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No insurance policy, necessarily                                                 mean  annual  precipitation  fell
        Recent data suggest that at a continental scale, those regions in Africa rich in tropical plant species (13) are also the most   below 1500 mm  in  evergreen
        sensitive to climate variability (6), implying that higher species richness does not necessarily lead to greater resilience.  tropical  forests  in  South  Amer-
                                                                                         ica,  Africa,  and  Southeast  Asia,
                                                                                         possibly indicating  a  tipping
           Vegetation sensitivity to climate variability    Plant species richness
                                                                                         point  about  to be  crossed.  By
                                                                                         contrast,  another  study  in the
                                                                                         Amazon rain  forests  found  only
                                                                                         evidence  of  gradual change  to
                                                                                         several  transitional  forest  states
             Sensitivity                                                                 in  response  to a lengthening  of
             index                                    Richness                           the  dry season  (10), rather  than
               100                                      ≥500                             an abrupt change from forest to
                                                        200–500                          another state.
                                                        80–200                            There  is  also  the fundamen-
               50                                                                        tal question of whether switch-
                                                        30–80
                                                                                         ing  between  alternative states
                                                        12–30
                                        0  800                                  0  800   is always necessarily a  bad
                                                        <12
               0                        km                                      km       thing.  Recent  studies  indicate
                                                                                         that  tropical  grasslands  persist
        disturbance  apparently  can also  influence  According to another biotic hypothesis, it   in a permanent transitional state and that
        the resilience of an ecosystem, and this varies   is the characteristics of the component spe-  the ability to switch between forest and sa-
        according to vegetation type. For example, in   cies (such as wood density,  rooting  depth,  vanna in response to perturbations under-
        tropical grasslands such as those in West Af-  and leaf-area index) that make ecosystems   pins their resilience (9).   Downloaded from
        rica, disturbance by fires buffers ecosystems   more resilient (1). Here, some clear trends   The  recent  studies  discussed  above  are
        from forest encroachment and thus promotes   are starting to emerge. For example, Green-  starting  to  test the  many  hypotheses  that
        grassland resilience (8). By contrast, frequent   wood et al. (14) found that across forested   exist  to explain  resilience  in terrestrial  eco-
        disturbance by fires in tropical forest-savanna   biomes, mortality rates after drought were   systems. They  reaffirm  the  complexity  of
        transition zones can lead to loss of resilience   lower for species with greater wood den-  resilience but also provide clear pointers for
        in forest communities (9).          sity  and  lower  specific  leaf  area.  A  global  future research and conservation. In tropical
          Another abiotic attribute hypothesized to   meta-analysis also identified  these  two  ecosystems, soil type, belowground processes,
        account for ecosystem resilience is soil type.   characteristics as important for withstand-  and rooting depth are potentially important   http://science.sciencemag.org/
        Again, there is some evidence  to  support  ing drought in tropical rain forests, whereas   areas of future research with direct manage-
        this. In Poorter et al.’s study, high soil fertil-  in tropical grasslands,  plants  with  deeper  ment  applications.  The  factors responsible
        ity had a positive influence on biomass re-  roots were more resilient to drought (3).  for  resilience  of  tropical  grasslands  are  an-
        covery in Neotropical secondary forest plots                            other knowledge gap needing more research.
        (5). Similarly, a modeling study that incor-  HOW CLOSE IS A SYSTEM TO   Given  the importance  of  terrestrial  ecosys-
        porated remote sensing and field data pre-  LOSING RESILIENCE?          tem  resilience to  natural  resource security
        dicts that rain forest situated on soils with   Determining  which biotic  and abiotic  fac-  and supply across the globe, research into the
        low clay  content  will  be  least  affected  by  tors contribute to resilient ecosystems is  attributes underpinning it should be high on   on March 1, 2018
        an increase in the length of the dry season   important for maintaining and enhanc-  any international agenda. j
        and will thus have  higher  resilience  (10).   ing them. However,  when  determining
                                                                                REFERENCES
        Belowground biotic attributes  may  also  be  conservation  strategies,  it  is  also  critically
                                                                                  1.   D. Hodgson, J. L. McDonald, D. J. Hosken, Trends Ecol. Evol.
        important in determining the resilience of   important to be able to identify  when  an  30, 503 (2015).
        an ecosystem; in particular, plants that have   ecosystem is about to lose its resilience and     2.   B. L. Timpane-Padgham, T. Beechie, T. Klinger, PLOS ONE
        root systems associated with mycorrhizal  cross a threshold from a desirable to an un-  12, e0173812 (2017).
