42

Conservation Biology, Pages 42–57Volume 16, No. 1, February 2002

Ecoregions in Context: a Critique with Special Reference to Indonesia

PAUL JEPSON* AND ROBERT J. WHITTAKER

School of Geography and the Environment, University of Oxford, Mansfield Road, Oxford OX1 3TB,United Kingdom

Abstract:

World Wildlife Fund–United States ( WWF ) is promoting an ecoregional framework internation-ally as a new hierarchical approach to organizing and prioritizing conservation efforts. We assessed WWFecoregions against existing frameworks: (1) the Dasmann-Udvardy ( World Conservation Union [IUCN]) Bio-geographical Representation Framework, (2) the Bailey Ecoregional Framework (U.S. Forest Service), and(3) the hotspot approach, as exemplified by the BirdLife Endemic Bird Area Approach and the WWF–IUCNCentres of Plant Diversity Program. We examined the genealogy of the schemes from three perspectives: meth-odological explicitness, transparency and repeatability, and whether the WWF–ecoregions system improveson existing schemes. We considered Indonesia as a case study and assessed the efficacy of each system in theIndonesian context. The existing planning frameworks achieved their objective; in general had explicit, trans-parent, and repeatable methods; and, in the case of the Dasmann-Udvardy system, attained an institutionalreality in Indonesia. The central purpose of the WWF–ecoregions framework is the same as the 25-year-oldDasmann-Udvardy system, and at the coarsest spatial scales it relies on similar spatial delineators ( biomesand faunal regions). The WWF methodology, however, employs a gestalt approach to defining ecoregionboundaries. In the Indonesian context the resulting map appears problematic both in terms of the underlyingrationale of the ecoregion approach and in terms of apparent conflict with preexisting protected-area design.We suggest, insofar as refined planning frameworks are needed, that an alternative route that builds onrather than competes with existing approaches would be to combine at the mesoscale the landform delinea-tors that characterize the Bailey ecoregion system with the existing macroscale ecoclimatic and biogeographicdelineators of the Dasmann-Udvardy system. We question the investment in developing and promoting theWWF–ecoregion scheme in Indonesia when the existing Dasmann-Udvardy system, used in conjunction withhotspot studies, provides a seemingly adequate system and when the reserve system itself is under consider-able pressure.

Ecoregiones en Contexto: una Critica con Especial Referencia a Indonesia

Resumen:

El Fondo Mundial para la Vida Silvestre ( WWF) de los Estados Unidos está promoviendo interna-cionalmente un marco de trabajo ecoregional como una nueva aproximación jerárquica a la organizacióny priorización de los esfuerzos de conservación. Evaluamos las ecoregiones de WWF contra marcos de tra-bajo existentes: 1) el marco de trabajo de representación Biogeográfica de la Dasmann-Udvardy (UniónMundial para la Conservación (IUCN), el marco de trabajo ecoregional Bailey (Servicio Forestal de USA) y3) la aproximación de la regiones problemáticas ejemplificado por la estrategia de Áreas para Aves Endémi-cas y el programa de centros para la diversidad de plantas de WWF/IUCN. Examinamos la genealogía de losesquemas desde tres perspectivas: nivel de claridad en la metodología, transparencia y repetibilidad y si elsistema de ecoregiones de la WWF mejora los esquemas existentes. Consideramos a Indonesia como un casode estudio y evaluamos la eficacia de cada sistema en el contexto de Indonesia. Los planes de trabajo exis-tentes alcanzaron su objetivo; en general tuvieron métodos explícitos, transparentes y repetibles; y en el casodel sistema Dasmann-Udvardy alcanzó una realidad institucional en Indonesia. El propósito central de losmarcos de trabajo de las ecoregiones de WWF es el mismo que el sistema de hace 25 años de Dasmann-Udvardy y a nivel de escalas espaciales amplias está basado en delineadores espaciales similares ( biomasa y

*

email paul.jepson@geog.ox.ac.ukPaper submitted September 17, 1999; revised manuscript accepted May 9, 2001.

Conservation BiologyVolume 16, No. 1, February 2002

Jepson & Whittaker Ecoregions in Context

43

regiones faunísticas). Sin embargo, la metodología de WWF emplea una metodología de configuración de el-ementos separados (gestalt) para definir los límites de las ecoregiones. En el contexto de Indonesia, los mapasresultantes parecen ser problemáticos tanto en términos de la racionalidad subyacente de la metodología deecoregión y en términos de un aparente conflicto con los diseños de las áreas protegidas existentes. Sugerimosque hasta el momento se necesitan planes refinados de marco de trabajo. Una ruta alternativa que con-struya, y no que compita con las metodologías existentes sería la combinación a nivel de mesoescala de losdelineadores de contornos que caracterizan el sistema de ecoregión Bailey con la macroescala ecoclimáticaexistente y los delineadores biogeográficos del sistema Dasmann-Udvardy, usados en conjunción con los estu-dios de regiones problemáticas. Esto provee un sistema aparentemente adecuado cuando el sistema de reser-

vas se encuentra bajo una considerable presión.

Introduction

Over the last 30 years, a variety of spatial frameworkshas been developed for the purpose of guiding conser-vation action internationally. The most recent is theecoregional approach developed by World WildlifeFund–United States ( WWF) (Dinerstein et al. 1995; Ol-son & Dinerstein 1998). It is being adopted and pro-moted widely by the WWF family of agencies (WWF In-ternational, WWF national organizations, and WWFcountry representative offices) and by international pro-grams of the U.S.–based, nongovernmental organizationThe Nature Conservancy (TNC). The combination of ad-vocacy power, human and financial resources, and inter-national project portfolios possessed by these two inter-national organizations make it likely that the ecoregionalframework will be adopted and used by other agencies,including the Global Environment Facility (GEF ) andgovernment conservation agencies in developing coun-tries. In other areas of natural resource management,misunderstanding of alternative spatial planning frame-works has resulted in inconsistency in their use and ulti-mate effectiveness (Omernik & Bailey 1997). Those in-volved in conservation planning on the ground thereforeneed to know what this scheme brings with it that pre-existing schemes do not.

Our review has three aims: (1) to compare the WWFecoregions with existing spatial frameworks for terres-trial conservation planning with a global perspective; (2)to assess scientific explicitness, transparency, and re-peatability of methods; and (3) to ask whether the WWFecoregions framework improves upon existing frame-works. To address these points we reviewed and at-tempted to define the purpose for which the variousschemes were devised.

We considered Indonesia as a case-study country withwhich to explore these questions. Indonesia is a suitablechoice because (1) it is one of the most biodiverse areason Earth; (2) the government has consistently been atthe forefront in adopting new spatial conservation plan-ning frameworks; and (3) as an archipelago comprisinglarge continental-shelf islands and oceanic islands, Indo-

nesia captures a wide range of biotic variation (it spanstwo zoogeographical regions, dry and ever-wet tropics,and its ecosystem variation ranges from tropical glacierto mangrove).

To meet the aims outlined above, we describe (inchronological order) the three most prominent catego-ries of spatial planning frameworks developed at the glo-bal scale: biogeographical provinces, hotspots, andecoregions. We summarize each framework with re-spect to their aims, rationale, and context of developmentand then assess their efficacy with reference to Indonesia’sterrestrial ecosystems. Our assessment is concerned prin-cipally with generic issues rather than specific boundaryquestions in Indonesia. As far as we have been able toestablish, no comparative overview of these differentschemes has been published, and we therefore hopethis contribution will stimulate debate among conserva-tionists in general, not merely those directly involved inplanning within the Indo-Malayan realm.

Biogeographical Representation

The Dasmann-Udvardy Framework

A central concern of the IUCN since its creation has beenthe need to establish a worldwide network of natural re-serves encompassing representative areas of the world’secosystems. In the 1960s there was widespread supportfor this “representation principle.” In response, Dasmann(1972, 1973) prepared for IUCN a hierarchical system thatdefines and classifies natural regions for the purpose ofconservation. His aim was to provide a system that gaveequal emphasis to the IUCN’s interests in conservingnatural ecosystems and vegetation types and the conser-vation of species. His solution was to establish a systemof classification of communities based on ecoclimatic fea-tures but emphasizing taxonomic differences (Table 1).

At the top level in the hierarchy, Dasmann (1972) chosethe biome system (e.g., tundra, taiga, deciduous broad-leaved forest) of Clements and Shelford (1939) becauseit is readily applicable globally, takes into account both

44

Ecoregions in Context Jepson & Whittaker

Conservation BiologyVolume 16, No. 1, February 2002

Tabl

e 1.

Com

para

tive

anal

ysis

of f

our

spat

ial c

onse

rvat

ion

plan

ning

fram

ewor

ks.

