ test: the winter, others chose to cross multiple mountains to df = ,P=.), with residents and some migrants still reach a suitable winter range. A simplistic conservation continuing to inhabit these forests. Five of  residents plan focused on protecting habitats along elevational remained only within mixed-conifer forests throughout bands on a given mountain slope would not protect those the year, two remained only within cool broadleaved individuals travelling between and across mountains. forests, two used both mixed conifer and cool broadleaved Moreover, tragopans migrate predominantly by walking, in- forests, and one used a mix of mixed conifer, conifer– terspersed with short bouts of running or flight through the broadleaved and cool broadleaved forest (Fig. b). forest. This raises the possibility that certain forms of land- use change could create barriers to the species’ migration, Selection of aspect Residents preferred south-west facing and underscores the need to protect contiguous tracts of slopes (n = ; Rayleigh test, P = .) at a mean aspect suitable habitat.

FIG. 4 Preferred aspect of (a) resident satyr tragopans and (b) migrants in Thrumshingla National Park (Fig. ) during the winter months (December–February). Mean aspect and confidence intervals are indicated by the dashed line and dashed arc, respectively.

The conservation requirements for birds that migrate from their sedentary counterparts in their choice of aspect primarily by walking may be more similar to those of mi- during winter, with migrants switching from south-east to grating ungulates than to birds that migrate by flying. As south-west facing slopes while residents continue to reside is the case with migratory ungulates (Bolger et al., ), on south-east facing slopes. We propose that protected barriers and land-use changes along migratory routes can areas, in addition to having representative habitat types, be detrimental to birds such as tragopans. Unlike ungulates should also have adequate representation of all aspects with- (Williamson & Williamson, ; Ben-Shahar, ; in a landscape. Protecting a diversity of aspects could also Mwangi, ), however, the tragopan’s ability to fly even benefit the conservation of a variety of other altitudinal mi- short distances may render it less susceptible to minor bar- grants (Poole et al., ; Zeng et al., ). riers such as fences. Over an annual cycle we observed that north-west facing Migratory tragopans used all forest types across their an- slopes were not preferred by any of the individuals, resident nual cycles, using mostly high-elevation fir and mixed coni- or migratory, that we tracked. Within Thrumshingla  fer forests in summer and cool broadleaved forests in winter. National Park almost % (c.  km ) of the total area has Residents and a few migrants continued to use mixed coni- a north-west aspect, and studies targeted at estimating the fer forests during winter, thus it is important to protect con- tragopan population size in the Park should perhaps contiguous tracts of all habitat types across all seasons. Similar sider excluding north-west facing slopes as suitable habitats. observations were made by Powell & Bjork (, ) for However, our sample size is small and we suggest that fur- altitudinally migrating birds in Costa Rica. ther study of aspect preference should be conducted to ver- Changes in habitat use from summer to winter were not a ify our assertion. Nonetheless, across protected areas in simple transition to lower-elevation forests (and vice-versa montane regions, if altitudinal migrants exhibit such prefer- during spring). Our results reveal a complex pattern where- ences, population estimates may need to be revised. by individuals also migrated within a single habitat type (i.e. Studies have shown continuing loss of forest cover across from higher-elevation mixed conifer forest to lower- many regions (Hansen et al., ). However, we remain op- elevation mixed conifer forest). Two individuals migrated timistic that migrants can persist in countries such as to lower elevations within a forest transition complex con- Bhutan where protected areas have retained their forests sisting