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Ecological Mechanisms Linking Protected Areas to Surrounding Lands

Author(s): Andrew J. Hansen and Ruth DefriesReviewed work(s):Source: Ecological Applications, Vol. 17, No. 4 (Jun., 2007), pp. 974-988Published by: Ecological Society of AmericaStable URL: http://www.jstor.org/stable/40061891 .

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ECOLOGICAL MECHANISMS

LINKING PROTECTED

AREAS

TO SURROUNDING LANDS

Andrew

J. Hansen1'3

and Ruth DeFries2

EcologyDepartment,

ontana

State

University,

ozeman,

Montana

59717-3460

USA

1

Departmentf Geography

nd Earth

Systems

cience

nterdisciplinary

enter,

181

Lefrak

Hall,

University

f

Maryland,

ollege

Park,

Maryland

0742 USA

Abstract. Land

use is

expanding

nd

intensifying

n the

unprotected

ands

surrounding

many

of the world's

protected

reas. The influence f

this

and

use

change

on

ecological

processes

s

poorly

understood. he

goal

of this

paper

is to draw on

ecological

theory

o

provide synthetic

rameworkor

understanding

ow and

use

change

round

protected

reas

may

alter

cologicalprocesses

nd

biodiversity

ithin

rotected

reas and to

provide

basis

for

dentifyingcientifically

ased

management

lternatives.We

first

resent conceptual

model of

protected

reas embeddedwithin

arger

cosystems

hatoften

nclude

urrounding

human land use. Drawing on case studies n this InvitedFeature,we then explorea

comprehensive

et

of

ecological

mechanisms

y

which and use

on

surrounding

ands

may

influence

cological processes

nd

biodiversity

ithin eserves. hese mechanisms

nvolve

changes

n

ecosystem

ize,

with

mplications

orminimum

ynamic

rea,

species-area

ffect,

and

trophic

tructure;

lteredflows f materials nd disturbancesnto and out of

reserves;

effects n crucialhabitats or easonal and

migration

movementsnd

population

ource/sink

dynamics;

nd

exposure

o humans

hrough unting, oaching,

xotics

pecies,

nd disease.

These

ecological

mechanisms

rovide

basis for

ssessing

he

vulnerability

f

protected

reas

to land use.

They

lso

suggest

riteria or

designing

egionalmanagement

o sustain

rotected

areas n the ontext f

surrounding

uman

and

use. These

design

riterianclude

maximizing

the

area of functional

habitats,

dentifying

nd

maintaining cological process

zones,

maintaining ey

migration

nd source

habitats,

nd

managing

human

proximity

nd

edge

effects.

Key

words:

ecological

rocesses;

cosystem

ize;

edge effects;

abitat;

and

use

change;management;

protectedreas; vulnerability.

974 INVITED FEATURE

^°l0*C

Vd^No™

Ecological

Applications,

(4),

2007,

pp.

974-988

© 2007

by

the the

Ecological

Society

f America

Introduction

Human

societieshave

long

set

aside tracts

f

land

to

conserve

ature

n theform

f

hunting

eserves,

eligious

forests,

nd common

grounds

Chandrashekara

nd

Sankar

1998).

The current

oncept

of national

parks

evolved n the

mid 1800s as

European

colonistswere

converting

ative

landscapes

to

farms,ranches,

and

cities

Schullery

997).

A

keygoal

was the

protection

f

nature.

By

minimizing

he nfluence f

humans,

natural

ecosystems

were

expected

to continue

to

maintain

ecologicalprocesses

nd native

pecies.

During

the 20th

century, rotected

reas became a

cornerstone f the

global

conservation

trategy.

New

protected

reas

continue

to be established: he total

number

lobally

has doubled since 1975

Ervin

2003a).

The term

protected

rea refers o

any

area of and or

sea

managed

for

the

persistence

f

biodiversity

nd

othernatural

processes

n

situ,

through

onstraints

n

incompatible

and uses

(Possingham

t al.

2006).

The

basic role

of

protected

reas

is to

separate

lements

f

biodiversity

rom

rocesses

hat hreaten

heir

xistence

in

the wild

(Margules

and

Pressey

2000).

Recent

assessments

have found that

most terrestrialeserves

are

adequately protected

within heir

borders

Bruner

et al.

2001,

DeFries

et al.

2005).

Despite

the

high

evel

of

protection

fforded

ational

parks

and

other

protected

areas,

many

are

not

functioning

s

originally

nvisioned.

ritical

cological

processes uch s fire,looding,

nd

climate

egimes

ave

been altered

Lawton

et al.

2001,

Pringle

001).

Exotic

species

are

increasingly

nvading

protected

areas

(Stohlgren

1998),

and

some native

species

have

gone

extinct

n

protected

reas

(Newmark

1987, 1995,

1996,

Rivard t

al.

2000,

Brashares

t al.

2001).

For

example,

11 of

13

national

parks

in the western

United States

have lost

large

mammal

species

since

park

establish-

ment,

with 5-21.4% of

original

pecies

ost

Parks

and

Harcourt

002).

Why

are

manyprotected

reas

not

functioning

ell,

despite dequate

management

within heir

borders?

A

major

reason

may

be thathuman

anduse is

expanding

Manuscript

eceived

August

005;

revised 2 March

2006;

accepted

17

July

006;

final ersion eceived 3

September

006.

Corresponding

ditor: M.

Friedl.

For

reprints

f this nvited

Feature,

ee footnote

,

p.

972.

3

E-mail:

[email protected]

974



June 007 LAND

USE CHANGE

AROUND PROTECTED

AREAS

975

and

intensifying

n the lands

surrounding rotected

areas,

resulting

n

changes

n

ecological

function

nd

biodiversity

ithin

rotected

reas.

Recent satellite-based

hange analyses

are

revealing

that human

populations

and intense

and use have

grown apidly

n recent ecades

around

manyprotected

areas (Hansen et al. 2004). In the tropics, road

construction,

onversion or

agriculture,

nd demand

fornatural esources re

leading

o

clearing

f

primary

forest around reserves

Mustard

et al.

2004)

and

increased

hunting

f native

species

(Escamilla

et al.

2000).

DeFries et al.

(2005)

found

that 66% of 198

reserves

n

the humid

tropics

had

undergone

oss of

forest abitat

n

the

surrounding

ands since

1980,

with

an

average

oss

rate

of 5%

per

decade

within

0 km of

the

boundary.

In other

areas,

increases in

wealth,

technology,

nd

populationdensity

re

leading

o more

rural ettlement

n

previously

ild areas. In the

United

States

ince

1950,

for

xample,

ural esidential

evelop-

mentwas the fastest rowingand use typeand now

covers25% of the ower 48 states

Brown

et al.

2005).

Some

protected

areas are

magnets

for such

rural

development

Chown

et al.

2003).

The counties round

theYellowstoneNational

Park,

for

xample,

re

among

the fastest

rowing

n

the United States

Rasker

and

Hansen

2000).

Even n

ong-established

ocieties uch as

in

China,

agricultural

nd urban and uses continue o

push

nto

unprotected

ildlands round

protected

reas

(Vina

et

al.

2007).

In recent

ecades,

cologists

ave

come

to realize hat

human

impacts

on

surrounding

ands

may

cross the

boundaries

nto

protected

reas

(Buechner

1987,

Das-

mann 1988, Schonewald-Cox1988). The creation of

buffer ones around

protected

reas was recommended

to minimize

egative

oundary

nfluences

Noss

1983).

Accordingly,

UNESCO's

Man and the

Biosphere

(MAB)

program

dvocated

managing

he ands around

protected

reas

along

a

gradient

f

decreasingly

ntense

land use

toward

protected

rea boundaries

UNESCO

1974).