Bio

geogr

aph

ic p

rovi

nce

E

ndem

ic b

ird a

rea

s (E

BA

s)

a

Eco

regi

on

s W

WF

ecore

gion

s

Dev

elo

pin

g ag

ency

Wo

rld

Co

nse

rvat

ion

Un

ion

Bir

dLi

fe I

nte

rnat

ion

alU

. S. F

ore

st S

ervi

ceW

orl

d W

ildlif

e Fu

nd

–U.S

.Le

ad a

uth

ors

R. F

. Das

man

n, M

. D. F

. Ud

vard

yC

. J. B

ibb

y, A

. J. S

tatt

ersf

ield

, M

. J. C

rosb

yR

. G. B

aile

y, C

. Om

ern

ik

D. M

. Ols

on

, E. D

iner

stei

n

Gen

eral

go

alco

nse

rve

glo

bal

hab

itat

an

d

spec

ies

div

ersi

tyid

enti

fy a

reas

ric

hes

t in

u

niq

ue

kin

ds

of

org

anis

m

for

pri

ori

ty a

ssig

nm

ent

of

con

serv

atio

n a

ctio

n

opti

mal

man

agem

ent o

f lan

d, d

efin

ed a

s en

suri

ng

that

all

lan

d u

ses

coin

cid

enta

lly s

ust

ain

res

ou

rce

pro

du

ctiv

ity

and

mai

nta

in e

cosy

stem

p

roce

ss a

nd

fu

nct

ion

pro

mo

te c

on

serv

atio

n o

f te

rres

tria

l, fr

esh

wat

er, a

nd

mar

ines

eco

syst

ems

har

bo

rin

g gl

ob

ally

imp

ort

ant

bio

div

ersi

ty a

nd

eco

logi

cal p

roce

sses

Op

erat

ion

al p

urp

ose

guid

e es

tab

lish

men

t of w

orl

dw

ide

net

wo

rk o

f n

atu

ral r

eser

ves

des

ign

ate

and

/or

stre

ngt

hen

m

anag

emen

t o

f p

rote

cted

ar

eas

in E

BA

s

Bai

ley:

ass

ist l

and

man

ager

s to

mee

t th

is

goal

; Om

ern

ik: e

ffec

tive

m

anag

emen

t o

f w

ater

qu

alit

y

sup

po

rt t

wo

-pro

nge

d s

trat

egy

of

esta

blis

hin

g p

rote

cted

are

as a

nd

ac

hie

vin

g su

stai

nab

le m

anag

emen

t in

th

e n

on

-res

erve

mat

rix

Key

des

ign

co

nsi

der

atio

ngi

ve e

qu

al s

tres

s to

str

uct

ura

l an

d

tax

on

om

ic d

iffe

ren

ces

of

eco

syst

ems

sele

ct a

mea

nin

gfu

l sca

le

dev

elo

p a

sys

tem

th

at c

lass

ifie

s la

nd

as

an

inte

grat

ed e

nti

ty b

ut

is s

till

suit

able

for

mul

tip

urp

ose

app

licat

ion

s

mo

ve a

way

fro

m o

rgan

izin

g co

nse

rvat

ion

wo

rk o

n t

he

bas

is o

f ge

op

olit

ical

bo

un

dar

ies

to p

lan

nin

g w

ith

in e

colo

gica

lly d

eriv

ed a

reas

Dat

es o

f dev

elo

pm

ent

1970

–197

519

89–1

992

1985

–199

619

91–p

rese

nt

Ap

pro

ach

b

glo

bal

(le

vel I

)

�

10

7

km

2

bio

geo

grap

hic

rea

lm (

regi

on

) cl

imat

ic &

tax

on

om

ic: b

iom

es

(Cle

men

ts &

Sh

elfo

rd 1

939)

su

bd

ivid

ed b

y fa

un

al r

egio

ns

(Wal

lace

187

6)

do

mai

n (

Bai

ley

) cl

imat

ic: f

ou

r zo

nes

ge

ner

ated

by

ove

rlay

ing

iso

ther

m

pat

tern

s (J

ames

195

9)

and

mo

istu

re

limit

s (S

cho

tt, i

n J

ames

195

9)

bio

geo

grap

hic

al z

on

e &

maj

or e

cosy

stem

ty

pe

tax

on

om

ic a

nd

eco

clim

atic

: m

ajo

r ec

osy

stem

typ

e (D

iner

stei

n e

t al.

1995

) d

isp

ense

d w

ith

in s

ub

seq

uen

t sc

hem

es; n

o c

lass

ific

atio

n c

ited

bu

t b

ioge

ogr

aph

ic z

on

es li

sted

fo

r La

tin

an

d N

ort

h A

mer

ica

(Din

erst

ein

et

al.

1995

; Ric

kett

s et

al.

1999

) eq

uiv

alen

t to

Wal

lace

(18

76)

Mac

ro e

cosy

stem

(lev

el I

I)10

6

(10

4

–10

8

) km

2

biog

eogr

aph

ical

(bi

otic

) p

rovi

nce

(7 d

ivis

ion

s) e

co-c

limat

ic &

ta

xo

no

mic

: clim

ax v

eget

atio

n

(Wea

ver

& C

lem

ents

193

8)

sub

div

ided

by

per

cen

t fa

un

al

sim

ilari

ty (

Hag

mei

er &

Stu

lts

1964

)

ecor

egio

n d

ivis

ion

s &

pro

vin

ces

(Bai

ley)

, le

vel I

I ec

ore

gio

n (

Om

ern

ik)

clim

atic

, fo

llow

ing

Kö

pp

en (

1931

) an

d T

rew

arth

a (1

968

) cl

imat

e cl

assi

fica

tio

n s

yste

ms

and

do

min

ant

po

ten

tial

veg

etat

ion

(K

üch

ler

1964

, 19

70)

maj

or

hab

itat

typ

e (9

div

isio

ns)

clim

atic

, m

od

ifie

d b

y o

ther

bio

ph

ysic

al

char

acte

rist

ics:

in L

atin

Am

eric

a fo

llow

s va

riou

s pre

limin

ary

sch

emes

(Din

erst

ein

et a

l. 19

95),

in N

ort

h A

mer

ica

follo

ws

Kü

chle

r (1

975)

, in

In

do

nes

ia f

ollo

ws

Wh

itm

ore

(19

84)

bas

ed o

n v

an S

teen

is

(195

7)

con

tin

ued

Conservation BiologyVolume 16, No. 1, February 2002

Jepson & Whittaker Ecoregions in Context

45

plants and animals, and broadly conforms to observablereality in areas not greatly modified by humans. Becausethe biome approach emphasizes ecological similaritiesat the expense of taxonomic difference, Dasmann(1972) divided the biomes of the world into regionalsubdivisions based on Wallace’s (1876) faunal regions(e.g., Palearctic, Ethiopian, Nearctic) and additional tran-sitional areas and biotic subdivisions that had long beenaccepted by biogeographers. These he termed “bioticrealms” (e.g., Indo-Malayan realm). Macro-scale (level II)units were termed “biotic provinces” and delineated bysubdividing a physiognomically defined climax vegeta-tion type at the level of the vegetation formation ofWeaver and Clements (1938) on the basis of a distinctivefauna (Dasmann 1973). Faunal distinctiveness is assessedby comparing the number of species in common be-tween areas divided by barriers that could have someconceivable distributional significance. Based on a re-view of various North American schemes ( Dasmanncites Goldman & Moore 1945; Blair 1950; Miller 1951;Hall & Kelson 1959; Hagmeier 1966) Dasmann consid-ered areas with 65% of their species in common to beseparate faunal provinces. Sixty-five percent is arbitrary,but because it is about two-thirds of the total speciescompliment, it constitutes a simple fraction of intuitivevalue. Dasmann (1972) recognized high mountains andmountainous islands (azonal features) as special situa-tions because vegetation and biota are likely to changemarkedly within short distances due to steep environ-mental gradients. Arbitrarily, he defined mountain rangesand island groups (e.g., Lesser Sundas) as separate bioticprovinces embedded within the system of provinceboundaries derived from his zonal methodology. His pro-visional list of biotic provinces (including Australisia andthe Antarctic) totaled 198.

To his chagrin, Dasmann (1973) found that the bio-geographer Udvardy (1969) had already published a de-tailed review of statistical methods for distinguishing bi-otic provinces, and that Hagmeier and Stults (1964) andHagmeier (1966) had made more exhaustive compari-sons than his but had arrived at the same conclusion inadopting the

�

65% similarity criterion. The IUCN thencommissioned Udvardy to develop and refine Das-mann’s system. Udvardy adjusted Dasmann’s terminol-ogy so that the highest level (biotic region) became the“biogeographic realm” and the second level (biotic prov-inces) became “biogeographic provinces.” In substance,Udvardy’s report (1975) was a reaffirmation of the scien-tific merits of Dasmann’s system. The resulting frame-work we call the Dasmann-Udvardy system.

Assessment of Biogeographic Representation in the Indonesian Context

The development of the biogeographic province frame-work coincided with the implementation in Indonesia of a

Tabl

e 1.

