Geoscience Reference
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as.they.respond.to.changes.in.climate.(but.see,.e.g.,.Smith.et.al..2000;.Russell.et.al..
2009)..In.some.instances,.one.or.more.aspects.of.a.species'.synecology.will.be.the.
means. by. which. climatic. inluence. may. either. expand. or. contract. a. species'. local.
(or. regional). distribution,. depending. on. whether. the. changed. climate. ultimately.
weakens. or. strengthens. the. species. in. its. interactions.. Examples. of. synecological.
interactions.include.disease.or.parasitism,.predation.pressure,.and.competition.from.
other.species.that.may.also.be.undergoing.changes.in.abundance.or.distribution.in.
response. to. climate. change. (LaVal. 2004;. Pounds. et. al.. 2006).. In. other. instances,.
synecological.factors.will.interact.in.concert.with.direct.(e.g.,.physiological).effects.
of.climate.on.organisms..For.many.species,.it.remains.unknown.what.combinations.
of. physiology,. diet,. water. stress,. symbiotic. relationships,. and. other. stressors. (e.g.,.
ire,.invasive.species,.harvest).produce.losses.in.distribution..More.rare.yet.is.quan-
tiication.of.the.relative.contributions.of.these.various.factors.in.various.geographic.
and.topographic.contexts..As.with.many.biological.phenomena,.synergy.with.other.
factors.appears.likely.to.be.the.norm.(Murphy.and.Weiss.1992;.Thomas.et.al..2004;.
Hulme.2005;.Root.and.Schneider.2006;.Parmesan.et.al..2011)..Changes.in.land.use.
and. resultant. landscape. fragmentation,. as. well. as. invasive. species. appear. poised.
to.be.factors.that.commonly.interact.with.climate.(Noss.2001;.Thomas.et.al..2004;.
Wilson.et.al..2005).
Although.there.have.been.many.predictions.of.which.life-history.traits.may.make.
individual. species. more. or. less. vulnerable. to. relatively. rapid. changes. in. climate,.
there.remains.a.need.to.broadly.and.rigorously.test.the.generalities.that.have.been.
postulated.(Keith.et.al..2008)..For.example,.there.still.remains.a.need.to.ind.traits,.
markers,. and. even. proxies. for. predicting. not. only. which. species,. but. also. which.
populations. within. a. species. may. be. most. vulnerable. to. climate-induced. losses.
(Beever.et.al..2012)..The.domain.of.vulnerability.assessment.has.already.spawned.
an.ever-enlarging.literature.on.species,.guilds,.and.regions.predicted.to.be.at.greater.
risk;.however,.relatively.few.such.assessments.have.been.quantitatively.validated.or.
veriied..Although.some.traits.are.intuitively.obvious.predictors.of.risk.(e.g.,.limited.
dispersal.capacity.and.low.reproductive.capacity.[Thomas.et.al..2004]),.other.rela-
tionships.may.yield.more.surprises..For.mountain-dwelling.vertebrates,.for.example,.
persistence.within.a.particular.mountain.range.might.relect.any.combination.of.hab-
itat-diversity.and.climatic.surrogates:.total.area.of.habitat.present,.topographic.diver-
sity. (e.g.,. availability. of. different. aspects),. presence. of. cold-air. pooling,. elevation.
of. the. highest. peak,. extent. of. perennial. lowing. water,. abundance. of. rock-ice. fea-
tures,.number.of.species.that.provide.habitat.structure.(e.g.,.conifers),.or.number.of.
major.habitat.types.(Johnson.1978;.Brown.1978;.Murphy.and.Weiss.1992;.Millar.and.
Westfall.2010)..Future.challenges.will.include.not.only.understanding.why.patterns.
of.loss.of.some.species.unexpectedly.depart.from.predictions.of.ecological.theory.
(e.g.,. “holes”. in. nestedness. analyses),. but. also. why. other. species. are. surprisingly.
resilient.. Even. for. demonstrated. relationships. between. organisms. and. climate,. we.
generally. do. not. know. where. the. “tipping. points,”. thresholds,. and. nonlinearities.
lie,. in. terms. of. climatic. tolerance—especially. when. multiple. factors. are. involved.
(even.if.all.are.related.to.climate)..From.the.opposite.side.of.the.coin,.it.is.virtually.
untested.which.species.(if.any).will.be.most.effectively.aided.by.assisted.migration,.
assuming.that.all.of.the.attendant.concerns.are.handled.satisfactorily.(Richardson.et.
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