Mountain Ecology: Organism Responses to Environmental Change, an Introduction Author(s): John R.

Haslett Reviewed work(s): Source: Global Ecology and Biogeography Letters, Vol. 6, No. 1, Mountain Ecology: Organism Responses to Environmental Change (Jan., 1997), pp. 3-6 Published by: Blackwell Publishing Stable URL: http://www.jstor.org/stable/2997522 . Accessed: 18/01/2012 07:34
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Letters1997) 6, 3-6 ( Global Ecology Biogeography and

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to responses environmental organism ecology: Mountain an change, introduction

JOHN R. HASLETT Austria 34, of of Institute Zoology,University Salzburg,Hellbrunnerstrasse A-5020 Salzburg,

climate Key words. Mountain habitats,landscape ecology,spatial and temporalscale, island biogeography, change,humanculturaltheory.

INTRODUCTION
spatial by Mountainsare characterized theirextreme caused by the heterogeneity. Quite apartfrom patterns topographyand ground cover altitudinalgradients, over a wide range of spatial scales to create interact These complex,nestedmosaics of habitatconditions. effects upon mosaics,in turn,oftenhave considerable on as the spatialdistributions organisms mountains of well as on their morphologies, growth,physiology, life cycles, behavioural patternsand the types and Such a betweenindividuals. intensities interactions of and highdegreeof spatialheterogeneity itsdominating ('alpine') influences majorwaysin whichmountain are environments to be consideredunique, set apart are from the equally harsh, but generally more homogeneous polar ('arctic') areas (Haslett,1996a). This 'landscape' approach to mountainecology is underpinned an awareness of the importanceof by scale-the size at whichtheland mosaic is interpreted. Traditionally, mountainshave been treatedsimplyas large scale landscapes-mosaics of habitatsand land of use thatreflect humanperceptions theenvironment. This way of thinking has been perpetuated some to degree by those using modern Geographical InformationSystemsat larger scales (e.g. Price & Heywood, 1994). An alternative is to consider mountain(or other)land mosaics fromthe 'organism point of view'. Here a wide range of spatial scales is to of included,frommillimetres hundreds kilometres, beetlesor birdsare thefocus depending upon whether of attention (e.g. Wiens, 1995; Haslett, 1996b). This multi-scale approach allows understanding of ecologicalprocesses(see Hansson, Fahrig& Merriam,
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than merely reflecting wishesof land the 1995) rather managersand planners(Haslett, 1996b). The papers in this Special Issue of Global Ecology clearlythe role Lettersdemonstrate and Biogeography in of spatial heterogeneity creatingthe uniqueness

of environments the communities and of mountain

thatinhabitthem,whether theybe plants, organisms Theyalso insects, smallmammalsor indeedourselves. emphasize the importanceof scale in any attempt how organisms,from individualsto at determining are populationsto communities, likelyto respondto changesto mountainenvironments.

OF THE VULNERABILITY MOUNTAIN ENVIRONMENTS: EVIDENCE FROM CONFLICTING PLANTS AND ANIMALS?
to are mountain environments subject change Although of from widevariety causes,bothnaturaland human a (Hewitt,1995), changesrelatedto a changingclimate a are presently major avenue of researchand are an themein the papers of this Special Issue. underlying consensusthat the There is now a generalscientific is to global climate likely changeovercomingdecades, in (e.g. at ratesunprecedented modernhumanhistory 1989). Mountainclimatesare characterized Schneider, by marked diurnal and seasonal cycles, with high overa wide rangeof spatialscales. Changes variability in to these climatesmay affect mountainecosystems two general ways, although these are also closely responses maybe physiological First, there interlinked. of arisingfrom the increased concentration carbon dioxide available for photosynthesis (Korner, 1994; 3