(Con

tinue

d)

Bio

geogr

aph

ic p

rovi

nce

E

ndem

ic b

ird a

rea

s (E

BA

s)

a

Eco

regi

on

s W

WF

ecore

gion

s

Mes

o e

cosy

stem

(l

evel

III

)10

3

(10

2

–10

7

) km

2

bio

geo

grap

hic

al u

nit

(b

iou

nit

) (4

0 d

ivis

ion

s) ta

xo

no

mic

: sam

e p

erce

nt

fau

nal

sim

ilari

ty

algo

rith

m a

pp

lied

by

Mac

Kin

no

n &

Win

d (

1981

) to

sm

alle

r ge

ogr

aph

ical

un

its

(Fig

. 1a)

EBA

(24

un

its)

tax

on

om

ic:

ove

rlay

dis

trib

uti

on

s o

f b

ird

sp

ecie

s w

ith

ran

ges

of

�

50,0

00 k

m

2

(T

erb

org

h &

W

inte

r 19

83)

lan

dsc

ape

mo

saic

(B

aile

y), l

evel

III

ec

ore

gio

n (

Om

ern

ik)

lan

dfo

rm

(geo

logy

an

d t

op

ogr

aph

y) in

form

ed

by

Ham

mon

d’s

(195

4, 1

964

) lan

dfor

m

clas

sifi

cati

on

sch

eme;

Om

ern

ik d

ivi-

sio

ns

also

info

rmed

by

lan

d-u

se

pat

tern

(A

nd

erso

n 1

970)

an

d v

ario

us

soils

map

s

eco

regi

on

(35

–40

div

isio

ns)

in N

ort

h

Am

eric

a ad

op

ts O

mer

nik

wit

h s

om

e m

od

ific

atio

ns;

in L

atin

Am

eric

a ad

op

ts

vari

ou

s n

atio

nal

sch

emes

; in

In

do

nes

ia

Wh

itm

ore

divi

sion

s an

d b

ioun

ts m

erge

d an

d m

od

ifie

d b

y EB

A b

ou

nd

arie

s

Exp

licit

nes

sex

celle

nt:

def

ined

pu

rpo

se, e

asy

to u

nd

erst

and

met

ho

do

logy

w

ith

sim

ple

alg

ori

thm

bas

ed o

nsp

ecie

s n

um

ber

dat

a, b

ut

anal

ysis

co

nfi

ned

to

bir

d a

nd

m

amm

al d

ata

goo

d: d

efin

ed p

urp

ose

, m

eth

od

s ea

sy t

o u

nd

er-

stan

d bu

t ba

sed

on a

rbit

rary

50,0

00-k

m

2

ran

ge c

rite

rio

n

and

ass

um

es m

eso

scal

e co

ngr

uen

ce w

ith

oth

er ta

xa

adeq

uat

e: d

efin

ed p

urp

ose

bu

t d

iffi

cult

to m

easu

re; m

eth

ods

diff

icul

t to

unde

r-st

and;

rob

ust

clas

sific

atio

ns

com

bin

ed,

bu

t d

ecis

ion

s o

n d

ivis

ion

s b

ased

on

su

bje

ctiv

e gr

ou

nd

s

limit

ed: p

urp

ose

all-e

mbr

acin

g; in

reg

ion

s w

ith

no

ex

isti

ng

sch

eme

(i.e

. ou

tsid

e N

ort

h A

mer

ica)

co

nd

uct

s ge

stal

t sy

nth

esis

of v

ario

us s

chem

es a

nd

crit

eria

a

A g

loba

l hots

pot

appro

ach

for

pla

nts

ha

s bee

n d

evel

oped

by

the

Worl

d C

on

serv

ati

on

Un

ion

an

d W

orl

d W

ildli

fe F

un

d i

n t

hei

r C

entr

es o

f P

lan

t D

iver

sity

Pro

gra

m. T

o d

ate

th

is h

as

ha

d li

mit

edef

fect

an

d t

her

efore

is

not

incl

uded

in

th

is t

able

.

b

Sum

ma

ry o

f te

rmin

olo

gy, h

iera

rch

ica

l det

erm

ina

tors

, an

d f

ou

nda

tion

stu

die

s a

t th

ree

spa

tia

l sc

ale

s.

46

Ecoregions in Context Jepson & Whittaker

Conservation BiologyVolume 16, No. 1, February 2002

two-phase (1974–1982) Food and Agriculture Organiza-tion (FAO) project that helped Indonesia establish nationalparks and expand the protected-area network (Blower1973; FAO 1977; Sumardja 1981). The national conserva-tion plan that was prepared to guide this process (MacKin-non & Artha 1982) was the first application of the globalDasmann-Udvardy system on a national scale.

In practice, it was realized that the biogeographicprovinces were at too coarse a scale to capture the bio-geographic variation in Indonesia, particularly in thetransitional island region of Wallacea (bridging the Ori-ental and Australasian faunal realms). In response, Mac-Kinnon and Wind (1981) applied the Dasmann-Udvardyalgorithm based on smaller geographic units to distin-guish between main regions within the larger islandsand island groups. This created a third tier in the hierar-chy, which was termed the biogeographical unit or “bio-unit” (Table 1; Fig. 1).

The national conservation plan for Indonesia (MacKin-non & Artha 1982) proposed a representative networkof reserves based on the following principles: (1) the

major biogeographic regions in the country and repre-sentative systems of reserves in each are identified; (2)within each biogeographic division, priority is given toestablishment of a major ecosystem reserve to includecontinuous habitat types and, if possible, the richest ex-ample of those habitats; (3) these large reserves are aug-mented with smaller reserves to protect special orunique additional habitat types or to cover regional vari-ations; and (4) small reserves are included to protectspecific sites of special beauty or interest. MacKinnonwas subsequently commissioned by the IUCN to pre-pare similar reviews for the Indo-Malayan and Afrotropi-cal realms, wherein he applied the same biogeographicspatial system and reserve design principles (MacKinnon& MacKinnon 1986

a

, 1986

b

). These principles are re-tained in the latest review of Indo-Malayan protected-area systems (MacKinnon 1997).

The subdivision into biounits further emphasizes taxo-nomic differences, but at the reserve selection level thisis balanced with MacKinnon and Artha’s (1982) secondand third design principles, which emphasize habitat

Figure 1. Dasmann-Udvardy biogeographic provinces for Indonesia, with the third-level “biounits” added to the sys-tem by MacKinnon and Wind (1981) overlain on the natural habitat-type boundaries redrawn with permission from MacKinnon (1997). Biogeographic provinces and biounits: 21, Sumatra (21a, south Sumatra; 21b, north Sumatra; 21c, Mentawi Islands; 21d, Nias and Batau islands; 21e, Simeuleu Islands; 21f, Enggano Island; 21g, Lingga Archipelago); 22, Java (22a, West Java; 22b, east Java; 22c, Bali Island); 25, Borneo (25b, southwest Borneo; 25e, central mountains; 25f, east Borneo; 25g, east Borneo; 25h, northwest Borneo); 24, Sulawesi (24a, central Su-lawesi; 24b, north Sulawesi; 24c, south Sulawesi; 24d, southeast Sulawesi; 24e, northeast Sulawesi; 24f, Sangihe-Ta-laud Islands); 23, Lesser Sundas; (23a, north Nusa Tenggara; 23b, Sumba Island; 23c, Timor and Wetar islands; 23d, Tanimbar); 13, Moluccas (13a, n. Maluku Islands; 13b, Obi; 13c, Buru; 13d, Ceram & Ambon; 13e, Kai Islands); P3, New Guinea; (P3a, Aru Islands; P3b, Western Islands; P3c, Geevlink Bay islands; P3d, Vogelkop; P3e, northwest New Guinea; P3f, southwest New Guinea; P3g, Snow Mountains; P3h, Star mountains; P3l, Trans-Fly).

Conservation BiologyVolume 16, No. 1, February 2002

Jepson & Whittaker Ecoregions in Context

47

representation (above). This is consistent with the aim ofthe Dasmann-Udvardy system to give equal emphasis tostructural and taxonomic features. Subsequent pro-tected-area reviews and biodiversity conservation strate-gies in Indonesia have adopted the same system and haveconfirmed the network of 80 key reserves to meet repre-sentation goals (Regional Physical Planning Programmefor Transmigration 1990; Government of Indonesia 1991;BAPPENAS 1993; KLH 1993). Since commencement ofthe FAO conservation project in 1974, Indonesia’s pro-tected-area network has been expanded from 170 re-serves covering 3.3 million ha (Sinaga, unpublished data)to 384 reserves covering 22.3 million ha ( PHPA 1999);with the exception of East Kalimantan, the Moluccas,and Nusa Tenggara, most of the 80 key reserves havebeen designated. Furthermore, it is only now, 20 years af-ter the design of the network, that forest loss in the fewbiounits still without reserves necessitates significant re-visions to the reserve configurations originally proposed.Many areas of economically valuable lowland habitatswere excised from proposed reserves at the time ofgazettement, but we consider this a reflection of the ten-dency of all governments to allocate land with limited ag-ricultural potential for biodiversity conservation, ratherthan a weakness with the system or prioritization per se.Thus, in Indonesia, at least, the Dasmann-Udvardy systemhas worked (Table 2). This assessment sets aside the vitalissues of and threats to reserve integrity.

