Climate-change impacts on understorey bamboo species and giant pandas in China s Qinling Mountains

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1 PUBLISHED ONLINE: 11 NOVEMBER 212 DOI: 1.138/NCLIMATE1727 Climte-chnge impcts on understorey moo species nd gint pnds in Chin s Qinling Mountins Mo-Ning Tunmu 1, Andrés Viñ 1, Julie A. Winkler 2, Yu Li 1, Weihu Xu 3, Zhiyun Ouyng 3 nd Jinguo Liu 1 * Climte chnge is thretening glol ecosystems through its impct on the survivl of individul species nd their ecologicl functions 1,2. Despite the importnt role of understorey plnts in forest ecosystems 3 5, climte impct ssessments on understorey plnts nd their role in supporting wildlife hitt re scrce in the literture. Here we ssess climte-chnge impcts on understorey moo species with n emphsis on their ecologicl function s food resource for endngered gint pnds (Ailuropod melnoleuc). An ensemle of moo distriution projections ssocited with multiple climtechnge projections nd moo dispersl scenrios indictes sustntil reduction in the distriutionl rnges of three dominnt moo species in the Qinling Mountins, Chin during the twenty-first century. As these three species comprise lmost the entire diet of the pnd popultion in the region, the projected chnges distriution suggest potentil shortge of food for this popultion, unless lterntive food sources ecome ville. Although the projections were developed under unvoidle simplifying ssumptions nd uncertinties, they indicte potentil chllenges for pnd conservtion nd underscore the importnce of incorporting interspecific interctions into climte-chnge impct ssessments nd ssocited conservtion plnning. Like mny understorey plnts 3,4, understorey moo species re n essentil component in mny forest ecosystems 6,7. They not only influence the species composition nd structurl complexity of forests 6,7, ut lso provide essentil food nd shelter for wildlife species, including one of the most endngered species in the world, the gint pnd 8,9. Deforesttion nd forest degrdtion re thretening the survivl of out hlf of ll moo species worldwide 1, nd climte chnge my present n dditionl significnt thret. Mny moo species re vulnerle to climte chnge ecuse their unusul extended sexul reproduction intervls (from 1 to 12 yr; ref. 11), long with limited seed dispersl ility 12, render them less cple of djusting their distriutions to the rpidly chnging climte projected to occur within this century 1. In ddition, unlike other moo species with rpidly growing underground stems, mny understorey moo species in mountin ecosystems hve limited vegettive dispersl ility (for exmple, out.2.35 m yr 1 for Bshni frgesii nd Frgesi roust) 13,14. Despite the vulnerility of moo species to climte chnge, knowledge of climte-chnge-induced dynmics of moo distriution nd their effects on other species is lcking. This leves n importnt knowledge gp of prticulr significnce for iodiversity conservtion efforts. Understorey moo is necessry component of gint pnd hitt ecuse pnds re extreme dietry specilists who devote most of their ctive time to feeding on lrge mounts of moo (up to 38 kg d 1 ; refs 8,9). The dependency of gint pnds on understorey moo not only ffects the present pnd distriution 15,16, ut is lso elieved to hve driven historicl chnges in pnd distriution, s moo distriutions shifted in response to climte fluctutions 8,9. Therefore, the tight moo pnd reltionship provides n excellent opportunity to exmine the potentil cscding effects of climte-chnge-driven shifts in plnt species distriutions on the distriution of trophiclly dependent wildlife species. Here we report n ssessment of the potentil impcts of climte chnge on moo distriution, nd emphsize the cscding effects on gint pnd hitt in the Qinling Mountins (locted in the northern prt of the present pnd distriutionl rnge 15 nd covering pproximtely qurter of the remining pnd hitt 17 ; Fig. 1). Together, three moo species, Qinling rrow (Frgesi qinlingensis), drgon-hed (F. drcocephl) nd wooden (B. frgesii), ccount for more thn 9% of moo cover in this region 15. They constitute the min diet of the pnd popultion in the region (out 27 pnd individuls corresponding to 17% of the entire wild pnd popultion 15 ). Although this pnd popultion lives in region tht, t present, hs rguly the est hitt conditions nd receives the est protection cross the entire pnd distriutionl rnge 