More

recently,

ethodshave been

developed

o

evaluate

the effectiveness f

protected

areas,

with

consideration

f human ctivities

n

surrounding

ands

(Hockings

2000,

TNC

2000,

Ervin

20036).

For some

protected

reas that are not

functioning dequately,

systematic

conservation

planning (Margules

and

Pressey 000) has been used to guide management f

the

regions

round

protected

reas to better chieve

conservation

bjectives

e.g., Pressey

t al.

2003).

Such

efforts

o

mitigate

boundary

influences

n

protected

reas

will be most effectivef based on

scientific

nderstanding

f the

underlying cological

mechanisms.

t can be difficult

o ascertain hemeans

by

which

human

activities outside

of

protected

areas,

sometimes

ens to

hundreds f kilometers

way,

can

impact

ecological

function and

biodiversity

within

protected

reas.

Knowledge

of these

ecological

con-

nections

could

help

to answer several

management-

oriented

uestions.

How

large

s the zone of influence

around

a

protected

rea?

Are all

locations

within

his

zone of

influence

quallyimportant

o

protected

rea

functioning?

hich

ecological processes

or

species

of

organisms

within

protected

areas are

particularly

sensitive

o

surrounding

and

use? Which

and

use

types

and

intensitiesre

most

ikely

o

have

negative

mpacts

within rotectedreas?

Advances

n

spatial

ecology

have

allowed an

increas-

ing

understanding

f the

ecological

mechanisms

on-

necting

protected

reas to

surrounding

ands.

Island

biogeography

heory,

or

xample,

provides

basis for

predicting

xtinction ates

of

species

as a

function

f

habitat

fragmentation

Brooks

et

al.

1999).

This

theory

can be

applied

to address the

effects

f habitat

oss

outside of

protected

reas on

species

richness

within

protected

reas

(DeFries

et al.

2005).

Metapopulation

theoryprovides

a basis

for

determining

hether

subpopulation

f a

species

within

protected

rea is

dependent

upon population

source

areas

located in

surroundingands (Sinclair1998,Hansen and Rotella

2002).

The

purpose

f this

paper

s to

drawfrom

iverse

studies of

spatial

ecology

to derive a

comprehensive

overview f the

ecological

mechanisms

y

which

and

use outside f

protected

reas

may

nfluence

cology

nd

biodiversity

ith

rotected

reas.

This

synthesis

s

meant

to enhance the

theoretical

nderpinning

f

efforts o

assess

the effectivenessf

protected

reas

(e.g.,

Parrish

et al.

2003)

and

systematic

onservation

lanning

cross

regions

ncluding rotected

reas

Margules

nd

Pressey

2000).

Our central hesis s

that

protected

reas are

often

parts

of

larger cosystems

nd that

and use

change

n

theunprotected ortionof theecosystemmayrescale

the

cosystem,

eading

o

changes

n

the

functioning

nd

biodiversity

within

the reserve. We first

present

a

conceptual

model of

protected

reas

embeddedwithin

larger cosystems

hat

ften nclude

urrounding

uman

land use. We

then

explore

the

key

ecological

mecha-

nisms

by

which his and use on

surrounding

ands

may

influence

cological processes

and

biodiversity

ithin

reserves. These mechanisms nvolve

ecosystem

ize,

ecological process

ones,

crucial

habitats,

nd

exposure

to humans.

A

concluding

ection

suggests

how these

ecological

mechanisms

rovide

basis for

ssessing

he

effectivenessf

protected

reas and

systematic

onser-

vationplanning crossprotectedreas and surrounding

lands.

The case studies

n

the

papers

that follow

provide

more detailed

xamples

of

ways

n

which and

use can

influence

rotected

reas. The

managementmplications

of these nteractions re

developed

further

n

DeFries

et al.

(2007).

Protected Areas as Parts of Larger

Ecosystems

Protected

reas sometimes xclude a

portion

of the

area that is

needed to maintain essential

ecological

processes

nd

organisms.

his

was

recognized

y

scien-

tists

tudyingarge

mammalswith

arge

home

ranges

hat



976

INVITED FEATURE

^'^dl^ri

Fig. 1.

Conceptual

model

llustrating

he effects f

land

use

change

on

ecosystem

unction,

a)

Protected reas as

part

of a

larger cosystem

ith

nergy,

materials,

nd/or

rganisms lowinghrough

he

cosystem,

b)

Land use

change

educes ffective

ize

of the

cosystem,

c)

Land use

change

lters

cological

flows,

d)

Land use

change

liminates

nique

habitats

nd

disrupts

ource-

sinkdynamics,e) Edgeeffects rom and use negativelynfluencehepark.

extended utsidenational

parks Wright

nd

Thompson

1935,

Craighead

1979,

Newmark

1985).

More

recently,

ecologists

have

established hat the

spatial

domains of

ecological

processes

such as natural disturbance nd

nutrient

yclingmay

extentoutside

park

boundaries

(Grumbine

1990).

Because

protected

reas were often

designated

based on factors other than

ecological

completeness,

uch as scenicvalue

(Pressey

1994,

Scott

et al.

2001),

they

ometimes o not include the areas

required

o maintain isturbance

egimes,

utrient

lows,

organismmovements,nd populationprocesseswithin

them

Fig.

la).

Following

stablishment,

rotected

reas

may

continue

o function s

parts

of

larger cosystems

because

surrounding

ands remain

undeveloped

and

continue to

provide

functionalhabitats.

f

land use

change

educes abitats

n

the

unprotectedortion

f the

ecosystem,cosystem

unction nd

biodiversity ay

be

degraded

within

he

protected

rea. The modern

oncept

of

ecosystem management grew

from the

goal

of

managing egional

andscapes

o maintain he

cological

integrity

f the

protected

reas that

they

ontain

Agee

and

Johnson

988,

Grumbine

994).

How can the

spatial

dimensions

of the effective

ecosystem ncompassing protected

rea be

quantified?

If the

goal

of the

protected

rea

is to maintain

native

species

and the


hat

they

require,

then he

patial

xtent

f the ffective

cosystem

ncludes

the area that

strongly

nfluences

hese

species

and

processes

Grumbine

1990).

This area can

be

mapped

based on theflows f

materials,

nergy,

nd

organisms.

Watershed oundaries re

often sed to define

he xtent

of

aquatic ecosystems

Pringle

2001).

Watersheds

encompass

he

rea of movement

f

ground

nd

surface

water. Water carriesnutrients uch as

nitrogen

nd

phosphorous,

which are critical

o

plant

and animal

growth.

Water also serves

s a conduit

for the move-

ments f

many quatic

and terrestrial

rganisms.

ence,

strong

nteractions

mong

many components

of an

ecosystem

may

occur

withinwatersheds.

Natural

dis-

turbances uch as wildfire

ove across

andscapes

from

initiation ones to run-out

zones and

differentially

influence

oils,

vegetation,

nd animal

habitatswithin

these ones

Baker

1992).

Ecosystem

oundaries

an be

delineated ased on

homogeneity

f disturbance

egimes

(Pickett

nd

Thompson

1978).

Similarly,many organ-



June 007

LAND

USE

CHANGE AROUND

PROTECTED AREAS

977

Fig. 2.

Depiction

of the GreaterYellowstone

cosystem

s defined ased

on the

biophysical radients,

aturaldisturbance

regimes,

nd

organism

movements

modified

romHansen et

al.

[2002]

and Gude et al.

[2006]).

Shown

are land

allocation,

movement

athways

for two

migratory

pecies,

areas of

high predicted

ird

diversity

biodiversity otspots),

nd

exurban

development

ruralhomes).

The

biodiversity odeling

mask

refers o ocations oo

high

n elevation o

be within

hedomain f

the

bird

diversity redictions.

isms move

predictably

cross the

andscape,

for

xam-

ple,

to

gain

access to seasonal resources.