The Dasmann-Udvardy system has the merit of a trans-parent and repeatable methodology with a genealogythat goes back to such authorities as A. R. Wallace andF. E. Clements. The delineation of biogeographic prov-inces and units is open to review as distributional datasets on faunal groups other than birds and mammals arecompleted, or after changes in taxonomy. Furthermore,the method provides for finer-scale subdivisions (Mac-Kinnon & Wind 1981).

The seven Dasmann-Udvardy biogeographic provincesof Indonesia accord with the main geographic, cultural,and economic developmental regions of Indonesia, andthey have been adopted widely as a framework for un-derstanding biological variation throughout the archipel-ago. They are taught in schools, they define coverage ofvolumes in the ecology of Indonesia series ( Whitten etal. 1987, 1996, 2000; MacKinnon et al. 1996; Monk et al.1997 ), and they provide the planning units for severalstrategies and overviews and the geographic units in var-ious tourist guidebooks. In short, biogeographic prov-inces have attained a social reality in Indonesia (Searle1996 ) (Table 2).

Hotspot Approaches

During the late 1970s, two attributes of biodiversitystarted to attract particular attention: species richness(the number of species in an area) and endemism (the

number of species in an area that occur nowhere else),and there were many prognoses of an impending massextinction of species (Myers 1979; Ehrlich & Ehrlich1981; Wilson 1988). These prognoses were based inpart on the simple observation that deforestation in thetropics was progressing at a rapid rate and was linked tothe “relaxation effect.” The effect is predicated on Mac-Arthur and Wilson’s (1963, 1967 ) equilibrium theory ofisland biogeography and postulates that species numberwill inevitably re-equilibrate to a lower number if habitatarea is reduced and isolation of patches increased (re-viewed by Whittaker 1998).

The notion of an impending “extinction crisis” led to asensible and obvious desire to target scarce conserva-tion resources and to give priority to establishing newreserves first in regions that (1) are exceptionally rich inspecies and/or unique species and (2) are under threat.Myers’s (1988, 1991) “hotspot analysis” was the firstsuch global study (updated by Myers et al. 2000). Al-though at too broad a scale (18—now 25—hotspots inthe world) to be of much practical use (Long et al. 1996),the idea inspired more detailed studies, notably theBirdLife International Endemic Bird Area (EBA) ApproachInternational Council for Bird Preservation ([ICBP] 1992),the IUCN–WWF Centres of Plant Diversity (IUCN & WWF1994), and, recently, the Global 200 ecoregions (Olson& Dinerstein 1998).

The IUCN–WWF Centres of Plant Diversity Approach

The IUCN–WWF Centres of Plant Diversity Program (IUCN& WWF 1994) sought to identify sites around the worldof greatest importance for plant conservation. Criteriawere developed for selecting sites that (1) are obviouslyrich in species, (2) are rich in endemic species, (3) arethreatened and/or (4) contain a diverse range of habi-tats, (5) have a gene pool of species useful to humans,and (6 ) contain species adapted to particular edaphicconditions. Sites were nominated and criteria applied onthe basis of expert review.

The Centres of Plant Diversity scheme is neitherwidely known nor widely used in Indonesia. Its strengthlies in it is ability to pinpoint areas and features of con-servation importance with scattered (azonal ) distribu-tions—notably l imestone massifs—which zonalschemes, such as those described in this paper, do notidentify. The main weakness of the approach lies in thecomparative incompleteness and unevenness of plantdistributional data relative to those for mammals andbirds: botanists tend to have collected in places of easyaccess. As a result, sites identified may reflect collectingeffort rather than truly exceptional levels of species rich-ness or endemism (cf. Nelson et al. 1990), and reservescome out as priority sites because this is where peoplehave collected.

Tabl

e 2.

Effic

acy

of fo

ur s

patia

l con

serv

atio

n pl

anni

ng fr

amew

orks

in I

ndon

esia

.

Cri

teri

aB

ioge

ogr

aph

ic p

rovi

nce

En

dem

ic b

ird a

rea

s (E

BA

s)E

core

gion

sW

orl

d W

ildli

fe F

un

d e

core

gion

s

Po

licy

& d

evel

op

men

tcr

eden

tial

ses

tab

lish

ed f

ram

ewo

rk f

or

gove

rnm

ent

and

do

no

r co

nse

rvat

ion

str

ateg

y an

d p

olic

y d

ocu

men

ts

wel

l kn

ow

n a

mo

ng

spec

ialis

tb

iod

iver

sity

pla

nn

ers

and

in lo

cal

pla

nn

ing

auth

ori

ties

of

Mal

uku

and

Eas

t N

usa

Ten

ggar

a

sim

ilar

con

cep

ts a

nd

cla

ssif

icat

ion

su

sed

in d

elin

eati

on

of

ph

ysio

grap

hic

al t

ypes

an

d la

nd

su

itab

ility

fo

r d

evel

op

men

t (s

ee R

ePP

Pro

T 1

991)

term

gai

nin

g re

cogn

itio

n, b

ut f

ew

unde

rsta

nd

its

mea

nin

g or

uti

lity

Scie

nti

fic

cred

enti

als

bas

ic f

ram

ewo

rk f

or

teac

hin

g m

acro

leve

l eco

logi

cal v

aria

tio

n

in s

cho

ols

an

d

un

iver

siti

es

wel

l kn

ow

n a

mo

ng

the

few

sci

enti

fic

orn

ith

olo

gist

s an

d s

pec

ialis

t co

nse

rvat

ion

pla

nn

ers

reso

urc

e-m

anag

emen

t d

isci

plin

esfa

mili

ar w

ith

clim

atic

, veg

etat

ion

, an

d la

nd

form

var

iab

les

no

ne

yet

Pu

blic

aw

aren

ess

seve

n p

rovi

nce

s w

ell k

no

wn

am

on

g In

do

nes

ian

s an

dre

late

d t

o t

he

maj

or

isla

nd

an

d

hen

ce c

ult

ura

l gro

up

s

at a

n o

per

atio

nal

leve

l th

e m

essa

ge

of

un

iqu

e b

ird

s in

EB

As

is p

rovi

ng

a p

ow

erfu

l mea

ns

of

gen

erat

ing

po

pu

lar

inte

rest

in c

on

serv

atio

n

limit

ed u

nd

erst

and

ing

no

ne

yet

Tan

gib

le c

on

serv

atio

n

ou

tco

mes

nat

ion

al c

onse

rvat

ion

pla

n e

nsh

rin

ed

the

rep

rese

nta

tion

pri

nci

ple

in

nat

ion

al p

olic

y; g

over

nm

ent p

olic

y to

est

ablis

h a

“m

inim

um s

et”

of 8

0 m

ajor

eco

syst

em r

eser

ves;

nea

rly

200

new

res

erve

s co

veri

ng

20

mill

ion

ha

adde

d to

th

e ex

isti

ng

rese

rves

sys

tem

; des

ire

to s

upp

ort

and

par

tici

pat

e in

th

e ab

ove

pro

gram

led

to fi

ve m

ajor

in

tern

atio

nal

con

serv

atio

n N

GO

s es

tabl

ish

ing

pro

gram

s in

Indo

nes

ia

crea

ted

a p

olic

y aw

aren

ess

of

the

con

serv

atio

n im

po

rtan

ce o

f th

e sm

all i

slan

d r

egio

ns

of

Less

erSu

nd

as a

nd

Mal

uku

; mai

n f

acto

rin

loca

tio

n a

nd

/or

des

ign

atio

n o

f se

vera

l new

res

erve

s an

d in

p

rom

oti

ng

fiel

d r

esea

rch

to

fill

d

istr

ibu

tio

nal

lacu

nae

str

engt

hen

ed

loca

l pri

de

and

sen

se o

f p

lace

on

is

lan

ds

wh

ere

EBA

s ar

e p

art

of

con

serv

atio

n p

lan

nin

g fr

amew

ork

no

t ap

plic

able

no

ne

yet

of

any

sub

stan

ce; h

as

intr

od

uce

d n

ew g

ener

atio

n o

f co

nse

rvat

ion

ists

fro

m s

oci

al o

r ec

on

om

ic b

ackg

rou

nd

s to

n

oti

on

s o

f ec

olo

gica

l rep

re-

sen

tati

on

; in

stru

men

tal i

n

exte

nd

ing

spat

ial p

rio

riti

zati

on

ap

pro

ach

to

th

e m

arin

e re

alm

Ove

rall

asse

ssm

ent

ach

ieve

d p

urp

ose

to

a r

emar

kab

le

exte

nt

ach

ievi

ng

pu

rpo

se a

nd

pro

vid

ing

use

ful a

dju

nct

to

th

e b

ioge

ogr

aph

ic

pro

vin

ce a

pp

roac

h

no

t ap

plic

able

app

ears

to

ad

d n

oth

ing

new

an

d

cou

ld c

om

pro

mis

e go

als

by

un

der

min

ing

the

esta

blis

hed

ap

pro

ach

es a

lrea

dy

adva

nce

d

in m

eeti

ng

com

par

able

go

als

Ass

essm

ent

of

futu

re v

alu

ep

rin

cip

le o

f re

pre

sen

tati

on

, co

nse

rvat

ion

’s jo

int

con

cern

s fo

r p

rese

rvat

ion

of

eco

syst

em t

ypes

an

d s

pec

ies

vari

atio

n, a

nd

cr

eati

on

of

rese

rves

to

mee

t th

ese

con

cern

s is

ex

pec

ted

to

st

and

th

e te

st o

f ti

me;