17, it is oth geogrphiclly (Fig. 1) nd geneticlly isolted 8 from other popultions. Our ssessment contins three min steps. First, using ioclimtic models to ssocite moo-presence loctions recorded in the field with ioclimtic vriles derived from oservtionl climtologies 18, we chrcterized nd mpped present suitle climte conditions for ech of the three moo species (Fig. 1). The predicted climticlly suitle res (CSAs) effectively cptured the oserved moo-presence loctions ccording to model-ccurcy evlutions (see Supplementry Methods). The sptil ptterns of predicted CSAs were lso consistent with our understnding of the present moo distriution 8,15 (tht is, Qinling rrow nd wooden moos re distriuted from 1,8 to 3, m, nd from 9 to 1,9 m, respectively, with the distriution of 1 Center for Systems Integrtion nd Sustinility, Deprtment of Fisheries nd Wildlife, Michign Stte University, 145 S. Hrrison Rod, Suite 115 Mnly Miles Building, Est Lnsing, Michign 48823, USA, 2 Deprtment of Geogrphy, Michign Stte University, Est Lnsing, Michign 48824, USA, 3 Stte Key Lortory of Urn nd Regionl Ecology, Reserch Center for Eco-Environmentl Sciences, Chinese Acdemy of Sciences, Beijing 185, Chin. Present ddress: Deprtment of Ecology nd Evolutionry Biology, Yle University, New Hven, Connecticut 652, USA. *e-mil: liuji@msu.edu. NATURE CLIMATE CHANGE ADVANCE ONLINE PUBLICATION Mcmilln Pulishers Limited. All rights reserved.

2 NATURE CLIMATE CHANGE DOI: 1.138/NCLIMATE1727 W N E Drgon-hed S (km) Qinling rrow Wooden Pnd hitt Nture reserve Elevtion High: 3,65 m Low: 6 m Figure 1 Loction nd topogrphy of the study re, nd the seline CSAs for the three moo species studied., The study re includes the Qinling Mountins nd surrounding res (5 5 )., The CSAs for the Qinling rrow, drgon-hed nd wooden moos were otined from the ioclimtic models under the seline climte, nd lue, red nd green colours indicte the CSAs for the three species, respectively. The mixtures of the three colours indicte overlps of the CSAs for individul species. The rightness of the colours shows the numer of ioclimtic models (mong the ten models with different presence dt prtitions) predicting pixels s suitle for ech species, with righter colours indicting lrger numer of models. drgon-hed moo overlpping etween 1,1 nd 2,3 m; Supplementry Fig. S1). In the second step, we pplied the ioclimtic models to future climte projections for three time slices during the twenty-first century (21 239, nd ) to evlute potentil temporl dynmics of the CSAs. We used the climte projections downscled to out 1 km 2 resolution from four generl circultion models (GCMs) for the Intergovernmentl Pnel on Climte Chnge (IPCC) third ssessment report (IPCC TAR) under SRES A2 nd B2 greenhouse-gs emissions scenrios (Supplementry Tle S1). All four GCMs projected wrming climte within the study re over this century, ut vritions in diurnl temperture rnge nd temperture nd precipittion sesonlity exist mong the GCM projections (Supplementry Fig. S2). We lso incorported downscled climte projections for the time slice of from 15 GCMs (Supplementry Tle S1) for the IPCC fourth ssessment report (IPCC AR4) under the A2 greenhouse-gs emissions scenrio. Although ville only for single time slice, downscled projections from dditionl GCMs etter cpture the uncertinty ssocited with future climte projections, nd, furthermore, llow for the differences in projections etween IPCC TAR nd IPCC AR4 GCMs to e ssessed. Our ioclimtic models with the climte projections from the four IPCC TAR GCMs indicte sustntil chnges in the CSAs during the twenty-first century with considerle vritions mong GCMs (Fig. 2 nd Supplementry Fig. S3). Ares in the north nd northwest of the Qinling Mountins ecome climticlly suitle for the three moo species under the climte projections from two GCMs (tht is, Commonwelth Scientific nd Industril Reserch Orgnistion (CSIRO)-Mk2 nd Hdley Centre Coupled Model, version 3 ()), especilly for the lower-elevtion species, tht is, the wooden moo (Fig. 2 nd Supplementry Fig. S3). However, within the Qinling Mountins region, our projections suggest tht the CSAs shift to, nd ecome restricted t, higher elevtions (Supplementry Fig. S1). The projected CSAs under the climte projections from the 15 AR4 GCMs lso indicte lrge GCM-relted uncertinty out the extent nd sptil distriution of CSAs of the three moo species (Supplementry