Human Impacts and the Global Distribution of Extinction Risk Author(s): Richard G. Davies, C. David L.

Orme, Valerie Olson, Gavin H. Thomas, Simon G. Ross, Tzung-Su Ding, Pamela C. Rasmussen, Ali J. Stattersfield, Peter M. Bennett, Tim M. Blackburn, Ian P. F. Owens, Kevin J. Gaston Reviewed work(s): Source: Proceedings: Biological Sciences, Vol. 273, No. 1598 (Sep. 7, 2006), pp. 2127-2133 Published by: The Royal Society Stable URL: http://www.jstor.org/stable/25223578 . Accessed: 05/11/2011 08:39
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PROCEEDINGS

Proc. R. Soc. B (2006) 273, 2127-2133 -OF-T7Y) THE ROYAL I Kn doi:10.1098/rspb.2006.3551

SOCIETY JAJJ

Published online8 June2006

Human
Richard Gavin Pamela Tim M.

and the global impacts risk of extinction

G. Davies1, H. Thomas4, C. David Simon Ali L. Orme2, G. Ross1,

distribution
Valerie Olson3, Ding5,

C. Rasmussen6, Blackburn4,

M. Bennett3, J. Stattersfield7, and Kevin Ian P. F. Owens2'8 J. Gaston1'*

Tzung-Su Peter

1 Biodiversity & Macroecology Group, Department ofAnimal & Plant Sciences, University of Sheffield, Sheffield S10 2TN, UK 2Division of Biology, Imperial College London, Silwood Park, Ascot, Berkshire SL5 7PY, UK 3Institute of Zoology, Zoological Society of London, Regent's Park, London NW1 4RY, UK 4School of Biosciences, University of Birmingham, Edgbaston, Birmingham B15 2TT, UK
5School of Forestry & Resource Conservation, National Taiwan University, 1, Sec 4,

Roosevelt Road, Taipei 106, Taiwan, ROC 6Michigan State University Museum, West Circle Drive, East Lansing, MI 48824-1045, USA 7BirdLife International, Wellbrook Court, Girton, Girton Road, Cambridge CB3 ONA, UK 8NERC Centre for Population Biology, Imperial College London, Silwood Park, Ascot, Berkshire SL5 7PY, UK Understanding
biology. high It remains human

the global geographical

controversial, or due however, to impacts

distribution
to what

of extinction
extent areas

risk is a key challenge

threat hotspots to Limits the

in conservation
simply because of and taxonomic

become

predisposing

ecological

conditions.

geographical
assessment distributions richness, Ecological biogeographic risk globally the mechanisms driving Keywords: the the of

extent,
the of birds

resolution
roles of on continents of

and quality
these and the global factors.

of previously
Here, we use islands of at

available
a new to show risk scale.

data have precluded

global that, database after on the controlling of

a full global
geographical for species impact. individual extinction on in

relative

continental pattern

best

gradients

predictors are of within

extinction

are measures converse and its

human for on

secondary which

importance variation These of

a global impact underline risk, and

The

is true influence

realms, is therefore driving current

in human results extinction

is reduced the the

underestimated. spatial patterns crisis. global

importance key role of

of a global anthropogenic

perspective factors

extinction risk;

extinction

biodiversity;

human

population;

species

richness;

threatened

species

1. INTRODUCTION
Understanding
risk and its

the geographical
causes are key

distribution
challenges in

of extinction
conservation

biology,
for ants the of

and are central

focus of conservation risk extinction

to determining
responses. across space impacts but

spatial priorities
Major include also determin not variation only in

thought to limit overall and individual species population numbers (Wright 1983), absolute species numbers which may influence food web structure and thus the likelihood of extinction cascades and surface (Gaston 2002),
topography which influences the occurrence of narrowly

distributed
Despite restrictions resolution

species
a to and the

(Richerson
of taxonomic of

St Lum
and

1980).
regional studies, extent, data, have

anthropogenic

environmental

number

valuable

predisposing 1996). The

agricultural introduced

(Forester St Machlis ecological conditions former include human population density,

and species urban and land-use, disease, species and exploitation, anthropogenic

geographical available

quality

previously

thus far largely precluded

major taxon of the relative predisposing ecological

a full global
roles of

assessment
impact

for a
and

human

climate
Predisposing exclusive energy

change
to, which the

(Soul?

1991; Forester
conditions of influence ambient

& Machlis
but

1996).
are not

factors

in determining

threatened

ecological

include,

availability to

environmental rates and

is thought

speciation

thus the occurrence

availability of productive

of neoendemics
environmental

(Rohde
energy

1992),
which

the
is

et al 1995; Balmford species richness (Kerr & Currie 2001; McKinney 2001; Norris St Pain 2002; Luck et al et al 2005). Given that the sensitivity 2004; Scharlemann of individual species to human population density has
been regions whether shown to vary within 2000), and between an factors biogeographical open question by (Woodroffe the relative it remains of

importance

indicated

*Author The

for correspondence

(k.j.gaston@sheffield.ac.uk). at http://dx.doi. material is available or via http://www.journals.royalsoc.ac.

individual regional
or globally. The within individual impact 2005 2127 ? 2006 and The

studies will generalize

frequently continents,

to other regions

supplementary org/10.1098/rspb.2006.3551 uk. 28 16 March 2006

electronic

ecological Royal

incomplete representation, of global variation in human as well as the distinct gradients, Society

Received Accepted

November

2128

R. G. D avies and others histories

to conservation

Global distribution of extinction risk occurring

this of

evolutionary
regions, relevance

of species
to

in different
The answer the

number

of threatened

species

in each

grid

cell.