Das

man

n-

Ud

vard

y sy

stem

a g

oo

d

hie

rarc

hic

al s

yste

m f

or

con

serv

atio

n p

lan

nin

g d

ow

n t

o

the

mes

oec

osy

stem

sca

le

kno

wle

dge

of

loca

tio

n o

f m

eso

- or

mic

roec

osys

tem

cen

ters

of e

nde

mis

m

is li

kely

to

incr

ease

in v

alu

e in

lin

e w

ith

incr

easi

ng

lan

dsc

ape

chan

ge

and

hu

man

po

pu

lati

on

gro

wth

; h

ots

po

t ap

pro

ach

es s

uch

as

EBA

s ar

e es

sen

tial

su

pp

lem

ents

to

“r

epre

sen

tati

ve”

app

roac

hes

fo

r m

acro

scal

e co

nse

rvat

ion

pla

nn

ing

and

str

ateg

izin

g

lack

of

con

sid

erat

ion

giv

en t

o

spec

ies

vari

atio

n m

ean

s th

is

syst

em is

un

suit

able

fo

r b

iod

iver

sity

co

nse

rvat

ion

p

lan

nin

g at

glo

bal

leve

l to

m

eso

scal

e; f

ocu

s o

n la

nd

form

b

elo

w t

he

mes

osc

ale

off

ers

op

po

rtu

nit

y to

co

mb

ine

con

cern

s re

lati

ng

to s

oci

al ju

stic

e an

d e

colo

gica

l in

tegr

ity

wit

h

trad

itio

nal

hab

itat

an

d s

pec

ies-

pre

serv

atio

n c

on

cern

s in

co

nse

rvat

ion

; lan

dfo

rm c

ou

ld

form

a f

ou

rth

-leve

l det

erm

inat

or

in t

he

Das

man

n-U

dva

rdy

syst

em

may

be

ben

efic

ial i

n a

reas

of

the

wo

rld

lack

ing

det

aile

d

pre

exis

tin

g sc

hem

es, b

ut

we

qu

esti

on

th

e m

erit

of

pro

mo

tin

g th

is a

pp

roac

h o

ver

exis

tin

g sc

hem

es in

In

do

nes

ia

Conservation BiologyVolume 16, No. 1, February 2002

Jepson & Whittaker Ecoregions in Context

49

BirdLife International’s Endemic Bird Area Approach

Contemporaneous with the Centres of Plant Diversityprogram, BirdLife International developed its EndemicBird Area Approach ( ICBP 1992; Stattersfield et al. 1998),which is the most complete example of a global hotspotanalysis (Long et al. 1996). This approach is based on thebelief that “first priority must be assigned to programs toconserve areas richest in unique kinds of organism” ( Ehr-lich 1988) and was inspired by pioneering studies of birddistribution in Africa (Hall & Moreau 1962), Colombia,and Ecuador (Terborgh & Winter 1983). The last studymapped bird species with ranges of 50,000 km

2

(an arbi-trary size) to locate areas of concentrated endemism thatwould be optimal for designation as reserves. TheBirdLife Biodiversity Project applied Terborgh and Win-ter’s 50,000-km

2

range criterion worldwide. For bird spe-cies meeting this criterion (2649 species, following Longet al. 1996), project researchers conducted a compre-hensive literature review and compiled a database ofgeographically referenced distributional records. Recordswere plotted and distributions of species overlaid. Areaswhere two or more such species co-occurred (an arbi-trary choice) were termed endemic bird areas. In regionswith complex distributional patterns, a divisive clusteranalysis of multivariate distributional data summarized bygrid square was performed to aid in identification of “nat-ural” groupings of species (ICBP 1992; Long et al. 1996).Although this is a repeatable methodology, the use of ar-bitrary criteria (e.g., size of range, number of endemics,degree of range overlap), as with all such schemes, inevi-tably has a bearing on precisely which areas are selected.

Like the Dasmann-Udvardy system, the EBA approachhas the merit of an explicit purpose and a transparentand repeatable methodology. The database on whichEBA boundaries were devised is freely available, and themethod could be developed to identify centers of ende-mism at different taxonomic levels or (with greater ef-fort) using a different range-size threshold. This allowsconservation recommendations derived from EBAs to beindependently assessed, revised, or refined.

Assessment of Endemic Bird Areas in Indonesia

BirdLife originally identified 221 EBAs worldwide, 24 ofwhich are in Indonesia ( ICBP 1992; Sujatnika et al.1995). In response, BirdLife launched its Indonesia pro-gram in partnership with the Directorate General of For-est Protection and Nature Conservation (PHPA) in 1992,with the purpose of securing the designation of reservesproposed under the national conservation plan in prior-ity EBAs ( Jepson 1995). The EBAs are not designed in ahierarchical system. As biogeographic entities, however,EBA boundaries in Indonesia nest within biogeographicprovinces and accord closely with the MacKinnon andWind (1981) biounits ( Jepson & Sujatnika 1997). Many

are located in the complex island region of Wallacea. Toidentify priority islands and reserves for conservation,BirdLife identified sub-EBAs by island(s) and by habitatby creating simple matrices of species presence againstisland/island group and habitat type (Sujatnika et al. 1995).

The EBA approach assisted in implementation of thenational conservation plan ( based on the Dasmann-Udvardy system) by prioritizing the proposed reservessited in areas of concentrated endemism. Prior to the es-tablishment of the PHPA–BirdLife Indonesia Programme,conservation efforts in Indonesia were focused on thelarge land masses of Sumatra, Kalimantan, Java, Irian Jaya,and Sulawesi (mainly in Minahasa). The Lesser Sundaand Moluccas biogeographic provinces received little at-tention, despite their long-recognized scientific and con-servation importance (e.g., Harper 1945). Between 1992and 1999, and as a result of the BirdLife initiative, two newnational parks were designated on Sumba ( Jepson et al.1996), four new key reserves were designated on Timor,a major GEF–financed project was prepared to establishfive new reserves in Maluku (since cancelled because ofpolitical and social instability), and proposals for desig-nating new reserves in Flores and Sumbawa have beenprepared ( Trainor et al. 2000; P.J. et al., unpublisheddata). The program inspired 10 university biological ex-peditions to remote islands and the “Action Sampiri”conservation project on Sangihe-Talaud (Riley 1997). InIndonesia the EBA approach is succeeding in directingnew conservation effort to centers of avian endemism.

When EBAs are assessed against the hotspot ap-proach’s dual criteria of richness/endemism and threat,a weakness is evident: the areas mentioned in the preced-ing paragraph (e.g., Timor, Sumba, Flores) are less threat-ened than areas in west Indonesia that are not includedwithin EBAs. For example, with 164 endemic bird spe-cies ( Wells 1985), the lowland ever-wet forest of theSunda shelf constitutes a center of endemism, but manyof these species have ranges above the 50,000-km

2

threshold for EBAs. These forest ecosystems are beingconverted to agriculture and estate crops at an alarmingrate. On Sumatra, there is little intact lowland forest re-maining outside reserves (Laumonier 1997), and evenwithin reserves this habitat is being degraded by illegallogging and fire. Yet EBAs in west Indonesia are all inmountainous regions that are relatively intact and se-cure. The new reserve currently being advocated in In-donesia is Sebuku-Sembukang in East Kalimantan, butthis priority was identified on the basis of a simple analy-sis of gaps in representation of habitats and species of in-ternational conservation interest (Momberg et al.1998

a

), not with the EBA approach.Endemic bird areas were promoted as centers of unique

biodiversity and thus as indicators of likely areas of ende-mism in other taxa (ICBP 1992). Although congruence ofbird endemism with centers of endemism in other groupsgenerally obtains at the macroscale, it often fails at finer

50

Ecoregions in Context Jepson & Whittaker

Conservation BiologyVolume 16, No. 1, February 2002

scales of analysis (cf. Bush 1994). At the scale of nationalconservation planning in Indonesia, expert consultationsuggests that existing data support the notion of congru-ence only for some small-mammal groups (D. Kitchener,personal communication) and swallowtail butterflies (P.J.,unpublished data). In the experience of the senior author,overstating the scope of the approach in the early 1990swas counterproductive because this drew criticism fromother conservation specialists when the approach was pre-sented in conservation fora. As a result, policy makers in In-donesia were wary of its merits and came to perceive EBAsas a specific approach for endemic bird conservation. Thisgoal was perceived as a relatively unimportant componentof the wider Indonesian biodiversity discourse because en-demic birds lack a clear utility value.