Ecosystem

boundaries an be defined ased on these

movements r

on the area required o maintain articular opulation

levels f these

rganisms

Newmark1985).

In

practice,defining

he actual boundaries of an

ecosystem

s

subjective.

Although

he

flowsof

water,

nutrients,

isturbance,

nd

organisms

re often nterre-

lated,

their

patial

dimensions re oftennot identical.

Water and nutrients

may

be well

represented

ithin

watershed

boundaries,

but

organismsmay migrate

among

watersheds.

ence,

t is often ifficulto define

a

particular cosystem oundary

hat

s

adequate

for ll

components

f the

ecosystem.

Also,

the

strength

f

interaction must be considered when

defining

an

ecosystem.

Ecological processes

and

organisms

n a

particular

ocation

are often

trongly

inked to

some

places,

weakly

inked o

other

places,

and not inked o

still ther laces.Forexample, limaten theCaribbean

is

heavily

nfluenced

y regional

factors

nd

is

weakly

linked o Saharan

Africavia

input

of wind-borne

oess

(Prospero

nd Lamb

2003).

Thus

ecosystem

oundaries

are

necessarily

bstractions

hat eflect

hehuman

hoice

of the

ecosystem

roperty

f focus

nd the

strength

f

interactions sed

in thedefinition.

In a

growing

numberof

examples,protected-area-

centered

cosystems

ave been defined

see

Meffe t al.

2004).

For

example,

the Greater

Serengeti

cosystem

has been defined

ased on

the

migratory

atterns

f

the

dominant

herbivore,

he wildebeest

Sinclair

1995;

see



978

INVITED FEATURE

EcologicalApplications

Vol.

17,

No. 4

Plate 1

Serengeti

ational

ark,

anzania,

as

designed

o ncludemost f he

migratory

ange

f

he

erengeti

ildebeest

herd.

Many

ther

rotected

reas nclude

nly portion

f rea

required

y

migratory

pecies.

Wildlifen uch

rotected

reas

s

especially

ulnerableo anduse ntensificationn the

urrounding

ands. hoto redit:

. Hansen.

Plate

1).

Although erengeti

National Park is insuffi-

cientto maintain he wildebeest nd other

migratory

mammalswithin

t,

the network f wild and semi-wild

public

and

private

ands across the Greater

Serengeti

Ecosystemmay

be

nearly arge nough

o maintain hese

populations

Packer

et al.

2005;

but see Serneels nd

Lambin2001). The GreaterEvergladesEcosystem as

been defined o

encompass

he massive

ontiguous

rea

of

freshwater

lowly

flowing

seaward

in

southern

Florida.

Everglades

National

Park includes

only

a

portion

f this

arge

watershed

nd

ecologicalprocesses

within

t are

strongly

nfluenced

y

unprotected

ands

higher

n

the watershed

NAS

2003).

The Greater

Yellowstone

cosystem,ncluding

ellowstoneNation-

al

Park,

was defined

argely y gradients

n

topography,

climate,

and

soils,

and the

resulting

movement of

wildfirend

organisms

Keiter

nd

Boyce

1991,

Hansen

et al.

2002,

Noss et al.

2002).

Centered

n the Yellow-

stone Plateau

and

surrounding

mountains,

natural

disturbance egimes nd organismsmove across the

elevational

gradient

from

valley

bottoms to

high

mountains

in

response

to climate and

vegetation

productivityFig.

2).

In these

ases,

knowledge

f the

spatial

domainof

strong cological

nteractionsetween

protected

reas and the

surrounding

reas

has allowed

for

pecification

f the

larger

ffective

cosystem.

he

term

greater cosystem

s often sed to describe hese

protected-area-centeredcosystems

Keiter

and

Boyce

1991).

Recognizing

hat

protected

reas are often

parts

of

larger cosystems

elps

to

clarify

he ffects f and use.

Agriculture,

ettlement,

nd otherhuman and uses

in

the

unprotected art

of

the

ecosystemmay

alter

the

flows of

energy,

materials,

nd

organisms

cross

the

ecosystem

n

ways

that

change

ecological

functioning

within the reserve

Fig.

lb-e).

If,

for

example,

the

portion

f the

cosystem

herewildfire

ends o

gnite

s

converted

griculture,

ire

may

less

frequently

pread

into theprotectedrea and altervegetationuccession.

Similarly,

f land use decreases

the area

of suitable

habitats or wildlife

opulation

elow

ome

threshold,

the

population

size

may

fall

to the

point

where

extinction

s

likely.

Moreover,

and use

near a

protected

area

may

introduce

novel disturbances

o

which the

ecosystem

s not

adequately dapted.

Human

hunting

and intense utdoor

recreationre

examples.

Giventhat

human and use is

rapidly

xpanding

nd

intensifying

n

the

unprotected

arts

of

many protected

rea

ecosys-

tems,

t is critical hatwe better nderstand

ts

effects.

Knowledge

of

the mechanisms

onnecting

and use

to

protected

areas

can

provide

an

objective

basis for

defininghespatialdomainof the effectivecosystem

encompassing

protected

reas

and for

managing

the

unprotected

ands to

maintain

cological

function

nd

biodiversity

ithin

rotected

reas.

Mechanisms

inkingLand Use

to Protected

Areas

Advances n

ecological

heory

ave

allowed

ncreased

understanding

f

how the

spatial

patterning

cross

landscapes

and

regions

influences

ocal

ecosystems

(Turner

t al.

2001).

Island

biogeography,

pecies-area

relationships,metapopulation

dynamics,

disturbance

ecology,

nd

landscape

ecology

have

increasingly

een

applied

to

questions

f conservation

iology,

ncluding



June

007

LAND USE CHANGE AROUND PROTECTED

AREAS

979

Table 1 General

mechanisms

y

which

and use

surrounding rotected

reas alters

cological

processes

within

eserves.

General

mechanism

and

type

Description

Examples

Change

neffectiveize of

reserve

Minimum

ynamic

rea

Temporal

tability

f serai

stages

s a function f the

Hurricanes

n

Puerto

Rico

area of the reserve elative o the ize of natural (Shugart1984).

disturbance.

Species-area

ffect

As wild habitats

n

surrounding

ands

are

destroyed,

he

Fragmented

orestsn

Kenya

functionalize of the reserve s decreased nd risk f

(Brooks

et al.

1999);

harvest

extinction

n the reserve s increased. of

primary

orest utside

Calakmul

Biosphere

Reserve,

Mexico

Vester

t al.

2007).

Trophic

tructure

Characteristic

patial

scales of

organisms

iffer ith Loss of

predators

n Barro

trophic

evel uch that

organisms

n

higher

evels re Colorado Island

Terborgh

lost

as

ecosystems

hrink. et al.

2001).

Change

n

ecological

flows

into and out of reserve

Initiation nd

run-out

Key ecologicalprocesses

move across

andscapes.

Fire

n

Yellowstone

National

zones

Initiation nd run-out ones for

disturbance

may

ie

Park

Hansen

and Rotella

outside

reserves.

2001).

Location

n

watershed

r

Land use

in

upper

watersheds r airsheds

may

alterflows Rainfall nMonte Verde loud

airshed intoreservesower nthe watershed r airshed. forestLawtonet al. 2001).

Loss of

crucial

habitat

outside

f reserve

Seasonal

and

migration

Lands outside f

reserves

may

contain

unique

habitats

Large

mammals

n

the Greater

habitats

that re

required y organisms

within eserves.

Serengeti

Serneels

nd

Organisms

equire

orridors

o

disperse mong

Lambin

2001);

antelope

n

reserves r to

migrate

rom eserves o

ephemeral

GreaterYellowstone

Berger

habitats.

2004).

Population

ource-sink

habitats

Increased

xposure

o

Unique

habitats

utside f reserves

re

population

ource Birds round Yellowstone

humans

t

park

edge

areas

required

o

maintain ink

populations

n National Park

Hansen

and

reserves.