of patches of mixed conifer and cool broadleaved for- (Thrumshingla National Park has an estimated % forest ests, whereas one female and one male migrated to cover) and large-scale commercial harvesting of forest with- higher-elevation fir forests from mixed-conifer forests dur- in protected areas is not permitted. All the migrants we ing winter. In addition to supporting the need to protect for- tracked remained within the boundaries of the Park, and est habitats with adequate representation of all habitat types, their migratory tracks covered c. one third of the Park’s our data also highlight the importance of individual differ- area. It may therefore be necessary to protect large areas ences and choice (Bolnick et al., ) in shaping migratory for the sake of relatively few migrants, and in this regard a patterns. detailed assessment of park sizes and migrant distribution In addition to protecting all habitat types, we propose an across montane regions would be of value. additional requirement for the effective conservation of al- We hypothesize that the movement of migrants to areas titudinally migrating species within montane environments: outside the Park boundaries may be restricted by human conservation of aspect diversity. Migratory tragopans differ settlements surrounding the Park. Additional studies are

necessary to determine the types and intensities of land uses BOLGER, D.T., NEWMARK, W.D., MORRISON, T.A. & DOAK, D.F. that tragopans can tolerate. Several migrants crossed the na- () The need for integrative approaches to understand and  – tional highway at multiple locations, suggesting that the conserve migratory ungulates. Ecology Letters, , . BOLNICK, D.I., SVANBÄCK, R., FORDYCE, J.A., YANG, L.H., DAVIS,J. highway is not currently a barrier to tragopan migration. M., HULSEY, C.D. & FORISTER, M.L. () The ecology of This could be because highways in Bhutan are narrow individuals: incidence and implications of individual specialization. (mean width ,  m). Increases in traffic intensity and high- The American Naturalist, , –. way width should therefore be monitored to track potential BOTH, C., BOUWHUIS, S., LESSELLS, C.M. & VISSER, M.E. () risks for migrants. Climate change and population declines in a long-distance migratory bird. Nature, , –. Our findings are relevant to discussions regarding the BOWLIN, M.S., BISSON, I.-A., SHAMOUN-BARANES, J., REICHARD,J. role of habitat corridors in protecting biodiversity. Given D., SAPIR, N., MARRA, P.P. et al. () Grand challenges in ongoing deforestation (Hansen et al., ) and other land- migration biology. Integrative & Comparative Biology, , –. use changes that impede migration, the concept of connect- BROOKS, T.M., BAKARR, M.I., BOUCHER, T., DA FONSECA, G.A.B.,  ing protected areas with habitat corridors has gained HILTON-TAYLOR, C., HOEKSTRA, J.M. et al. ( ) Coverage provided by the global protected-area system: is it enough? prominence. However, much remains to be learned regard- BioScience, , –.  ing the effectiveness of such corridors (Hobbs, ; Beier & BROWN, D.D., KAYS, R., WIKELSKI, M., WILSON,R.&KLIMLEY, A.P. Noss, ). Our finding that female tragopans tend to mi- () Observing the unwatchable through acceleration logging of grate in a south-easterly direction whereas males have no animal behavior. Animal Biotelemetry, , .  particular orientation suggests that corridors are likely to BURGESS, N.D. & MLINGWA, C.O.F. ( ) Evidence for altitudinal migration of forest birds between montane Eastern Arc and lowland be no more than partially successful in protecting tragopan forests in East Africa. Ostrich, , –. migrations. For male tragopans in particular, corridors may CADE, B.S. & HOFFMAN, R.W. () Differential migration of blue prove to be a poor substitute for large, continuous blocks of grouse in Colorado. The Auk, , –. forest. CANTÚ-SALAZAR, L., ORME, C.D.L., RASMUSSEN, P.C., BLACKBURN, Given that most montane bird species of Bhutan and the T.M. & GASTON, K.J. () The performance of the global Himalayas are altitudinal migrants, and given the complex protected area system in capturing vertebrate geographic ranges. Biodiversity and Conservation, , –. patterns of migration and habitat use exhibited by a single CHAVES-CAMPOS, J., ARÉVALO, J.E. & ARAYA,M.