Despite these shortcomings, the EBA approach becameestablished in Indonesia as a distinct system complimen-tary to the biogeographic-region system. The value ofEBAs lies in drawing attention to the existence of discretecenters of avian endemism and in generating local sup-port for reserve designation. This last point is worth elab-oration. In Indonesia, approval by the provincial governorand district officer is required for reserve designation toprogress. BirdLife employees have found that such offi-cials respond with pride and interest when informed thattheir territory supports an assemblage of bird speciesfound nowhere else on Earth, and this has been instru-mental in securing their support for the new reserves.

Ecoregional Approaches

The term

ecoregion

was introduced into the arena ofU.S. land-management planning and conservation by R. G.Bailey (e.g., 1983, 1996) and the U.S. Forest Service (1993)ECOMAP project, who developed and refined a hierar-chical system for the purpose of optimizing land-man-agement goals within the United States. Another impor-tant contributor to the development of the ecoregionconcept is J. M. Omernik (1987, 1995). Internationally,Bailey (e.g., 1989) extended his ecoregions approach tothe world at the macroecosystem level of resolution.The conservation science program of WWF–United Statessubsequently developed a mesoscale ecoregional classi-fication for Latin America (Dinerstein et al. 1995) andembarked on a project to develop ecoregion maps forthe rest of the world (Dinerstein 1999).

Ecoregions: Bailey and Omernik Frameworks

Unlike the Dasmann-Udvardy system, the Bailey (1996,1998) and Omernik (1987, 1995) hierarchical ecosystemclassification systems do not aim to incorporate taxonomicdistinctions but rather focus on characteristics of ecosys-tem structure. This reflects their purpose, namely the opti-mal management of land and water (Omernik & Bailey1997), defined as ensuring that all land uses coincidentally

sustain resource productivity and maintain ecosystem pro-cesses and functions.

Bailey’s system aims to delineate at a given level bound-aries of ecosystems that control the process and func-tion of ecosystems at the next level down. He adopts the“controlling factor method” (Bailey 1996 ) by which aspatial hierarchy is constructed by successive subdivi-sion of large ecosystems on the basis of controlling factorsoperating at different scales. Ecoregions are delineated atthree levels. At the macroscale, climate is considered theprincipal controlling factor. In the top tier, four domains,or ecoclimatic zones of the Earth, are delimited—humidtropical, humid temperate, polar, and dry—by simpleoverlay of global thermal and moisture patterns ( James1959 ). These domains are subdivided on the basis ofKöppen’s system of climatic classification (as modifiedby Trewartha (1968) into second-tier ecoregions, termeddivisions, of which there are 31 globally. Bailey’s divisionalsystem distinguishes between zonal and azonal ecore-gions. Azonal ecoregions are, for example, wetlands oralpine ecosystems that can occur in any zone where theappropriate geomorphology occurs; for instance, thezonal “icecap division” is matched by the azonal “icecapregime mountains.” At the macroscale, Indonesia hasfour divisions: savanna, savanna regime mountains, rain-forest, and rainforest regime mountains. The divisionsmay also in turn be subdivided into “provinces” on thebasis of macrofeatures of the vegetation that reflectmore refined climatic differences. This province level isreferenced but not presented by Bailey (1996).

Bailey (1996 ) proposes further refinements to hisscheme, as follows (Table 1). At the mesoscale, he con-siders landform the principal determinant of potentialvegetation and uses Hammond’s (1954, 1964) scheme ofland classification, informed by Küchler’s (1964, 1970)maps of potential vegetation in the United States to deter-mine the limits of various mesoecosystems, termed land-scape mosaics. This is the third level of the Baileyscheme. These are further subdivided into smaller micro-ecosystems based on edaphic factors. Finally, Bailey pro-poses dividing all contemporary ecosystems into fourclasses, reflecting the degree of human transformation,based on the system of Milanova and Kushlin (1993).

Omernik’s system is essentially the same. The same de-lineators are used at the macroscale to produce what heterms a level II ecoregion. In Omernik’s scheme, level IIIecoregions divisions are informed by land-use pattern(Anderson 1970) and various soil maps in addition to Ham-mond’s land-use and Küchler’s vegetation maps (Omernik1987). The decision criteria for combining these data lay-ers in both schemes are subjective ( Wright et al. 1998).

WWF Ecoregions

The WWF ecoregions are more inclusive in purpose thanthose of the Bailey and Omernik systems. The WWF ecore-

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gions aim to create a superior biogeographic unit for con-servation planning at regional scales to meet the four maingoals of biodiversity conservation, as defined by Noss(1992) and Noss and Cooperrider (1994): (1) representa-tion of all distinct natural communities within a network ofprotected areas, (2) maintenance of ecological and evolu-tionary processes, (3) maintenance of populations of spe-cies, and (4) conservation of large blocks of natural habitat.Ecoregion-based conservation (the practical application ofthe framework) aims to promote these goals by employinga “two pronged strategy of establishing protected areas andachieving sustainable management of the lands and watersoutside protected areas” (Ricketts et al. 1999).

The WWF ecoregion approach combines two ele-ments: (1) mapping of regional patterns of biodiversity(i.e., delineation of ecoregion boundaries) and (2) prior-itization of ecoregions for conservation action (Wikra-manayake et al. 2001). Here, we focus on the first ele-ment. The second element, a form of hotspot analysis,uses transparent and repeatable indices of biological dis-tinctiveness and conservation status (Dinerstein et al.1995; Ricketts et al. 1999; Wikramanayake et al. 2001) toidentify priority ecoregions known as the Global 200ecoregions (Olson & Dinerstein 1998; WWF 2000). Inseveral cases, ecoregions have been aggregated to createa single Global 200 ecoregion. The basis of aggregationis not evident in the literature cited.

An ecoregion is defined in the WWF scheme as “an eco-system of regional extent” (Dinerstein et al. 1995), whichwe take to mean a mesoscale ecosystem (10

2

–10

7

km

2

)that controls and is defined by smaller constituent ecosys-tems. The WWF ecoregion framework is conceived as “ahierarchy based on habitat types” (Dinerstein et al. 1995).Dinerstein et al. (1995, their Figs. 1 & 2) place major eco-system types at the top level. These are subdivided intomajor habitat types said to equate broadly with biomes(Olson & Dinerstein 1998) and are overlaid with biogeo-graphic realm (e.g., Nearctic, Indian Ocean) to delineatethird-level ecoregion boundaries. In subsequent studies(Ricketts et al. 1999, their Fig. 2.3; Wikramanayake et al.2001, their Box 2.1), the hierarchy is represented moresimply as biogeographic realm (they use the term

zone

)subdivided by major habitat type (biome). Confusingly,the Global 200 accounts (Olson & Dinerstein 1998; WWF2000) state the reverse, major habitat types subdivided bybiogeographic realm.

In practice, WWF ecoregions are delineated by com-bining boundaries of existing regional schemes. ForLatin America, various schemes were combined (Diner-stein et al. 1995); for North America, Omernik’s (1987 )ecoregion maps were adopted (Ricketts et al. 1999); forthe Asia-Pacific volume, MacKinnon’s biounit and origi-nal forest cover maps (Fig. 1) were combined ( Wikra-manayake et al. 2001), all with modifications. Modifica-tions to existing schemes are described in ecoregionalaccounts and were made on the basis of refinement of

“first cut” boundaries following consultation with re-gional experts.

The intent of the WWF ecoregion approach is to mapboundaries of ecosystems at the mesoscale. It employsmap overlay methods at the macroscale. At the meso-scale, it employs a gestalt approach in which regionalboundaries are drawn intuitively around areas that ap-pear homogenous (Bailey 1996), combining various ex-isting zonal and azonal schemes not necessarily with thesame genealogies. A key assumption at the mesoscale isthat boundaries of original vegetation types equate toboundaries within which key ecological flows and link-ages operate internally, as opposed to externally.

This is difficult to substantiate with certainty becausethe terminology adopted in the WWF ecoregion frame-work is vague. First, key ecobiogeographical terms—

biome

,

bioregion

,

major habitat type

—are not linked tofoundational definitions, which introduces a degree ofsubjectivity and confusion. Second, macroscale unitssuch as bioregion are frequently defined in terms ofamalgamation of ecoregions, which is counter to normalpractice in the field and introduces circularity. Third,new methodological terms are introduced but not de-fined, which obscures method. Use of the term

stratify

is a case in point.

Stratify

should refer to layering morethan once, but in WWF ecoregion discourse it means fit-ting clusters of ecoregion boundaries within boundariesof major habitat types.