Rotella

2002).

Hunting/poaching;

Negative

human

nfluences rom he reserve

eriphery

Eurasian

badgers

n

Donana

exotics/disease

extend ome

distance

nto

protected

reas.

Park

Revilla

et al.

2001);

spread

of disease from

etsto lions n

Serengeti

ational

Park

Packer

et al.

1999).

the

design

of nature

eserves

Pressey

t al.

1993,

Noss

and

Cooperrider

994,

Prendergast

t al.

1999).

These

bodies

of

theory

an

also be

applied

to

understanding

how

changes

in the

unprotected

parts

of

greater

ecosystems

may

influence

rotected

reas.

Here we

synthesize

cross

these bodies

of

theory

o

develop

a

simple

conceptual

framework

or

understanding

ow

changes

urroundingrotected

reas alter

he

cological

processes

within

hem.

According

o our

framework,

four

general

mechanismsinkhuman land uses with

ecological

function

ithin

rotected

reas.

These

mech-

anisms

nvolve

ffective

ize of

the

ecosystem,

lows

f

ecological

process

ones,

crucial

habitats,

nd

exposure

to humans

t

reserve

dges

Table

1).

We

will

describe,

for ach

of

mechanisms,

hevarious

forms

n which

hey

may

be

expressed,

he

conceptual

basis,

and

illustrative

examples.

Effective

ize

of ecosystem

We refer

o effective

ize of

the

ecosystem

s the

area

that

ncludes

cological processes

nd

organisms

integral

o the

protected

rea. This often s correlated

with he rea

of wild and semi-wild abitats

within

nd

surrounding

the

protected

area.

By reducing

this

effective

ize,

land use can

negatively

nfluence oth


and

communitydiversity

nd

structure

Fig.

lb).

Minimum

ynamic

rea.

Island

biogeography

heory

suggests

hatthe number f

species

n a nature eserve

results from

the balance between colonization

and

extinction.f protected reas are increasinglysolated

from xternal

olonization

ources,

extinction ill hen

become the dominant

population

process affecting

equilibrium

n reserves

nd

species

numbers

will

decline

to a

new

level,

according

to Pickett nd

Thompson

(1978:28).

Hence it

s

critical

o maintain ecolonization

sourceswithin

rotected

reas. Natural

disturbances a

key

force

n

driving atch

dynamics

within nd

among

protected

reas

and resources vailable

to

organisms.

Landslides, floods,

wildfires,

nd hurricanes nitiate

succession nd maintain

esources

or

pecies

ssociated

with ach serai

stage.

Bormann

nd Likens

1979)

used



980

INVITED FEATURE


Vol.

17,

No.

4

the term

shifting teady-state quilibrium

o define

landscapes

where isturbance

as

adequate

to maintain

each serai

stage

n

relatively

onstant

proportion

ver

time. The location of disturbance hifts cross the

landscape

over

time,

but the

representation

f

early-

and late-seral

patches

across

the

landscape

remains

within steadystate. Such landscapes can support

relatively igh

numbers f

organisms

ecause

recoloni-

zation sources re

continuously

maintained or

pecies

requiring

ither

arly-

r late-seral onditions.

Pickett nd

Thompson

1978:27)

defined minimum

dynamic

area as the smallest area with a natural

disturbance

egime,

which

maintains

nternal ecoloni-

zation

sources,

and hence minimizes xtinction.

n

other

words,

minimum

ynamic

rea is the mallest rea

withinwhich he

natural isturbance

egime

maintains

shiftingteady-statequilibrium.

As land use

change

reducesthe

effectiveize of

the

ecosystem

ontaining protected

rea,

the

ecosystem

s

increasinglyikely o fall below the minimum ynamic

area

(Baker 1989).

n this

ase,

the

protected

rea itself

will be too

small to maintain

dynamic teady-state

equilibrium

nderthe nfluence f

naturaldisturbance.

At this

point,

ecolonization ourcesfor

pecies

re lost

and

extinctionates

will

rise.How

big

does

an

ecosystem

need be to maintain

dynamic teady-statequilibrium?

Shugart

(1984)

and Baker

(1992)

suggested

that a

landscape

needs to be at

least 50 times

arger

han the

area of the

largest

disturbance to maintain this

equilibrium.

We

are not aware of

any

studies that

have docu-

mented

change

n

effective

cosystem

ize

resulting

n

loss ofminimumynamicrea and loss ofspecieswithin

a

protected

rea. Baker

1989)

found hat he

Boundary

Waters Canoe Area in

northernMinnesota was not

large enough

to maintain a fire-induced

teady-state

equilibrium.

Wimberly

t al.

(2000)

found that the

minimum

ynamic

rea induced

by

fire

n

the

Oregon

Coast

Range

in

pre-European

settlement imes was

larger

than the

old-growth

orest eservesmaintained

today. McCarthy

nd

Lindenmayer

1999, 2000)

used

spatially

xplicit opulation iability

models o estimate

the

minimumize of

protected

reas needed o maintain

viable

populations

of forest

marsupials

n Australia

undervariousdisturbance

egimes.

Species-area effects. A well-known enetof island

biogography heory

s that the number f

species

that

are found n an oceanic

sland or

in a

habitat

ragment

is a function f

its area.

A

large

body

of

empirical

evidence

ndicates hat the number f

species

termed

species

richness),

,

increaseswith rea

A,

according

o

the

quation

S

=

cAz where and

z

are constants

e.g.,

Rosenzweig

995).

Hence,

pecies

ichnessncreaseswith

island or

habitat

rea,

at a

decelerating

ate for

arger

areas.

The

primary xplanation

s that

given pecies

s

less

ikely

o

go

extinct

f

the area of suitablehabitat s

large

enough

to

provide

the resources

o

allow

for a

population

ize

larger

han a minimum iable

popula-

tion below

which risk of

extinctions

elevated

Pimm

et al.

1988).

The

species-area

elationship

as been used

to

predict

the

consequences

of

reducing

the size of

a habitat

through

onversion o

intensiveand

uses

for

review,

see Cowlishaw

1999).

A

contraction

n habitat

from ts

original rea to its new area is predicted o lead to a

decline from

he

original

number f

species

to a new

total based

on the size of the

fragment.

his

number s

expected

to be further

educed

through

ime as

the

effects f

isolation ead to

local extinctions

ithin he

fragment,

ue

to small

population

izes.

n

a test

f this

approach,

Brooks

et al.

(1999) surveyed

irds

n

upland

forest

fragments

n

Kenya. They compared

current

species

richness

orforest irdswith hat

from he

time

prior

o habitat

fragmentation,sing

museum

records.

They

found hat ach of

the fivehabitat

fragments

ad

undergone

extinctions f

forest birds and

that the

number f extinctions as

close to that

predicted,

ased

on thechange n area.

Following

habitat

ragmentation,

herelaxation

o the

new reduced

species

richness

may

take decades

to

centuries r more

Burkey

1995,

Brooks

et al.

1999).

The term extinction ebt s used

to denote henumber

of

species

that are

expected

o become extinct

s the

communitydjusts

o a

new,

maller,

rea of habitat.

n

the New World

tropics,

deforestation s

sufficiently

recent

that few extinctions

have

yet

occurred.

In

confirmation f

the

species-area approach,

however,

Brooks

and Balmford

1996)

and

Brooks et

al.

(1997)

found that the

predicted

number of

extinctions

or

Atlantic orests f South

America nd insular outheast

Asia closelymatched he numbers f speciescurrently

listed s threatened

ith xtinction.

n

tropical

orests f

Africa,

Cowlishaw

1999)

predicted

hatcurrent

efor-

estationwill

eventually

esult

n

the xtinction

f >30%

of

the forest

rimate

auna

n each of several ountries.