                                                                                  3.   K. J. Willis et al., in State of the World’s Plants, 2017 Report,
        fungi may have greater resilience to water   desirable stable state. Several methods have
                                                                                   K. J. Willis, Ed. (Royal Botanic Gardens, Kew, 2017),
        stress in tropical dry forests (11).  been proposed to do this. For example,  at  pp. 42–49.
          There  is  also  a  suite  of  biotic  factors to  the continental scale, Hirota et al. (15) have     4.   L. E. S. Cole, S. A. Bhagwat, K. J. Willis, Nat. Commun.5,
                                                                                   3906 (2014).
        consider.  Possibly  the  most  widely  cited is  shown that tropical and subtropical ecosys-
                                                                                  5.   L. Poorter et al., Nature530, 211 (2016).
        the insurance hypothesis. This suggests that   tems in Africa, Australia, and South Amer-    6.   A. W. R. Seddon, M. Macias-Fauria, P. R. Long, D. Benz,
        more  biodiverse ecosystems  will  be more  ica switch to a  savanna  state  when  forest  K. J. Willis, Nature531, 229 (2016).
                                                                                  7.   J. Verbesselt et al., Nat. Clim. Change6, 1028 (2016).
        resilient  to  environmental  perturbations  cover is less than 60%. This has direct im-
                                                                                  8.   P. Laris, S. Dadashi, A. Jo, S. Wechsler, Plant Ecol.217, 583
        because  they contain  a  greater  number  of  plications for the management of tropical  (2016).
        species available to replace functions carried   forests, where deforestation is a huge issue.     9.   I. Oliveras, Y. Malhi, Philos. Trans. R. Soc. B Biol. Sci.371,
        out by lost species. This certainly appears to   Another  proposed  method  is  to  examine  20150308 (2016).
                                                                                  10.   N. M. Levine et al., Proc. Natl. Acad. Sci. U.S.A.113, 793
        be the case at the community level in some   recovery rates from disturbances on the basis   (2016).
        ecosystems (12)  but  does  not  necessarily  of the hypothesis that the closer a system is     11.   K. Allen et al., Environ. Res. Lett.12, 023001 (2017).
                                                                                  12.   M. Hisano, E. B. Searle, H. Y. H. Chen, Biol. Rev.93, 439 (2018).
                                                                   recovery
        hold
                                                                          rate
                                                                the
                                                           slower
                 continental
      GRAPHIC: J. YOU/SCIENCE  regions with the highest tropical plant spe-  will be (1). This approach appears to work in     14.   S. Greenwood et al., Ecol. Lett.20, 539 (2017).
                                   example,
                                            to a threshold,
                           scale. For
                                                        the
            at the
                                                                                  13.   H. P. Linder, Front. Ecol. Evol. 10.3389/fevo.2014.00038
                                                                                   (2014).
        cies richness in Africa (see the figure, right)
                                            models but only seems to hold true for some
                                                                                  15.   M. Hirota, M. Holmgren, E. H. Van Nes, M. Scheffer, Science
                                 to climate
            appear
                 to be
                      most
                          sensitive
        (13)
                                                                example, Verbes-
                                                    ecosystems. For
                                            terrestrial
                                                                                   334, 232 (2011).
        perturbations
                   (see
                                   (6)—the
                                            selt et al. (7) found that recovery rates from
                               left)
                       the
                          figure,
        opposite finding to the insurance hypothesis.
                                                                                                 10.1126/science.aar5439
        SCIENCE  sciencemag.org             perturbations  slowed  down  sharply  once  2 MARCH 2018 • VOL 359 ISSUE 6379    989
                                                       Published by AAAS
   DA_0302Perspectives.indd   989                                                                            2/28/18   11:03 AM
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