Assessment of the WWF Ecoregions for Indonesia

Our assessment of WWF ecoregions is based on the In-donesia sections of Figs. 2.1.b and 2.1c of Wikramanay-ake et al. (2001), reproduced here as Fig. 2. This repre-sents a refinement of “first cut” ecoregion boundariesthat have circulated in Indonesia conservation-planningcircles since 1996 and were used by Yayasan WWF–Indonesia to reorganize and develop new strategic direc-tions (see Momberg et al. 1998

b

, their Map 1).At the time of their introduction to Indonesia, the Das-

mann-Udvardy and EBA schemes were the first of theirtype and guided new conservation programs. In con-trast, the WWF ecoregions represent an alternative schemeintended to improve the performance of existing conser-vation programs. Our assessment therefore focuses onwhether ecoregions compliment or improve upon exist-ing frameworks.

According to Wikramanayake et al. (2001), the ecore-gion delineation in the Asia Pacific is similar to the Das-mann-Udvardy hierarchy. MacKinnon’s (1997 ) maps ofbiounits and original forest vegetation were used as thegeneral guide, modified by EBA boundaries and the com-ments of three regional experts, A. J. Whitten, T. C.Whitmore, and D. Madulid. Comparison of Figs. 1 and 2shows that ecoregion boundaries correspond to a syn-thesis of MacKinnon (1997, his Map 2) in western Indo-

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nesia, and EBA boundaries (Fig. 2) in Wallacea and to alesser extent western New Guinea ( Irian Jaya). Thereare several modifications to this broad picture, differingin significance, that are described but not fully justifiedby Wikramanayake et al. (2001).

The aim of delineating mesoecosystems (as definedabove) is not consistently achieved in Indonesia, as illus-trated by three cases. (1) The 1.8 million-ha tectonic basincharacterized by the Middle Mahakam wetland system(Voss 1983) in East Kalimantan is unquestionably a dis-tinct mesoecosystem or ecoregion in the Bailey-Omerniksense. Yet Wikramanayake et al. (2001) divide it into acomplex of three ecoregions on the basis of dominantvegetation formations (heath forest, peat swamp forest,and freshwater swamp forests). These are constituent ec-osystems. (2) Two small island systems (Sangihe-Talaudand Bangai-Sula) are combined with areas below 1000 mon Sulawesi in a single ecoregion. It is inconceivable thatthere are distinct ecological flows and linkages internal toand controlling an area with this pattern. (3) Montaneecoregions are delineated on the basis of the 1000-m con-tour because it approximates a change in forest type.There are several problems with this: for example, the ac-tual elevation of forest-type change varies substantially asa function of the Massenerhebung (mass-elevation) ef-fect ( Whitmore 1984; Richards 1996); faunal assemblagesthat are the target of conservation action are not confinedin distribution to above or below a single elevationalthreshold (Md. Nor 2001); and factors controlling ecosys-tem function generally coincide with mountain landscapeunits rather than being bounded by an arbitrary elevation.

The use of major habitat type in Indonesian ecoregionnames is consistent with WWF’s desire to stress habitatrepresentation. In reality, WWF ecoregion boundariessimplify MacKinnon’s map (1997, Fig. 2) of vegetationtypes, particularly in the Wallacea region. MacKinnon’shabitat type boundaries (Fig. 1) are based on Whitmore’s(1984) vegetation formations. Spatially these may be (1)large, homogenous areas such as lowland everwet for-est, (2) smaller scattered patches such as limestone for-ests, or (3) linear features such as beach forests. Homoge-nous-area formations appear at macro- and mesoscales,scattered formations at the mesoscale, and some linearformations only at finer scales. As a result, WWF ecore-gions emphasize homogeneous major habitat types but ig-nore scattered and linear formations, or treat these asdistinct habitat types within an ecoregion. Moreover,Whitmore’s (1984) scheme is an ever-wet forest vegetationclassification: dry-tropical forest formations have not yetbeen systematically classified in Southeast Asia. Conse-quently, WWF ecoregions underrepresent dry-tropicalforest habitats that are priorities for conservation actionglobally (Green et al. 1996). Thus, Indonesian ecoregionsarguably do not place greater emphasis on habitat repre-sentation and may reflect expediency—which existingclassifications can be mapped at the desired scale—at theexpense of other ecological and biogeographical patterns.

Indonesia already has a protected-area networkplanned and justified by the Dasmann-Udvardy frame-work operationalized by MacKinnon. In practice, con-servation planners in Indonesia capture habitat variabil-ity in reserve networks by listing and locating vegetation

Figure 2. Endemic bird areas of Indonesia, redrawn with permission from International Council for Bird Preser-vation (1992): 1, Sumatra; 2, Enggano; 3, Bornean mountains; 4, Java and Bali forests; 5, Javan coastal zone; 6, Sulawesi lowlands; 7, Sulawesi mountains; 8, Sangihe-Talaud; 9, north Nusa Tenggara; 10, Sumba; 11, Timor and Wetar; 12, Banda Sea Islands; 13, Seram; 14, Buru; 15, Banggai and Sula; 16, north Molucca (including Halma-hera); 17, west Papuan lowlands; 18, west Papuan mountains; 19, Geelvink islands; 20, north Papuan lowlands; 21, north Papuan mountains; 22, central Papuan ranges; 23, south Papuan lowlands; 24, Trans-Fly.

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formations and associations occurring within each bio-unit at the time a reserve is designed. This is expedientbecause it avoids the need for a hierarchical system ofvegetation, which would be required to capture themassive vegetation heterogeneity in Indonesia and whichwould be a complex task to attempt.

We are unclear how introduction of WWF ecoregionswill strengthen the design of a protected-area system inIndonesia. As currently presented, ecoregions risk weak-ening the system by implying that past planning wasbased on inferior science or information and by break-ing the link between spatial frameworks and clear andpractical reserve-design principles. In addition, there is amore subtle issue. The goal of conserving large blocks ofnatural habitats has been pursued in Indonesia by advo-cating major ecosystem reserves. These are justified onthe basis of watershed protection, the conservation im-portance of intact ecological gradients, and area require-ments of viable populations of megafauna. In short,these reserves and surrounding forest landscapes havebeen presented as functioning ecological entities (e.g.,the Leuser Ecosystem). Ecoregions are also so pre-sented, but delineation on the basis of habitat bound-aries results in the division of at least 11 major ecosys-

tem reserves. This creates two alternative ecologicalmanagement units, the ecoregion and the major ecosys-tem reserve. Embarking on such major reorganization ata time when ecosystem reserves in Indonesia are undersiege from exploitation interests is inadvisable.

Replacement of existing conservation planning frame-works with ecoregions could have important implica-tions for the targeting of future conservation resources.For example, representation of the Sulawesi lowlands ashomogenous ( Wikramanayake et al. 2001) may suggestthat targeting resources to any peninsula is equally good,whereas in reality the northern (Minahasa) peninsula isthe highest priority for conservation of large mammalsand endemic species ( Whitten et al. 1987; Sujatnika etal. 1995). Creating mangrove ecoregions in the Sunda-shelf and New Guinea bioregions but not in Wallaceamay lead some to conclude that Wallacea lacks man-groves worthy of conservation investments, which is notthe case. The ecoregion complexes in east and southKalimantan could create the misleading impression thatinvesting in the provinces where they occur will con-serve more biodiversity than similar investment in ap-parently simpler regions such as the Lesser Sundas or Su-lawesi.

Figure 3. World Wildlife Fund ecoregions of the Indo-Pacific, redrawn with permission from Wikramanayake et al. (2001): 82, Sumatran lowland rainforests; 83, Sumatran montane rainforests; 84, Mentawi Islands rainforest; 85, Sumatran peat-swamp forests; 86, Bornean peat-swamp forests; 88, Sumatran freshwater swamp forests; 89, Southern Borneo freshwater swamp forests; 90, Sundaland heath forests; 93, western Java rainforests; 94, eastern Java-Bali montane rainforests; 95, Borneo montane rainforest; 96, Borneo lowland rainforests; 105, Sumatran tropical pine forests; 107, Sunda shelf mangrove; 109, Sulawesi lowland rainforest; 110, Sulawesi montane rain-forest; 111, Lesser Sundas deciduous forests; 112, Timor and Wetar deciduous forests; 113, Sumba deciduous for-ests; 114, Halmahera rainforest; 115, Buru rainforest; 116, Seram rainforest; 117, Banda Sea Islands moist decidu-ous forests; 118, Vogelkop montane forests; 119, Vogelkop-Aru lowland forests; 120, Biak-Numfoor rainforests; 121, Japen rainforests; 122, northern New Guinea montane rainforests; 123, northern New Guinea lowland rainforest and freshwater swamp forest; 125, central Range montane rainforests; 127, southern New Guinea freshwater swamp forests; 128, southern New Guinea lowland rainforest; 129, New Guinea mangroves.