The

implication

f the

species-area

relationship

or

protected

reas is that the

number of

species

in a

protected

rea

will

decline as

the effectiveize of the

reserve s reduced

through

estruction

f the

unpro-

tectedhabitats

urrounding

he reserve.

his

point

was

illustrated

y

Pimm nd Raven

2000). They

focused n

biodiversity

otspots

around the world

identified

y

Myers

t al.

(2000).

These

hotspots

ave

already

uffered

disproportionateoss

of

primary egetation,meaning

thatthe

many pecies hey

ontain re

under

particular

threat f extinction.

sing

the

pecies-area

elationship,

Pimm and Raven

(2000) predicted

hat

more

species

would be lost

f

onlyhotspots

ow

n a

protected

tatus

were aved than

f

all

hotspot

habitats

both

nside nd

outside

protected

reas)

were aved.

The

species-area pproach

was

applied

to three f the

case

study

ocations

reviewed

n

this

nvited Feature:

Maasai East

Africa,

Southern

Yucatan,

and

Greater

Yellowstone

H.

L.

Rustigian

t

al.,

unpublished

anu-

script).

Habitats

hathad been

deforested

nd converted

to

agriculture,

ettlements,

r rural

dispersed

homes



June

007 LAND

USE CHANGE

AROUND

PROTECTED

AREAS

981

Table 2. Loss of habitat

ince

pre-European

ettlementimes

nd

predicted

xtinctionsf birds

nd

mammals

under

urrent

remaining

abitat nd habitat

emaining

f

all

unprotected

ands are

converted o

human and

uses.

No.

(and %)

species

predicted

xtinct

Total

Original

Original

Underfull

Location

area

(km2)

habitat ntact

richness

Current

conversion

Maasailand,East Africa 193405 55% 756 107(14.2%) 262(34.7%)

Greater

Yellowstone,

USA

95

363

89%

284

14(4.9%)

26(9.2%)

Mayan

Forest,

outhern ucatan Peninsula

120 109

70%

315

26

(8.3%)

75

(23.8%)

Notes:

Data are fromH. L.

Rustigian

t

al.

{unpublished

anuscript).

ichness s the

number f

species.

since

pre-European

ettlementimeswere

onsidered ot

suitablefornativebird and mammal

pecies.

Based on

loss of habitats rom

re-European

ettlement

imes,

he

ecosystem

roundeach

park, hey redicted

loss of

5-

14%

of

species among

the sites

(Table

2).

If

all

unprotected

abitats were converted o human land

uses,

9-35%

of

species

were

predicted

o be lost.

A

limitation f the

species-area pproach

to estimat-

ing fragmentationffectss thatspeciesdiffern their

tolerance

o the

type

nd

intensity

f human

and

use.

Many

native

species

find suitable habitat in human-

altered

andscapes

and some of these

species

become

more abundant

under certain and uses

(McKinney

2002).

The

approach

willbe most ffectivef t s

applied

to

species

hat

re unable to tolerate hehuman-induced

changes

to the

unprotected

ortion

of the

ecosystem.

The

approach

will lso be more ccurate

f

he

uality

f

losthabitats

s considered. ester t al.

(2007)

found hat

tall

primary

forest

has been

disproportionately

e-

stroyed

n the southern ucatan

region.

everal

pecies

of

trees nd

butterfliesre

uniquely

ssociated

with his

foresttype and may have been disproportionately

affected

y

ts

oss in area.

Trophic

tructure.

A

third

onsequence

of

reducing

the effective

ize of nature reserves s an altered

representation

f

organisms

t various

levels of the

foodchain.

Perhaps

the most

typical

ase is the oss of

high-levelpredators

and the release of meso-level

predators

r herbivores. ome

range

size and

density

are

associated

with level

in

the food chain.

Top

predators

tend to have

relatively arge

home

range

requirements

nd low densities.

onsequently, hey

re

particularly

ensitive o extinction s effective eserve

size

decreases

Schonewald-Cox

1988,

Woodroffe nd

Ginsberg 998).

Perhaps

the

most direct

evidence that

trophic

structure

n

protected

reas varies

with

ffective

ize of

ecosystem

omes from orrelational

tudies of

species

extinction

n

protected

reas of

differing

ize. Rivard

etal.

(2000)

found

hat xtinction

atesof mammals

n

Canadian

national

arks

were ssociatedboth

with

ark

area and

with he

extent f intense anduse outsideof

parks.

Moreover,

they

found

that

species

with

large

home

ranges,

ypical

f

species

t

higher rophic

evels,

were more

likely

to suffer xtinction.Evidence for

similar

patterns

n

aquatic systems

omes

from Post

et al.

(2000),

who

studied

rophic

tructure

n 25 north-

temperate

akes.

They

found that

the

length

f

food

chains was

positively

elated o

ecosystem

ize

(size

of

the

lake).

Specifically,

igher

rophic

evels

were more

commonly

ound n

larger

akes.

In

some

systems,

hese

top predators

nfluence he

abundance of

organisms

ower

n

the food

chain with

cascading

ffects

hroughout

he

ecosystem.

ence,

oss

of

the

top predators

may

allow

meso

predators

or

herbivores o become ncreasinglybundant.Thiseffect

on

trophic

tructure as

documentedn

a

study

f sland

habitat

fragments

reated

by

a water

impoundment

project

n

Venezuela

Terborgh

t al.

2001).

Vertebrate

predators

went

extincton small-

and

medium-sized

islandsbut remained n

larger

slands.

Densities

f seed

predators

nd herbivores

ere

10-100 times

higher

han

thoseon the

mainland,

robably

ue to the

release rom

predation.

These

changes

cascaded

through

he food

chain,

resulting

n

severe

reductions

n

densitiesof

canopy

tree

eedlings

nd

saplings.

Trophic

cascades

associated with

top predators

re

also

suggested

n

the

GreaterYellowstone

Ecosystem.

The recent eintroductionfthewolfCanus upis), top

predator

hathad

been extinct or60

years,

ppears

to

be

expanding

the

scavenger

community,

reducing

mesocarnivoresnd

ungulate opulation

izes,

releasing

the

riparian lant community

hatwas

overbrowsed

y

ungulates,

and

allowing

for

expansion

of

riparian-

dependent

ird ommunities

Ripple

and Beschta

004).

Thus,

reduction n

the effectiveize of a

protected

area is

predicted

o result n losses in

species

due

to

change

n

landscapedynamics,

pecies-area

ffects,

nd

loss of

top

carnivores. s the

unprotected

ands

around

nature reserves re

increasingly

onverted o

intense

human and

uses,

ffective

cosystem

ize s reduced.

or

relativelyewprotectedreas do we know theeffective

size of the

cosystem

nd the

rateof oss to human

and

use.

Rustigan

t al.

{unpublished

ata)

found

that,

for

three ase

study andscapes,

he oss rate ince

presettle-

ment

times was substantial:

11%, 30%,

and 45% for

Greater

Yellowstone,

Mayan

Forest,

and Greater

Serengeti cosystems

have

been converted o

intense

humanuses.

Ecologicalprocess

ones

Just s

organisms

move cross

andscapes

t character-

istic

cales,

cologicalprocesses

esult n

flows f

energy

and materials

long predictable athways.

hese flows



982

INVITED FEATURE


Vol.

17,

No.

4

may

be

important

o

ecological

functionn

influencing

local

ecological rocesses

uch

s

primary roductivity

r

habitat

uitability.

o the xtent hat and use conversion

and intensificationlters

ecological

flows across the

landscape,

t

may

mpact

he

ecologicalfunctioning

nd

biodiversity

ithin

rotected

reas

Fig.

lc).

Disturbance nitiation nd run-out ones. Disturb-

ances tend o be initiated

n

particular

andscape ettings

and move to other ocations

n

the

landscape.