Fig. S4). However, compred with the CSA projections ssocited with the TAR GCMs (Supplementry Fig. S3i l), those ssocited with the newer GCMs show higher degree of consensus s ll projections indicte reduction of CSA extent (Supplementry Fig. S4). In the finl step of the ssessment, we estimted the mount of pnd hitt on the sis of the projected CSAs of the three moo species. To cpture the uncertinty of the ility of moo species to occupy their CSAs, we used two moo dispersl scenrios representing opposite nd extreme situtions (tht is, no dispersl nd unlimited dispersl). We then otined the extent of suitle pnd hitt y comining the occupied CSAs of the three moo species under the ssumption tht the occurrence of t lest one species is sufficient for providing food nd thus constituting hitt for the pnds. Results suggest tht lmost the entire pnd hitt in the region my dispper y the end of the twenty-first century (8 1% projected decreses in medin re) if moo species cnnot colonize new CSAs eyond their present distriutionl rnges (Fig. 3,d). With unlimited moo dispersl ility, considerle mount of pnd hitt is projected to persist over the entire century, ut only if future climte is closer to the projections from two of the four GCMs ( nd ), nd then only with lower greenhouse-gs emissions scenrio (tht is, SRES B2; Fig. 3,c). In ddition, ll pnd hitt projections ssocited with the 15 IPCC AR4 GCMs show sustntil hitt losses (59 1% projected decreses in medin re), with out hlf of the projections indicting n lmost complete loss y the middle of this century under oth moo dispersl scenrios (Fig. 4). Compred with the hitt projections derived from the IPCC TAR GCMs, those ssocited with the AR4 GCMs, including those otined from newer versions of the TAR GCMs (CSIRO-Mk3. versus nd Cndin Glol Climte Model (CGCM)3.1 versus ), indicte more pronounced losses of pnd hitt (Fig. 4). This suggests tht the projected hitt expnsion ssocited with two of the older GCMs ( nd ; Fig. 3) my e overly optimistic (Fig. 4). Potentilly mking the sitution more difficult for the pnd, no increse in CSAs for the three moo species within present nture 2 NATURE CLIMATE CHANGE ADVANCE ONLINE PUBLICATION Mcmilln Pulishers Limited. All rights reserved.

3 NATURE CLIMATE CHANGE DOI: 1.138/NCLIMATE1727 LETTERS c d A2 e f g h B2 Drgon Qinling Wooden Figure 2 Projected future distriutions of CSAs in for the three moo species studied under the climte projections from four IPCC TAR GCMs (Supplementry Tle S1). h, The distriutions were projected using ioclimtic models under the SRES A2 ( d) nd B2 (e h) greenhouse-gs emissions scenrios. Blue, red nd green colours indicte the CSAs for the Qinling rrow, drgon-hed nd wooden moos, respectively. See the legend of Fig. 1 for detiled informtion on the colour representtion. For comprison, the outline of the CSAs for the seline climte for ll three species is shown in white. : coupled model from the Center for Climte System Reserch nd the Ntionl Institute for Environmentl Studies. Unlimited dispersl No dispersl A c 2 d 2 B Figure 3 Temporl dynmics of the projected chnges in the re of gint pnd hitt over the twenty-first century. The chnges (%) re reltive to the re of pnd hitt (CSAs for the three moo species studied comined) under the seline climte. d, Projections of pnd hitt were otined from the ioclimtic models for the three moo species under the unlimited- (,c) nd no- (,d) moo-dispersl scenrios using multiple future climte projections for three time slices from four IPCC TAR GCMs (Supplementry Tle S1), nd under the SRES A2 (,) nd B2 (c,d) greenhouse-gs emissions scenrios. Ten ioclimtic models were uilt for ech moo species. The points indicte the medin vlue nd the verticl rs indicte the rnge of projected vlues otined from the comintions of those models (tht is, ten ioclimtic models for the Qinling rrow moo 1 for the drgon-hed moo 1 for the wooden moo, thus 1, comintions). reserves is projected under ll model projections cross this century (Supplementry Fig. S5). This suggests tht lthough some res my ecome climticlly suitle for the moo species under chnged climte (Fig. 2 nd Supplementry Fig. S3), most of them my lie outside the present network of nture reserves. In ddition, the remining CSAs re projected to e distnt from present NATURE CLIMATE CHANGE ADVANCE ONLINE PUBLICATION Mcmilln Pulishers Limited. All rights reserved.