4 lohn Haslett R.
Price& Haslett,1995). Second,there maybe responses of organisms thephysical to climatic changes:changes in the means, variances and spatial distributions of and light parameterssuch as temperature, humidity Both of these sets of issues have important intensity. for at consequences mountain organisms a widevariety of spatial scales and organisational levels. Responsesofmountain plantsto changesin climate, or indeed to variations in any aspect of the environment, of fundamental are because importance, alterations the formand/orcontentof vegetation to mosaics also constitute changesto habitatmosaics for both plantsand animals.In a studyof the vegetation of theOlympicMountains(USA), Peterson, Schreiner & Buckingham (1997) use palaeontological and biogeographicaldata to identifyresponses of the vegetation previousclimaticchanges,and use this to to as information gainan insight to how thevegetation will react to futureclimates. They argue that the is diversity habitatsin mountains a major sourceof of resilience climatechange and that the exceptional to mountain ecosystemsin the face of environmental change. This is despite the fact that animals are generallymuch more mobile than plants, so that intuitively animalpopulationsmight expected be be to morerobust. Boggs& Murphy (1997),usingbutterflies, show that althoughthe morevagile mountainspecies have the abilityto move theirdistributions, tracking the changes in vegetationand conditionscaused by global warming, less mobile speciesare at riskof the in extinction such circumstances. Importantly, these authorsnote that changesin the variance(alterations not to the frequency occurrence), just themeans of of to cause the climatic conditions, are sufficient population extinctions. A similar argument is developedby Haslett (1997) in relationto fliesof the in of family Syrphidae theAlps. Here,numbers species and individualshave been correlatedwith the form and complexityof the habitat mosaic (taken as a dynamicentityof its own, not just a collectionof patches) at relevantspatial scales. Changes to the microclimatic mosaic,particularly degreeofmixing the betweenpatches,are likelyto have significant effects on populationsizes and speciesassemblagestructure. One of themostwidespread consequencesof global is warming on mountain environments an overall reductionin area, with the total loss of particular habitats. As the different altitudinalzones migrate habitats reduced are the upwards, 'islands'ofmountain in size, and the uppermostsimplydisappear when thereis no longer any land surfacehigh enough to to accommodatethem.This leads ultimately changes in population densities,species ranges and regional species distribution patterns of biogeographic as dimensions and significance, illustrated thestudy by of North Americanbutterflies Boggs & Murphy by of (1997). Indeed, the characteristics mountainsand mountainranges can be sufficiently insularto allow to speciesdistributions be analysedusingthe theories and ideas of island biogeography.In an island biogeographicalstudy of the non-volantmammals in in NorthAmerica, occurring themountains western Lomolino & Davis (1997) suggestthat our abilityto and assess causalityin biogeography perceive patterns may be stronglyinfluencedby our own limited perceptionof spatial scale (here intra-archipelago to large-scaleregional) in combinationwith the use of different parameters ecological description of (species richness, community nestedness and distribution of patterns singlespecies).One important implication is that a limitedperceptionof scale is also likelyto affect understanding extinction of our rateson islands,

within small variety habitat of conditions relatively

distances facilitates survival of plant species by a mosaicthatusuallyincludessome providing dynamic 'safe sites'. These sites are the sources of later recolonization. Going a stagefurther, authorsalso the make the pointthatmostforms humanland use in of mountain environmentstend to reduce spatial heterogeneity and thus also reduce the inherent resilience the system. of Some support thisviewis provided theresults for by of a studyof the forest typesof the Swiss Alps. This workemploysa verydifferent, indeedalmostopposite approach to the previousone. Fischlin& Gyalistras (1997) take Global Climate Models as theirstarting the to point,down-scaling predictions local levels on in real mountain terrains pre-defined climatescenarios that also include seasonal changes.They have found thatup to a certain limittheir model predicts thatthe forestsare robust in the face of changingclimatic parameters. Indeed, at one of theirsites,even major to changes did not resultin any greatalteration the forest type.However,results further of simulations at other sites do suggestthat a whole range of forest responses is possible, from no change to complete collapse oftheforest canopy,depending upon thelocal situation. to Contrary the above vegetational studies, two the includedin thisSpecial Issue thatfocus investigations is upon insectssuggestthat spatial heterogeneity an factorthat contributes the fragility to important of