Inter-

actions between the location

where disturbance

gets

started

initiation ones)

and locations wheredisturb-

ances

move

to

run-out ones)

nfluencehenature f the

disturbance

egime

n an area

(Baker

1992).

In south-

western

Montana,

for

example, ightning

trikes ccur

across

the

andscape,

but more

frequentlygnite

ires n

dry

valley-bottom rasslands

than in moist conifer

forests

n

the

uplands

Arno

and Gruell

1983).

These

fires

hen

preadupslope

o the onifer orests.

hus,

the

juxtaposing

of

grasslands

nd conifer orests

trongly

influences he regionalfireregime.Local disturbance

regimes

an best be maintained n

protected

reas that

include the

disturbance

nitiation ones within their

boundaries

Baker

1992).

In the case of the Montana

example,

a

protected

rea

placed only

in the

upland

conifers

may

uffer oreor less

frequent

ire,

epending

on

the

management

f the

valley-bottom rasslands

outsideof the

protected

rea.

It is also

important

o include disturbance un-out

zones within

heboundaries f

protected

reas.

Run-out

zones

may

ontain

nique

biotic onditions

nd habitat

patterns mportant

o


nd

organ-

isms.

For

example,

flood

severity

ften ncreases

rom

headwaters o large floodplains. he largescoursand

bare

gravel

ars and themosaic of serai

tages

hatform

on

floodplains upporthigh

evels

of

biodiversity

Saab

1999).

A

protected

rea that does not contain this

disturbance un-out one

will

not include hese

unique

riparian

egetation

ommunities.

n

protected

reas that

omit either the initiationor run-out

zones,

human

manipulation

f

disturbance

may

be

required

o main-

tain

landscape patterns

nd

organisms

Baker

1992,

Arcese nd

Sinclair

1997).

Location in watershed r airshed. Protected reas

may

be

heavily

nfluenced

y

hydrologic

lows

and

weather

ystems.

or

example, rotected

reas

through-

out the world are threatenedycumulative lterations

in

hydrologic onnectivity

ithin he

larger

andscape

(Pringle

2001).

Humans are

altering

hydrologic

lows

directly y

dams,

water

diversions,

roundwater

xtrac-

tion,

and

irrigation,

nd

indirectly y altering

and

cover,

which

may change

ratesof

transpiration,

unoff,

and soil

storage.

These flows f water

ransport nergy,

nutrients,ediments,

nd

organisms.

The location of a

given protected

rea

within

a

watershed,

elative o

regional quifers

nd wind and

precipitationatterns,

an

play

a

key

role

n ts

response

to

human disturbance ransmitted

hrough

he

hydro-

logic cycle.

Protected

reas located

n

middle

nd lower

watersheds ften

experience

ltered

flow

regimes

nd

inputs

f exotic

rganisms

nd

pollution

rom

pstream.

The Colorado

River

withinGrand

Canyon

National

Park,

for

example,

has

undergone

dramatic

trans-

formation ue

to dams

and intense

and use

in the

headwaters

f thewatershed

Cohn

2001).

Alteredwater

temperaturesnd loss of flooddeposition f sediments

have

changed

habitats,

eading

to a substantial

educ-

tion of nativefishes.

Waterborne

eeds of

exotic

plants

such s saltcedar

Tamarix

pp.)

have ed to

entirely

ew

riparian

ommunities

nd loss

of native

iparian

pecies

such as the

Willow

Flycatcher

Empidonax

raillii).

Protected

reas

in

upper

watersheds,

n

contrast,

re

vulnerable o

land use lower

n the watershed.

hese

human ctivities

may provide

vectors

or xotic

pecies

and disease to

penetrate

he

upper

watershed.

heymay

also result

n

genetic

solationof

populations

n head-

waters.

n the northern

ocky

Mountains,USA,

native

west-slope

utthroat

rout

Oncorhynchus

larki

ewisi)

historicallyccupiedentirewatersheds.ubpopulations

in headwaters

may

have been

dependent

on source

populations

n lowlands.

Mainstream

opulations

were

forced

o

extinction

y

the

ntroductionf exotic

trout

species

nd

possibly

y

habitat

hanges

ssociated

with

irrigation

nd

other ntense and

uses.

Consequently,

populations

urviving

n

headwater

treams

n

national

parks

are

subject

to a

high probability

f

extinction,

probably

because

they

no

longer

receive

mmigrants

from ource

populations

n lowland streams

Shepard

et

al.

1997).

This discussion

of watershed

ffects lso

applies

to

airsheds.

Change

n

regional

anduse

may

alter limate

and nutrient epositionwithin downwindprotected

areas considerable

distances

away

(Lawton

et

al.

2001).

Tropical

montane cloud

forests

n

Central

America

depend upon prolonged

mmersion

n clouds.

Clearing

f forests

n Costa

Rica's Caribbean

owlands

appears

to have reduced

cloud cover

and increased

cloud

height

n

cloud

forests,

uch

as

in Monte

Verde

National

Park,

ltering cosystem

unctionnd

possibly

contributing

o

thedecline f 20-50

species

f

frogs

nd

toads

in Monte Verde

National Park

Nair

et al.

2003).

Such

changes

n

ecosystem rocesses

are

especially

difficult or

managers

of

protected

reas

to

perceive

because

they may

resultfrom

changes

in the

air or

watershed t longdistances rom heprotectedrea.

Crucial

habitats

Protected

reas

may

not

contain

the full suite

of

habitats

equired

y organisms

o meet

heir nnual

ife

history requirements.

easonally important

habitats

may

ie outside

heboundaries

f

protected

reas.

Land

use in the

unprotected

ortion

f

ecosystems

may

alter

or

destroy

hese easonal

habitats,

s

well as

movement

corridors

onnecting

hese habitats

to

protected

reas

(Fig.

Id).

This situation s common

because

of the nonrandom

location of

protected

areas

relative to

biophysical



June 007

LAND USE CHANGE

AROUND PROTECTED

AREAS

983

conditions

and habitats. Protected areas

are often

located

in

relatively

harsh

biophysical

settings

nd

represent

he colder or

hotter, rier,

more

topograph-

icallycomplex,

nd/or

ess

productive ortions

f the

broader

cosystems

n

which

hey

ie

Scott

et al.

2001).

Intense uman and

use,

n

contrast,

s often

entered n

more equitable and productive landscape settings

(Seabloom

et al.

2002,

Huston

2005).

Consequently,

the

unprotected ortions

of

ecosystems

ftencontain

habitats crucial for

organisms

hat reside within

he

protected

reas for

ortions

f the

year.

ntenseanduse

may

be

disproportionately

enteredon these

unpro-

tected rucialhabitats

Hansen

and Rotella

2002).

Seasonal

and

migration

habitats. Animals

often

move across the

andscape

easonally

o obtain

required

resources. or

protected

reas at

higher

levations,

ey

winter

ranges

are often outside the boundaries of

protected

reas.

Similarly,

rotected

reas

in

more arid

regions

ften o not containwet-season abitats.Land

usemayalter heseunprotectedeasonal habitats r the

movement

athways

etween easonal habitats.

Populations

of several

arge-mammal pecies

within

the Maasai Mara

Reserve nd AmboseliNational Park

in

Kenya

have declined

n

abundance

during

he

past

30

years

n the face of

rapid

ntensificationf land use

in

the

urroundingegions

A.

J.Hansen et

al.,

unpublished

report).

f

the 15

species

nalyzed,

ight

pecies

n

the

Massai Mara Reserve

and one

species

in

Amboseli

declined

ignificantlyuring

his ime.The declineswere

severe or

everal

pecies

nd

ranged

rom .7% to 2.5%

of the

population

size

per

annum. These

population

changes

were

statistically

ssociated with human use

factors in the wet-season habitats outside of the

protected

rea

boundaries.

n

tropical

orests f

Borneo,

Indonesia,

long-distance

migrations

f bearded

pigs

have been

disrupted

y

the

ogging

f the

dipterocarp

trees

whose fruits re

prime

food sources for the

pigs

(Curran

t al.