4 NATURE CLIMATE CHANGE DOI: 1.138/NCLIMATE IPCC TAR GCMs IPCC AR4 GCMs GISS-AOM CNRM-CM3 GFDL-CM2.1 CSIRO-Mk3. IPSL-CM4 MRI-.3.2 GFDL-CM2. FGOALS-g1. CGCM3.1 (T47) ECHO-G CGCM3.1 (T63) MIROC3. (medres) MIROC3.2 (hires) BCCR-BCM2 PCM GISS-AOM CSIRO-Mk3. CNRM-CM3 IPSL-CM4 GFDL-CM2.1 MRI-.3.2 FGOALS-g1. GFDL-CM2. CGCM3.1 (T47) CGCM3.1 (T63) ECHO-G MIROC3. (medres) MIROC3.2 (hires) BCCR-BCM2 PCM Figure 4 GCM-relted uncertinty of projected chnges in gint pnd hitt re for the time slice of under the SRES A2 greenhouse-gs emissions scenrio. The percentge chnges re reltive to the re of pnd hitt (CSAs for the three moo species studied comined) under the seline climte.,, The projections were otined from ioclimtic models for the three moo species under the unlimited- () nd no- () moo-dispersl scenrios nd under multiple future climte projections from four IPCC TAR nd 15 AR4 GCMs (Supplementry Tle S1). Ten ioclimtic models were uilt for ech moo species. Ech oxplot shows the mximum, 75th percentile, medin, 25th percentile nd minimum of the projected vlues otined from the comintions of those models. BCCR-BCM: Bjerknes Centre for Climte Reserch-Bergen Climte Model; CNRM: Centre Ntionl de Recherches Meteorologiques; ECHO-G: coupled ECHAM4 tmosphere nd HOPE ocen model; FGOALS: Flexile Glol Ocen-Atmosphere-Lnd System model; GFDL: Geophysicl Fluid Dynmics Lortory; GISS-AOM: Goddrd Institute for Spce Studies- Atmosphere Ocen Model; hires: high resolution; IPSL: Institut Pierre Simon Lplce; MIROC: The Model for Interdisciplinry Reserch on Climte; medres: medin resolution; MIR-CGCM: Meteorologicl Reserch Institute-Coupled Atmosphere Ocen Generl Circultion Model; PCM: Prllel Climte Model. moo distriutionl rnges nd pnd hitt (Fig. 2g,h). With long history of intense humn ctivities surrounding the Qinling Mountins, keeping the projected CSAs from humn disturnce, nd mintining or re-estlishing ecologicl connectivity mong those res will e difficult. This frgmented lndscpe my thus hinder the rnge shifts of moos nd pnds in response to climte chnge. As glol icon of iodiversity conservtion, the gint pnd hs ttrcted unprlleled conservtion efforts nd resources for reducing its hitt loss nd degrdtion due to lnd use/cover chnge. However, the projected loss nd shift of CSAs for the moo species during the twenty-first century indicte tht climte chnge my ecome n dditionl thret to the gint pnds, thus posing chllenge for their conservtion. As the moo species evluted constitute, t present, lmost the entire diet of gint pnds in this region 8,15, the pnds my fce shortge of food, unless they cn find lterntive food resources. Although gint pnds hve survived lrge-re die-offs of single moo species y shifting their home rnges nd forging on non-ffected moo species 8,16, they my fce regionl extinction if climte chnge, s projected y this study, induces simultneous die-offs of multiple moo species. Although the geogrphic isoltion of the study re my prevent the dispersl of moo species from other mountin regions, other species, especilly those growing t lower elevtions, might hve the potentil to occupy the res dominted t present y the three species evluted. However, lmost ll lower-elevtion species in the study re elong to the genus Phyllostchys 19, which the gint pnd t present voids consuming 2. The only other species known to constitute suitle food for pnds in this region (tht is, Frgesi nitid) 15 is distriuted t present t very low densities in smll ptches. Long-term monitoring nd further studies re needed to ssess the potentil of this species to sustitute the three dominnt moo species under chnged climte s n lterntive food resource for the pnd popultion in the study re. Like ll other ssessments of