() Altitudinal species, the satyr tragopan, it seems clear that more research movements and conservation of bare-necked umbrellabird on altitudinal migrants is needed to understand fully what Cephalopterus glabricollis of the Tilarán Mountains, Costa Rica. Bird will be required to ensure their future in an era of wide- Conservation International, , –.  spread land-use and climate change. CHETTRI, N., SHAKYA, B., THAPA,R.&SHARMA,E.( ) Status of a protected area system in the Hindu Kush–Himalayas: an analysis of PA coverage. International Journal of Biodiversity Science and Management, , –. Acknowledgements CITES () The CITES Appendices. Http://www.cites.org/eng/app/ index.shtml [accessed  August ]. We are grateful to the International Max Planck Research ERVIN,J.() Rapid assessment of protected area management  – School for Organismal Biology and the Max Planck effectiveness in four countries. BioScience, , . Institute for Ornithology, Radolfzell, for funding this work. FAABORG, J., HOLMES, R.T., ANDERS, A.D., BILDSTEIN, K.L., DUGGER, K.M., GAUTHREAUX,JR, S.A. et al. () Recent advances in understanding migration systems of New World land birds. Ecological Monographs, , –. References GRIMMET, R., INSKIPP,C.&INSKIPP,T.() Birds of Nepal. Princeton University Press, Princeton, USA. BEIER,P.&NOSS, R.F. () Do habitat corridors provide HANSEN, M.C., POTAPOV, P.V., MOORE, R., HANCHER, M., connectivity? Conservation Biology, , –. TURUBANOVA, S.A., TYUKAVINA, A. et al. () High-resolution   – BEN-SHAHAR,R.() Does fencing reduce the carrying capacity global maps of st-century forest cover change. Science, , for populations of large herbivores? Journal of Tropical Ecology, , . –. HARRIS, G., THIRGOOD, S., HOPCRAFT, J.G.C., CROMSIGT, J.P.G.M. &  BIRDLIFE INTERNATIONAL (a) Tragopan satyra.InThe IUCN Red BERGER,J.( ) Global decline in aggregated migrations of large List of Threatened Species v. .. Http://www.iucnredlist.org terrestrial mammals. Endangered Species Research, , –. [accessed  August ]. HAWKINS,P.(). Bio-logging and animal welfare: practical  BIRDLIFE INTERNATIONAL (b) Species factsheet: Satyr tragopan refinements. Memoirs of National Institute of Polar Research, , Tragopan satyra. Http://www.birdlife.org/datazone/ –. speciesfactsheet.php?id= [accessed  November ]. HESS, S.C., LEOPOLD, C.R., MISAJON, K., HU,D.&JEFFREY, J.J. () BIRDLIFE INTERNATIONAL () Endemic Bird Area factsheet: Restoration of movement patterns of the Hawaiian goose. The Eastern Himalayas. Http://www.birdlife.org/datazone/ebafactsheet. Wilson Journal of Ornithology, , –. php?id= [accessed  May ]. HOBBS, R.J. () The role of corridors in conservation: solution or  – BLYTH, S., GROOMBRIDGE, B., LYSENKO, I., MILES,L.&NEWTON,A. bandwagon? Trends in Ecology & Evolution, , . () Mountain Watch: Environmental Change & Sustainable HOLLAND, R.A., WIKELSKI,M.&WILCOVE, D.S. () How and Development in Mountains. UNEP–WCMC, Cambridge, UK. why do insects migrate? Science, , –.

INOUYE, D.W., BARR, B., ARMITAGE, K.B. & INOUYE, B.D. () POWELL, G.V.N. & BJORK, R.D. () Habitat linkages and the Climate change is affecting altitudinal migrants and hibernating conservation of tropical biodiversity as indicated by seasonal species. Proceedings of the National Academy of Sciences of the migrations of three-wattled bellbirds. Conservation Biology, , – United States of America, , –. . INSKIPP, C., INSKIPP,T.&GRIMMET,R.() Birds of Bhutan. RDEVELOPMENT CORE TEAM () R: A Language and Environment Christopher Helm Publishers Ltd, London, UK. for Statistical Computing. R Foundation for Statistical Computing, KAPOS, V., RHIND, J., EDWARDS, M., PRICE, M.F., RAVILIOUS, C., Vienna, Austria. KREMSA,V.etal.() Developing a map of the world’s mountain ROEVER, C.L., VAN AARDE, R.J. & LEGGETT,K.() forests. In Forests in Sustainable Mountain Development: A State of Functional connectivity within conservation networks: Knowledge Report for  (eds M.F. Price & N. Butt), pp. –. delineating corridors for African elephants. Biological Conservation, CABI Publishing, Wallingford, UK. , –. LAYMON, S.A. () Altitudinal migration movements of spotted owls SHRESTHA, U.B., GAUTAM,S.