1999).

Berger

2004)

documented hat he

225-km

movement orridor

for

pronghorn

ntelope

(Antilocapra

mericana)

betweensummer nd winter

range

n GreaterYellowstone

passes

through

1 km

wide

bottleneck

hat s now threatened

ithnatural

gas

development.

Population

ource-sink abitats.

The crucialhabitats

outside of

protected

reas

may

be

especially

rich

in

resources nd may act as population source areas.

These habitats

may

allow

subpopulations

o

produce

surplus offspring

hat

disperse

to less

productive

habitats

n

protected

reas and allow

persistence

f the

subpopulations

n

the

reserves. or

example,

Hansen

and Rotella

(2002)

found

that bird

populations

n the

Greater Yellowstone

Ecosystem

were concentrated

n

small

hotspots

n

productive,

owland

settings

utside

protected

areas.

These source habitats

have been

disproportionately

sed for

griculture

nd rural

home

sites

(Gude

et al.

2007).

This

intense

and use has

converted these low-elevation source areas

for the

populations

to sink areas and

reduced the

viability

of

subpopulations

n

the more

marginal

habitats n

protected

reas.

Similarly,

Arcese and

Sinclair

1997)

suggested

hatmostof

Serengeti

ational

Park s a

sink

for he ion

population

nd that

he

pecies

s

maintained

there

because of

connectivity

ith

the

Ngorongoro

Conservation

Area,

which s a

population

ource area

for ion.

The

migratory

ovements f

many

organisms

cross

greaterecosystems

re

often

quite

obvious

to

park

managers

and local

people.

Hence,

the

problem

of

unprotected

easonal habitats

has

received

onsiderable

attention n

many

regions.

Designation

nd

protection

of

migration

orridors s

increasingly

idely

used to

minimize r

mitigate

onflicts

etween

human and use

and

migrating

ildlife

e.g.,

Miller t al.

2001).

Proximity

o

humans

Human

presence

n the

periphery

f

protected

reas

may

cause

changes

n

ecosystem

rocesses

nd

biodiver-

sity hat xtend arying istancesnto heprotectedrea

(Fig.

le).

Some of these

edge

effects

esult

n

habitat

change.

For

example,

learing

f forests

o the

edge

of

the

protected

area

boundary may

lead to

elevated

disturbance ates and

high

levels of

forest

mortality

within he forest

eserve

Laurance

et

al.

2000).

Other

types

of

edge

effects o not

cause

visible

changes

in

habitat,

but

have

strong

influences n

organisms

n

protected

reas

nonetheless.

unting

nd

poaching

often xtend

the

footprint

f human settle-

ments nto

adjacent

protected

reas

(Escamilla

et al.

2000,

Revilla et al.

2001).

For

example,

n the western

border of

Serengeti

National

Park,

poaching

was

estimated oextend pto 25km nto hepark Campbell

and Hofer

1995).

Exotic

organisms

nd disease

lso

may

spread

fromborder ommunitiesnto

protected

reas.

Bison

{Bison bison)

in

Yellowstone

National Park

contracted

rucellosis

while

ommingling

ith

ivestock

in

winter

range

outside of the

park

(Yellowstone

National

Park

1997).

This has led to a

substantial

management hallenge

now thatthe disease has been

largely

radicated

from

ivestockherds

n

the

United

States.

Similarly,

ions in

Serengeti

National

Park

underwent ramatic

opulation

eclines rom

he anine

distemper

hat

they

contracted

from

domestic

dogs

living

outside the

park

(Packer

et al.

1999).

Human

recreation is sometimeselevated near borders of

protected

areas and

may displace

wildlife

Hansen

et al.

2005).

Many

of these

edge

effects re

proportional

o the

density

f

the

adjacent

human

population

Woodroffe

and

Ginsberg

998,

Brashares

t

al.

2001).

Hence,

these

effects

may

be increased under human

population

growth

round

protected

reas.

Implications or

Conservation and

Management

This frameworkf

mechanisms

inking

and use with

protected

areas can

enhance the

various

ongoing

conservation nd

management pproaches.

The frame-



984 INVITED FEATURE

Ecological

Applications

Vol.

17,

No. 4

Table 3.

Criteria or

managing egional

andscapes

to reduce the

mpacts

of land use

change

outside

of

protected

reas

on

ecologicalprocesses

nd

biodiversity

ithin eserves.

Mechanism

Type

Design

criteria

Change

n effective

ize of reserve

species-area

ffect;

minimum

ynamic

rea;

trophic

maximize

rea of functional

structure

habitats

Changes

n

ecological

flows nto disturbancenitiation nd run-out

ones;

placement

identify

nd maintain

cological

and out of reserve in watershed r airshed process ones

Loss of crucialhabitat utsideof

ephemeral

abitats;

ispersal

r

migration

abitats;

maintain

eymigration

nd

reserve

population

ource inkhabitats

sourcehabitats

Increased

xposure

o human

poaching;displacement;

xotics/disease manage

human

proximity

nd

activity

t

reserve

dge

edge

effects

work

provides

conceptual

basis for:

1)

mapping

he

boundariesof the effective

cosystem

ncompassing

protected

rea;

(2)

monitoring

nd

assessingmanage-

ment

ffectiveness;

3)

systematic

onservation

lanning

and

management

f the effective

cosystem;

nd

(4)

assessingvulnerability

f

protected

reas to land use

change.We willdiscuss ach of these pplications.

Ecosystem

oundaries

Managers

need to

be able to

quantify

he boundaries

of the ffective

cosystem

ncompassing protected

rea

in

order to know where

to monitor and use

change,

assess

management

ffectiveness,

nd

implement egion-

al

conservation

trategies

o maintain

he

protected

rea.

The

ecosystem

oundaries hould nclude he reas that

are

strongly

connected to the

protected

area in

ecological

processes

or

organism

movements and

population

processes.

The mechanisms

framework

provides

a

conceptual

basis for

mapping

these con-

nections nd identifyingheboundaries f the effective

ecosystem.

he area of

seminatural abitats ontribut-

ing

to effective

cosystem

ize can be

mapped using

remote-sensing

echniques

Rogan

and Chen

2004).

The

spatial

and

temporaldynamics

f natural disturbance

regimes

can be

mapped

from historic records or

projected

with

omputer

imulationmodels

e.g.,

Baker

1989).

Movements of

organisms

can be

quantified

through

se of

telemetry

nd othermethods

e.g., Berger

2004).

Mapping

of

population processes

and

source-

sink

dynamics

equires

othfield tudies nd

simulation

modeling e.g.,

Hansen and Rotella

2002). Quantifica-

tion

of human

edge

effects an be

done

with human

surveys nd other ssessmentmethods Campbelland

Hoffer

995).

n

addition o these

uantitative

methods,

expert opinion

often will be

needed to delineate

ecologicallymeaningful

oundaries.

Management

ffectiveness

nd

monitoring

After a

period

of focus on

the creation of new

protected

reas,

many

managers

nd

conservationists

are

attending

o the assessment f how

well

existing

protected

reas are

working.

or

example,

the World

Commission

n Protected

reas

WCPA)

has

developed

a

six-step rocess

for

assessingmanagement

ffective-

ness.

The

process

begins

with

stablishing

he context

of

existing

values

and

threats,

progresses

through

planning

nd allocation of

resources

inputs),

nd,

as

a

result of

management

actions

(process),

eventually

produces goods

and services

outputs)

that result

n

impacts

or outcomes

Hockings

2003:826).