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Given these problems, and the fact that in their NorthAmerican scheme WWF has adopted geomorphologyand topography as delineators and in the names of theecoregions, we are unclear why they have not done thesame in Indonesia and the Asia-Pacific realm. Adoptionof landform as a primary mesoscale ecoregion delinea-tor, as recommended by Bailey, would have made a dis-tinct and useful contribution and avoided several of theproblems described above. A landform analysis of Indo-nesia was conducted by RePPProT (1990) and is avail-able.

Discussion

The first aim of our paper was to place WWF ecoregionsin context by assessing how the alternative systems dif-fer. The Dasmann-Udvardy biogeographic frameworkand Bailey-Omernik ecoregion frameworks are distinctin terms of methodology (captured in the terms used)and purpose. The former combines ecoclimatic and tax-onomic delineators with the purpose of achieving bio-logical representation in a global reserve network, and itemphases taxonomic delineators. The latter emphasizesecoclimatic indicators to achieve sustainable land use.The WWF ecoregions approach aims to do both andadopts methodological aspects of each spatial frame-work. At level 3, the mesoscale, the Dasmann-Udvardyapproach as extended by MacKinnon employs taxo-nomic delineators, the Bailey and Omernik ecoregionscheme employs landform and topographic delineators,and the WWF ecoregions approach uses a gestalt synthe-sis of available regional schemes, which unfortunatelyhave uneven geographical coverage.

It is important to distinguish between systematic plan-ning frameworks and strategic planning approaches.The existing schemes are primarily spatial planningframeworks that represent three basic conservation ap-proaches: (1) representation of biodiversity attributes innetworks or reserves (Dasmann-Udvardy system), (2)protection of special elements such as centers of speciesendemism or richness (EBAs, centers of plant diversity),and (3) land-use planning within ecologically defined ar-eas (Bailey-Omernik ecoregions). The zonal schemes as-sume that the actions for which they are designed (des-ignation of reserves, optimal management of land) willbe pursued within each spatial unit. The azonal frame-works assume that each hotspot merits conservation ac-tion. In this sense, the distinctive contribution of WWFecoregions is the application of strategic-planning crite-ria to a zonal framework. The value of the WWF contri-bution therefore depends on the scientific merits ofboth azonal and zonal aspects. Our critique has consid-ered only the zonal framework.

Our second concern is scientific explicitness, trans-parency, and repeatability of methods. These are impor-

tant features of systematic conservation planning toolsthat guard against planning decisions having more to dowith political, organizational, and technological expedi-ency than persistence of biodiversity ( Pressey 1999).We have shown that preexisting schemes generallymeet these criteria. By adopting an existing ecoregionscheme (Omernik 1987 ), the WWF ecoregions of NorthAmerica (Ricketts et al. 1999) appear to meet these cri-teria (but see Wright et al. 1998).

We argue that the WWF ecoregion methodology is un-clear in important respects and that the assumption thatmajor habitat boundaries equate to mesoscale ecosystemboundaries is flawed in several instances. We are con-cerned that ecoregion delineation on the basis of gestaltsynthesis and “expert” review may not meet our threemethodological criteria well enough for ecoregions toform the foundation of global and national conservationplanning decisions. Specifically, the claim that “several”other regional-scale assessments (i.e., Dinerstein et al.1995; Ricketts et al. 1999) use the same ecoregion delin-eation scheme cannot be justified. Whereas in NorthAmerica the WWF adopted the Bailey-Omernik scheme(Ricketts et al. 1999), in Indonesia it adopted the Das-mann-Udvardy scheme as modified by MacKinnon ( Wik-ramanayake et al. 2001). Consequently, prioritizationamong ecoregions globally (Olson & Dinerstein 1998) issuspect because similar regions are not compared glo-bally.

The development of systematic approaches is produc-ing powerful decision-support tools and has the poten-tial to improve the public accountability of conservationagencies. One measure of explicitness and transparency,however, is consistency in terminology, which is poorin this field. Within our comparatively brief review, wehave encountered an enormous variety of terms ( Table1). The potential for confusion and inefficiency in devel-opment of conservation planning and implementationappears considerable. We therefore call for greater stan-dardization of terminology in the field and for explicitreferencing of terminology and delineators to existingschemes and classifications.

Our third goal was to ask if WWF ecoregions improveupon existing schemes, for which we took Indonesia asa case study. We focused on the utility of the schemesfor guiding tangible, on-the-ground conservation out-comes. Our assessment ( Table 2 ) is that the existingschemes of Dasmann-Udvardy (adapted by Mackinnonand colleagues), complimented by EBA and reserve-designprinciples, for all their individual weaknesses, have pro-vided a successful mesoscale conservation planningframework. They have informed and motivated a majorexpansion of the protected-area network, established abiogeographic perception of space in government de-partments, and educated Indonesian society.

The WWF ecoregions “seek to advance biodiversityconservation planning beyond previous approaches,

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such as hotspots” (Dinerstein et al. 1995:2) to “achieverepresentation of habitat types on a global scale” (Olson& Dinerstein 1998:502) and to move conservation be-yond “reactive efforts and into a realm of visionary, pro-active approaches at regional and landscape scales”—or“big conservation” as they now define it ( Wikramanaykeet al. 2001). Our review demonstrates that systems toachieve representation predate hotspot approaches andthat conservation planning to achieve habitat represen-tation has been the norm in Indonesia for 25 years. Con-servation planners have long thought beyond politicalboundaries and at landscape scales. The Bentuang-Kari-mun transnational reserve (designated in 1995) and theKerinci-Seblat National Park (declared in 1982), whichspans four provincial (state) boundaries, are testimonyto this point. The 1982 Indonesian national conservationplan was proactive and visionary. Furthermore, deliver-ing on this vision, which is already advanced, is the besthope of meeting biodiversity conservation goals. This isespecially so given that spatial planning in the widerlandscape may be obsolete for the next decade or morebecause Mafia-like networks involved in illegal loggingand land grabs constitute the de facto institutional ar-rangement for natural resource management in manynewly decentralized districts ( Jepson et al. 2001). Fur-thermore, given that conservation resources are scarce,ever more detailed ecoreogional work may take fundsfrom field conservation.

Nevertheless, we believe there is scope to modernizethe Dasmann-Udvardy system. We suggest that Bailey’sthird-tier delineator of landform mosaic or landscapelevel can readily be incorporated into the Dasmann-Udvardy system as a third- or fourth-level delineator (Ta-ble 1), and we see five principal benefits of doing this.First, professional conservation planners in Indonesiause several maps (and systems) in conjunction with oneanother. A landform-based map would provide a wel-come new resource to replace existing physiographicmaps (e.g., RePPProT 1990) that are sometimes foundlacking for conservation planning purposes. Second,landscape units are suitable for biogeographical analysisat increasingly discrete scales. Third, landscape providesa framework for considering ecological linkages and pro-cesses at the scale at which they operate (Brunckhorst& Rollings 1999). Fourth, humans change landscapes,and landscapes embody culture, so threats to biodiver-sity operate at this level. This dynamic is a basic idea inemerging integrated implementation approaches suchas bioregional management ( Miller 1995). Finally, allagencies involved in natural resource or land manage-ment ( and society as a whole) perceive large areas ascollections of landscapes, thereby providing the basisfor the interagency cooperation necessary for successfulconservation planning and implementation.

Ironically, the fact that state-planned conversion ofnatural landscapes constitutes a major threat to biodiver-

sity in Indonesia means that the overlay maps of physi-ography, geology, and land use necessary to the applica-tion of Bailey’s third-level delineators, are well-developed and generally available. Unfortunately, WWFecoregions do not make this step but instead appear tocombine existing frameworks based on a gestalt meth-odology that renders their output less explicit.

Given that the existing schemes, despite their flaws,have achieved their purpose in Indonesia and are trans-parent in their methods and operational criteria, wequestion the wisdom of introducing the WWF ecore-gions system. The tangible additional benefits that itpromises—extension to the aquatic and marine realmsand introduction of an ecologically based planningframework to a new generation of decision-makers—ap-pear limited in the Indonesian context and may be asreadily attainable within the existing conservation plan-ning framework. Moreover, there is a risk that the intro-duction of a new scheme may undermine and delay im-plementation of existing schemes that are well along inachieving comparable goals.

In other parts of the world where long-establishedconservation planning frameworks are lacking, there maybe a stronger case for introducing the WWF ecoregionsframework, providing that the methodological concernsraised here (and see Wright et al. 1998) are addressed.

Acknowledgments

We thank C. Bibby, W. Duckworth, A. Hamilton, S.Schmitt, and J. Tordoff for discussion of earlier versionsof this paper, and R. E. Pressey, T. C. Whitmore, A. J.Whitten, and three anonymous reviewers for construc-tive comments during the review process. The views ex-pressed are our own. We thank BirdLife International,the Asian Conservation Bureau, and World WildlifeFund–United States for permission to reproduce the maps.

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