potentil climte-chnge impcts on species distriutions, the present study is limited y numer of underlying ssumptions nd uncertinty in model projections 21. The extended sexul reproduction intervls of the moo species evluted (for exmple, 7 75 yers nd out 5 yers for the wooden nd Qinling rrow moos, respectively) 22 mke niche conservtism 21 resonle ssumption. In ddition, our projection ensemle hs cptured, t lest prtilly, the uncertinty ssocited with future climte projections nd moo dispersl ility. However, the ensemle cnnot ddress ll ssumptions or cpture the full rnge of ssocited uncertinty. For exmple, our definition of gint pnd hitt is sed on two simplifying ssumptions tht my introduce dditionl uncertinty. We ssumed tht pnd hitt is determined only y the presence of moo species nd the presence of ny one of the three species evluted is sufficient to support the pnd s demnd for food. However, the presence of moo is not the only fctor determining pnd hitt. Mny other fctors, such s forest cover, humn ctivities nd pnds climtic tolernce nd dispersl ility, further limit pnd distriution 8,23. Similrly, ecuse gint pnds forge different moo species in different sesons 8, single moo species my not e le to support gint pnds yer-round. Therefore, with these ssumptions, our ssessments proly overestimte the extent of pnd hitt or underestimte pnd hitt loss due to climte chnge. In ddition, our models did not ccount for the potentil influence of overstorey tree species on moos responses to chnged climte, or cpture the potentil enefits of the relese of moo species from their nturl enemies under chnged climte 24. Nevertheless, lthough chrcteristics of the forest cnopy my lter resource vilility nd microclimte for moo species t smll sptil scles, t lrger scles climte conditions re usully the dominnt determinnt of species distriution 25. Furthermore, lthough the moo popultion my enefit from enemy relese, these effects on the gint pnd my e very limited ecuse pnds do not strongly compete for moo with other nturl enemies of moo species 8. 4 NATURE CLIMATE CHANGE ADVANCE ONLINE PUBLICATION Mcmilln Pulishers Limited. All rights reserved.

5 NATURE CLIMATE CHANGE DOI: 1.138/NCLIMATE1727 Mny studies hve oserved or projected climte-chngeinduced distriutionl shifts or extinction of individul species 1,2,26. However, the cscding effects of these chnges through interspecific interctions, especilly those cross trophic levels, re usully neglected 27. This my result in n underestimtion of the ecologicl impcts of climte chnge, thus compromising the enefits otined from present iodiversity conservtion efforts. This study underscores the importnce of incorporting these cscding effects not only in impct ssessments ut lso in conservtion plnning in the fce of climte chnge. Methods We collected moo-presence dt in 293 field plots covering the elevtionl rnge of the distriutions of three dominnt moo species within the Qinling Mountins, which form the northern limit of the present gint pnd distriutionl rnge (out 21,3 km 2 ). We used gridded ioclimtic vriles (3 3 rc s), which were sed on wether sttion records from 195 to 2, otined from the WorldClim dtse 18 to represent the seline climte in the study re. Using the mximum entropy lgorithm 28, we generted ioclimtic models to mp seline CSAs for ech moo species sed on the presence dt nd seven ioclimtic vriles (Supplementry Tle S2) tht re iologiclly nd sttisticlly importnt for chrcterizing the three moo species CSAs (Supplementry Methods). To incorporte the uncertinty introduced y potentil smpling is, we generted 1 ioclimtic models for ech species with different dt prtitions, ech of which contined rndomly selected 7% of the moo presence dt. The remining 