This

ap-

proach may

be based on results

f

questionnaires

f

managersnd other takeholders r on quantitativeata

from

ecological

measurement.

Our

frameworkof

mechanisms

rovides

a

conceptual

basis for

portions

of the

process.

Context an be

evaluated

by

assessing

threats rom

and use relative o the

places

and

processes

identified

n

our

ecological

mechanisms.

Planning

nd

management process

an

be aimed at

maintaining

he

connections nd functions

dentified

y

our mechanisms.

The

U.S.

National Park Service

and the Canadian

Park Service

Parks

Canada

Agency

2005)

have

each

established

nventory

nd

monitoring

I&M)

programs

aimed at

assessing

he

condition f

parks

nd determin-

ing management

ffectiveness.

ithin he U.S.

National

Park Service &M Program,ur mechanismsramework

has been used to

guide

selection

of

monitoring

indicators,

elimitate he

effective

cosystem,

nd

guide

analysis

of trends

n threats nd

ecological

response

(D.

A.

Jones t

al.,

unpublished

anuscript).

Regionalmanagement

It

is

apparent

hat

manyprotected

reas

may

become

degradedby

and use and other actors

ccurring

n the

unprotected

arts

of the

surrounding

cosystem.

hus,

maintaining

rotected

reas often

will

require

ome evel

of conservation-oriented

anagement

n the

unpro-

tected

portion

f

the

ecosystem. Systematic

onserva-

tionplanning Margulesand Pressey 000) provides

coordinated

pproach

for ssessment

nd

management

across

regional andscapes.

Our

mechanisms ramework

provides design

criteria

for

regional

management

(Table

3).

Knowledge

of land use

patterns,

he

spatial

dynamics

of these

ecological

mechanisms,

nd the

responses

f

ecological processesprovide

context o

identify

laces

n

the

unprotected arts

f the

cosystem

that re most ritical or

maintainingcological

function

within

protected

reas.

Management

hould focus

on

maintaining

ffective

cosystem

ize,

ecological process

zones,

rucialhabitats or

rganisms,

nd on

minimizing

negative

uman

dge

effects.

oupled

withunderstand-



June 007 LAND USE CHANGE

AROUND PROTECTED AREAS

985

Table 4.

Varying

ffects f differentand

use

types

n

ecological

mechanisms

ltering

eserves.

Effective

Ecological

process

Crucial

Edge

Type

of and use

change

reserve ize

zones/flows

habitats

effects

Resource xtraction

Logging

x

xx

Mining x x

Poaching

x

Food

production

Subsistence

arming

x

Small-scale

arming

x

x

x

x

Large-scale

ommercial

arming

x

x

x

x

Recreation

Tourism

x

Infrastructure

Roads/other

ransport

x

x

Dams x

x

Residential/commercial

Settlements

x

Urban/suburban

x x x

x

Note:An x indicates hat landuse

type

s

likely

o nvoke he

pecified

cological

mechanism

and

to influence

cosystem

unction nd

biodiversity

ithin

rotected

reas.

ing

of the socioeconomic

dynamics

in

the

region

(DeFries

et al.

2007),

the

mechanisms

ffer

compre-

hensive

pproach

for

nderstanding

nd

managing

hese

vitallymportant egions

o maintain

cological

function

while

minimizing

negative impacts

on

surrounding

human

ommunities.

Vulnerabilityfprotected

reas to

surrounding

and

use

With imited esources

or

onservation,

t s

necessary

to identifyhich rotectedreas are mostvulnerableo

land use

change

so that

mitigation

trategies

an be

focused

n these reas

Wilson

t al.

2005).

Three lasses

of factorsthat

may

influence he

vulnerability

f a

protected

area

to land use intensification

re: the

ecological properties

f

the

protected

rea and sur-

rounding

cosystem;

he

type

and rates of land use

conversion

nd

intensification;

nd the

properties

f the

surrounding

uman communities.

We

suggest

hat the

most vulnerable

rotected

reas

will be those with he

following

haracteristics.

1)

The

protected

rea

is small or

poorly placed

relative o

minimum

ynamic

rea of

disturbance;

hape

of

species-area

curves;

biophysicalgradients

nd the

resulting

reas

of

organism

movements;

nd watershed

size.

2)

The

protected

rea is

in

close

proximity

o dense

human

populations,

ntense and use

in

critical

ortions

of the

cosystem,

r is

likely

o come n close

proximity

underfuture

and use

change.

3)

The

surrounding

uman

community

acks incen-

tives or resources

for

forwarding

cological

goals

of

protected

reas.

The

first

oint

follows rom

he

fact

that

the

spatial

extent

f

ecosystem rocesses

may

differ rom

lace

to

place.

For

example, patial

patterns

f

precipitation

ay

determine

whether

rganisms

migrate

over small

or

large

areas.

Thus,

the

key

to

assessingvulnerability

f

protected

reas

s to

evaluatenot the

bsolute ize of the

area,

but its size relative o

the effective

cosystem

t

exists within.

Quantitative

ssessmentof the

spatial

extent f

biophysical

actors,

ydrologic

lows,

istur-

bance,

and

organism

movements elative

o size and

location of the

protected

rea

provides

context

for

assessing

he

vulnerability

f the

protected

rea to land

use

change

n

the

unprotected ortion

f the

cosystem.

The

intensity

nd

type

of land

use

in

surrounding

lands also

differs

among

protected

areas. Parks

surrounded

y

intense and

uses,

such as

urban and

suburban,

are more vulnerable than

those set in a

wilderness ontext.

Also,

land use

types

differn

their

likely

nfluence n

protected

reas. Land uses such

as

commercial

armingmay

elicit all four of the

mecha-

nisms

previously

described,

whereas others such as

tourism r

poaching may

involve

single

mechanism

(Table 4).

The socioeconomic fabric of surroundinghuman

communities

robably

lso influences he

vulnerability

of

protected

reas. Protected reas located in areas

where

surrounding

ommunities

ely

on bushmeator

forest

products

are

likely

to be more vulnerable.

Protected reas also

vary

n

theenforcementf

policies

to

protect

eserves

Bruner

t al.

2001).

For

example,

elephantpopulations

fared better

n

African

ountries

thatwere ble to

control

oaching rior

o the

vory

an

in

the 1980s

Leakey

and Morell

2001).

Protected reas

surrounded

y

human

communities hat benefit

rom

them

may

be less vulnerable ue to a

higher

ikelihood



986 INVITED FEATURE


Vol.

17,

No.

4

of

regional-scale

management

o forward he

goals

of

the

protected

rea

(Rasker

and Hansen

2000).

Finally,

we

suggest

hat he onfluence

f

these

actors

has

a

larger

effect n the

protected

rea than

the

additiveeffect f the individualfactors.

With further

refinementnd

testing,

uch criteria ould

provide

a

basis forevaluating he global network f protected

areas and for

identifying

hose that

are

the

highest

priority

f conservation

ttention,

ased on

vulnerabil-

ity

to

land

use

change.

Within

regional

ontext,

he

principles

lso

can

be

applied

to

strategic

and use

management

o conserve lements

f the

andscape

most

crucial to reserve

unction,

hile

allowing

and use to

fulfill uman needs on those

portions

f

the

andscape

less crucial o

the

functioning

f reserves.

The case studies in this Invited Feature

provide

examples

of

regional

assessments and

management

implications

round several

protected

reas

in

varying

ecological

and socioeconomic

ettings.

he

concluding

paper by DeFries et al. (2007) explores n detail the

interactions

etween

protected

reas and

local

people

and

opportunities

or

chieving egional-scalemanage-

ment.

Acknowledgments

We thank he uthors nthe nvited eaturefor

timulating

discussions n nature eserve ssues.This workwas

supported

by

the NASA Land

Cover Land

Use

Program

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