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Mountain introduction 5 ecology,

and therefore also our judgementof the vulnerability or robustness mountain of habitat'islands'in response to environmental change. (Note that the botanical paper by Petersonet al. (1997), in whichmountains were argued to be resilient,involves a parallel the methodology; use of information based upon the past as an aid to explaining present future and patterns of speciesdistributions). Given the evidence presentedin the papers cited it above, and including bothplantand animalstudies, maybe suggested thattoo muchreliance upon 'human scales of study' is the main source of many of the present uncertainties the surrounding degreeof impact of environmental changeon mountainecosystems. linearprogressions, systems the respond beingpulled to a in many different ways at once by exhibiting high in degreeof flexibility, whichstrategies may changeto of allow the systemto absorb the effects unforeseen 'surprise' events. putforward Price& Thompson As by (1997), the similarities between culturaltheoryand ecological theoryon mountainsare striking, and the approach shouldnot be ignored.

TAILPIECE
to This brief introductory paperhas attempted provide a perspective some of the critical of issuesinvolvedin understanding plant and animal responsesto changes in mountain environments, particularly changes in climate. Spatial complexityand scale have come the of to with dynamic nature strongly thefore, together scales.Whether spatialmosaicsoverdifferent temporal 'top down' or 'bottom up' researchapproaches to ecological questionsof climatechange are preferable remains subjectfordebate(Root & Schneider, a 1995), but it may be suggestedthat because of the wide a rangeof spatialand temporalscales involved, closer integration the two types of strategy of would be in particularly fruitful mountainsituations. Mountain ecosystemsare argued here to have importance special, functional as componentsof our planet, and also to havevalue as 'model'systems within the realms of ecological and biogeographical theory. This meanstheyare worthpreserving. Despite thefact that many mountainareas have been designatedas otherforms of National Parks,or have been afforded protectedstatus, the presently accepted large-scale, patchcontent approachesto conservation management are likely to prove inappropriatein situations of environmental change. If the characteristic 'spatial of is thenit biodiversity' mountains to be maintained, and will be necessary alter land use, management to to conservation strategies place much more emphasis on the importance complexity of and scale-the very aspects of mountain ecology that make mountains unique.

HUMAN INTERPRETATIONS: MOUNTAIN ECOLOGY MEETS CULTURAL THEORY

If it is necessary be waryof overlyanthropogenic to it interpretations naturein mountainsituations, is of on ironicthatecosystem certainly dynamics mountains seem to have a lot in common with human socioin culturalsystems the same areas. Classical views of both types of system assume some unidirectional this changefroman initialto a finalstate.In ecology, is exemplified successionalprogressions towardsa by 'climax'community, whilein the social scienceslinear to changesfromtraditional modern,or capitalismto are examples of 'grand theories' that communism, infer changein a singledirection. Recent experience has shown that in ecology, rather it successionis not a simplelinearprogression, is a complex,dynamic,and oftencyclic,systemof mosaic patches (e.g. Remmert,1991) verysimilarto those of mountainlandscapes discussed withinthis modernculturaltheory views Special Issue. Similarly, social systemsas non-linear, involvingnever ending sequencesof transitional states(e.g. Holling, 1986). In an explorationof the parallels and interactions between ecology and human culture, Price & Thompson(1997) examinetheresponses thehuman of inhabitants of mountain regions to the changing in environments whichtheylive,as shownby theways in whichthey'manage'their resources. Usingexamples ofa Himalayanand a Swissalpinevillage, theseauthors argue that mountain socio-culturalsystemscannot operate as straightforwardly implied by 'simple' as theories such as presentedin 'the tragedy of the commons' (Hardin, 1968, 1994). Instead of simple

ACKNOWLEDGMENTS
I thankJohnLee and Hazel Normanfortheir friendly support before and during the VI International CongressofEcologyin Manchester, wheretheauthors of thismountainecologySpecial Issue came together.