3% were used for vlidting the models nd determining thresholds (corresponding to 1% omission error) to convert the continuous model outputs (presence proility) into inry vlues (suitle or unsuitle). We then projected the sptil distriution of the CSAs over the twenty-first century y pplying the ioclimtic models to future climte projections tht were sttisticlly downscled to 3 3 rc s resolution from 19 GCMs (Supplementry Tle S1) sed on the sptil interpoltion of nomlies etween 3-yer verges of temperture nd precipittion for future nd present-dy simultions 29. We otined the portion of CSAs occupied y ech moo species under two dispersl scenrios. With the unlimited-dispersl scenrio moo species cn occupy ll CSAs, wheres with the no-dispersl scenrio it cnnot occupy the res eyond its seline CSAs. Although neither of these two scenrios reflects the rel dispersl ility of the moo species, they cpture the rnge of uncertinty in terms of moo dispersl ility. Finlly, we comined the occupied CSAs of the three moo species to define suitle pnd hitt nd clculted the percentge chnge of the re from the seline vlues for every comintion of the CSA projections for the three moo species (1 1 1 ioclimtic models 2 dispersl scenrios) under multiple future climte projections (4 (or 15) GCMs 2 (or 1) greenhouse-gs emissions scenrios). See Supplementry Informtion for detiled methods nd ssocited references. Received 23 Jnury 212; ccepted 1 Octoer 212; pulished online 11 Novemer 212 References 1. IPCC Climte Chnge 27: Impcts, Adpttion nd Vulnerility (eds Prry, M. L., Cnzini, O. F., Plutikof, J. P., vn der Linden, P. J. & Hnson, C. E.) (Cmridge Univ. Press, 27). 2. Prmesn, C. Ecologicl nd evolutionry responses to recent climte chnge. Annu. Rev. Ecol. Evol. Syst. 37, (26). 3. Gillim, F. S. The ecologicl significnce of the herceous lyer in temperte forest ecosystems. Bioscience 57, (27). 4. Nilsson, M-C. & Wrdle, D. A. 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Rpid rnge shifts of species ssocited with high levels of climte wrming. Science 333, (211). 27. Vn der Putten, W. H., Mcel, M. & Visser, M. E. Predicting species distriution nd undnce responses to climte chnge: Why it is essentil to include iotic interctions cross trophic levels. Phil. Trns. R. Soc. B 365, (21). 28. Phillips, S. J., Anderson, R. P. & Schpire, R. E. Mximum entropy modeling of species geogrphic distriutions. Ecol. Model. 19, (26). 29. Rmirez, J. & Jrvis, A. High Resolution Sttisticlly Downscled Future Climte Surfces (The Interntionl Centre for Tropicl Agriculture, 28). Acknowledgements We thnk G. Dng, X. Du, J. Meng nd Y. Wng for their invlule ssistnce during field cmpigns. We lso thnk the Forestry Deprtment of Shnxi Province nd dministrtions of nture reserves in the Qinling Mountins for support with fieldwork logistics. This study ws supported y the Ntionl Aeronutics nd Spce Administrtion (Terrestril Ecology nd Biodiversity progrmme nd the Erth nd Spce Science Fellowship progrmme) nd the US Ntionl Science Foundtion (Dynmics of Coupled Nturl nd Humn Systems progrmme nd Prtnership for Interntionl Reserch nd Eduction). Author contriutions M-N.T., A.V., J.A.W nd J.L. conceived the ides nd designed the nlyses; M-N.T., A.V., Y.L., W.X. nd Z.O. collected the dt; M-N.T. uilt the models nd performed the dt nlyses; ll uthors wrote the pper. Additionl informtion Supplementry informtion is ville in the online version of the pper. Reprints nd permissions informtion is ville online t Correspondence nd requests for mterils should e ddressed to J.L. Competing finncil interests The uthors declre no competing finncil interests. NATURE CLIMATE CHANGE ADVANCE ONLINE PUBLICATION Mcmilln Pulishers Limited. All rights reserved.