&BAWA, K.S. () Widespread climate in the Sierra Nevada, California. The Condor, , –. change in the Himalayas and associated changes in local ecosystems. MARTIN, T.G., CHADÈS, I., ARCESE, P., MARRA, P.P., POSSINGHAM,H. PLoS ONE, (), e. P. & N ORRIS, D.R. () Optimal conservation of migratory STILES, F.G. () Altitudinal movements of birds on the Caribbean species. PLoS ONE, (), e. slope of Costa Rica: implications for conservation. In Tropical MINISTRY OF AGRICULTURE AND FORESTS () Landcover Atlas of Rainforests: Diversity and Conservation (eds F. Almeda & Bhutan. Ministry of Agriculture and Forests, Thimphu, Bhutan. C.M. Pringle), pp. –. California Academy of Sciences, MØLLER, A.P., RUBOLINI,D.&LEHIKOINEN,E.() Populations of San Francisco, USA. migratory bird species that did not show a phenological response to TERBORGH, J., VAN SCHAIK, C., DAVENPORT,L.&RAO,M.() climate change are declining. Proceedings of the National Academy Making Parks Work: Strategies for Preserving Tropical Nature. of Sciences of the United States of America, , –. Island Press, Washington, DC, USA. MOORE, F.R., GAUTHREAUX,JR, S.A., KERLINGER,P.&SIMONS, T.R. THIRGOOD, S., MOSSER, A., THAM, S., HOPCRAFT, G., MWANGOMO, () Habitat requirements during migration: important link in E., MLENGEYA, T. et al. () Can parks protect migratory conservation. In Ecology and Management of Neotropical Migratory ungulates? The case of the Serengeti wildebeest. Animal Birds: A Synthesis and Review of Critical Issues (eds T.E. Martin & D. Conservation, , –. M. Finch), pp. –. Oxford University Press, New York, USA. WILCOVE, D.S. () No Way Home: The Decline of the World’s Great MOVEBANK () Https://www.movebank.org/ [accessed  August Animal Migrations. Island Press, Washington, DC, USA. ]. WILCOVE, D.S. & WIKELSKI, M.C. () Going, going, gone: is MWANGI, E.M. () Large herbivore dynamics in the face of animal migration disappearing. PLoS Biology, (), e. insularization: the case of Lake Nakuru National Park, Kenya. WILLIAMSON,D.&WILLIAMSON,J.() Botswana’s fences and the African Journal of Ecology, , –. depletion of Kalahari wildlife. Oryx, , –. NOGUÉS-BRAVO, D., ARAÚJO, M.B., ERREA, M.P. & WILSON, R.P., SHEPARD, E.L.C. & LIEBSCH,N.() Prying into the MARTÍNEZ-RICA, J.P. () Exposure of global mountain systems intimate details of animal lives: use of a daily diary on animals. to climate warming during the st century. Global Environmental Endangered Species Research, , –. Change, , –. ZELLER, K.A., MCGARIGAL,K.&WHITELEY, A.R. () Estimating NORBU, N., WIKELSKI, M.C., WILCOVE, D.S., PARTECKE, J., UGYEN landscape resistance to movement: a review. Landscape Ecology, , TENZIN, U. et al. () Partial altitudinal migration of a Himalayan –. forest pheasant. PLoS ONE, (), e. ZENG, Z., BECK,P.&WANG,T.() Effects of plant phenology and PANDIT, M.K., SODHI, N.S., KOH, L.P., BHASKAR,A.&BROOK, B.W. solar radiation on seasonal movement of golden takin in the Qinling () Unreported yet massive deforestation driving loss of endemic Mountains, China. Journal of Mammalogy, , –. biodiversity in Indian Himalaya. Biodiversity and Conservation, , –. POOLE, K.G., STUART-SMITH,K.&TESKE, I.E. () Wintering strategies by mountain goats in interior mountains. Canadian Biographical sketches Journal of Zoology, , –. POWELL, G.V.N. & BJORK, R.D. () Implications of altitudinal NAWANG NORBU uses science and policy to inform conservation and migration for conservation strategies to protect tropical biodiversity: management of forests. UGYEN is interested in biodiversity assess- a case study of the resplendent quetzal Pharomachrus mocinno at ment, and management of threatened wildlife species. M ARTIN Monteverde, Costa Rica. Bird Conservation International, , –. W IKELSKI studies global wildlife migration systems and their impacts POWELL, G.V.N. & BJORK,R.() Implications of intratropical on the environment and society. D AVID W ILCOVE combines research migration on reserve design: case study using Pharomachrus in ecology and the social sciences for the protection of global biodiver- mocinno. Conservation Biology, , –. sity and migratory phenomena.