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R. 6 John Haslett
Reinhard Bornkamm,Pehr Enckell and Paul Giller the have been of greathelp in getting EEF Specialist Group on High AltitudeEcology offthe ground.My sincerethanks also to all the authors and to those collegueswho acted as referees. ecosystems: of C.S. Holling, (1986)Theresilience terrestrial local surprise and global change. Sustainable (ed. development thebiosphere by W. C. Clarkand of Press, University R.E. Munn),pp. 292-320.Cambridge Cambridge. on changes high Korner, (1994)Impactofatmospheric C. in changing Mountain environments altitude vegetation. London. Routledge, climates by M. Beniston). (ed. scale M.V. & Davis, R. (1997) Biogeographic Lomolino, of forest mammals western andbiodiversitymountain of 6, GlobalEcol. Biogeogr. Letts, 57-76. NorthAmerica. N.M. E.G. & Buckingham, Peterson, D.L., Schreiner, Spatial and vegetation climate: and (1997) Gradients, Ecol. Biogeogr. Global in temporal dynamics mountains. Letts, 7-17. 6, Price,M.F. & Haslett,J.R. (1995) Climatechangeand issues Mountains risk:current at mountain ecosystems. in environmental studies by J.R.Allan),pp. 73-97. (ed. Manohar, New Delhi. Price, M.F. & Heywood,D.I. (eds) (1994) Mountain 309 environmentsgeographic and information systems, pp. & London. Taylor Francis, M. Price,M.F. & Thompson, (1997) The complexlife: EcoL Global ecosystems. Humanlandusesin mountain 6, Biogeogr Letts, 77-90. H. concept of Remmert, (ed.) (1991) The mosaic-cycle Heidelberg. ecosystems, 168pp. Springer-Verlag, 85). (Ecologicalstudies S.H. Root,T.L. & Schneider, (1995)Ecologyand climate: Science, 269, and implications. Research strategies 334-341. effect: science and S.H. Schneider, (1989)The greenhouse 243,771-781. policy. Science, Wiens,J.A. (1995) Landscape mosaics and ecological and Mosaic landscapes ecological processes (ed. theory. pp. by L. Hansson,L. Fahrigand G. Merriam), 1-26. Chapmanand Hall, London.

REFERENCES
Boggs, C.L. & Murphy,D.D. (1997) Community composition in mountain ecosystems: climatic Global butterfly distributions. determinantsmontane of Ecol. Biogeogr. Letts.6, 39-48. of D. impacts Fischlin, & Gyalistras, (1997)Assessing A. in climatic changeon forests the Alps. GlobalEcol. Biogeogr. Letts.6, 19-37. L. G. Hansson,L., Fahrig, & Merriam, (eds)(1995)Mosaic 356 processes, pp. Chapman landscapes and ecological and Hall, London. G. of Science, Hardin, (1968) The tragedy thecommons. 162, 1243-1248. Hardin, G. (1994) The tragedyof the unmanaged Ecol. Evol.9, 199. commons. Trends & biodiversity: patterns, Haslett, (1996a)Arctic alpine J.R. 1995.(ed. consequences. Springer, causesand ecological 379, 688 by F.S. Chapin,III and C. Korner).Nature, (book review). J.R. ecology diversifies. Trends Haslett, (1996b)Landscape Ecol. Evol.11,521-522(book reveiw). and Haslett,J.R. (1997) Insectcommunities the spatial Ecol. Biogeogr. Global of habitats. complexity mountain Letts, 49-56. 6, Hewitt,K. (1995) Hazards and disasterin mountain of environments: Problemsin the geography risk. in studies at issues environmental Mountains risk:current New Delhi. (ed. byJ.R.Allan),pp.98-128.Manohar,

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