Biogeographic

further

contribute

uncertainty.

realms were
ecoregions ical, final map

delimited
(Nearctic,

using

the World Wildlife

Olson grid Neotropical, et al cells 2001).

Fund
The

policy-making

Palaearctic,

Afrotrop

lies both regional

causes human extinction new breeding continental

in its indication of the wider applicability of of ultimate findings and in the confirmation
globally. Here, we present of global on grid spatial scales. an analysis patterns We use of of of a the and determinants ecological risk at continental and on of the extant on an geographical bird species

Indo-Malayan, dataset used or Antarctica, for as any of these land

and Australasian; for analyses since realms. area shelf, markedly threatening with et al. omitted

operating and

Oceania available defined edge known

environmental true than omitted kinds and affecting

falling within not data were islands, from these the are of

Remaining further also the were in

oceanic 200 km

located

database ranges

distribution continents at

continental to differ and

since

intensities constituent (Manne

islands

equal-area

a resolution

evolutionary avifauna, et al 2004).

processes continental 2004;

comparable
We impact skewing ance of tested and

to Io latitude X longitude
equal ecological numbers condition of

(Orme et al. 2005).

of avoid human a priori

in comparison 1999; Finally, Blackburn

locations

predictors so as to

Duncan

St Blackburn

so as to avoid to the

bias

in terms

of the contribution grid the final cells with

analyses predictors

in favour from

of finding the greater import one over the other. category

of coastal less than

land-area 50%

regression omitted

models, from

land-cover

were

dataset.

Building
importance species human power

on previous
in richness, population shaping for

demonstrations
spatial indices of patterns human

of their potential
of impact activity GDP), threatened we used (b) Environmental Data for the eight (see same data selected were environmental each grid as GDP, re-projected the species and human and impact

density, domestic

economic product,

parity

gross

(purchase and extent of

predictors to the

above) equal-area

re-sampled data. and log10

richness agricultural were all

agricultural and urban land-area (Kerr St Currie 1995; et ah 2001; McKinney Balmford 2001; Norris & Pain et ah 2005). For 2002; Luck et ah 2004; Scharlemann
ecological a measure gradients, of we available used mean ambient annual energy, temperature while as for

Human urban

population land-area, for and the

density, and

NDVI, range

elevation

transformed Sources environmental density (CIESIN for 127

analysis. resolutions are as follows: for from 1995 human units, have at of the eight selected population resolution census on data

raw

variables km-2) derived administrative estimates

(a) human 2.5 arc-min

productive Vegetation
variability cover types

Difference energy we used the Normalized Index (NDVI). In addition to topographical

(elevation (habitat range), diversity) in our we used number of measure land of as an alternative

(people 2003) 105

population and been based adjusted for

national to match country;

population the UN

that estimated

habitat heterogeneity.
spurious variables

To minimize
analyses, we

the risk of including

built a multivariate

national

population

each

minimum
methods response

adequate model
that accounted To for variable.

(MAM)
spatial the

based
results

on regression
in the global of our

(b) purchase power parity GDP data (US J) for 1990 at 0.25? resolution (CIESIN 2005); (c) agricultural land-area (km2);
(d) urban land-cover and types using built-up (habitat remotely land-area diversity) sensed (km2); occurring data for and (e) number in a grid cell, the 12-month of all

autocorrelation

test whether

model
a

could have been predicted

geographical to construct extent,

by analyses conducted
we used separately the for same

at
six

computed

smaller

period
resolution (USGS) 2003a); elevation

between April
classified 25-category (f ) elevation within each

1992 and March

to the US land-cover range

1993 at 30 arc-s
Survey (USGS minimum

methodology

models

Geological classification minus arc-s

major biogeographic

realms

(Olson et ah 2001).

(m), maximum from 30

2. MATERIAL AND METHODS (a) Species data

The analyses presented maps using et al. 2005, for a variety in press) here 9626 are extant, based on a database bird species details avian of distribution constructed see Orme recognized sources of published and following

data grid cell, mean data annual temperature (?C) for 20036); (g) (USGS from at 10 min resolution 1961-1990 the period interpolated et al 2002); annual station means remotely (New (h) mean at 0.25? resolution 1982-1996 for the period sensed NDVI resolution II 2005). the sum of Agricultural all agricultural land-area land-use (above) classes

(for a standard

Initiative (ISLSCP as was computed

taxonomy
range vector a Behrmann This grid

(Sibley StMonroe
maps were converted at projection cell size

1990). The polygon breeding

to an equal-area resolution Io longitude of grid using a cell to 96486.2m. Io latitude

from the USGS

and pasture; 3, dryland/irrigated mosaic; dize the 6,

data (USGS 2003a)

irrigated cropland cropland and pasture; mosaic). terrestrial each areas was map and

(2, dryland cropland

pasture; 4, mixed 5, cropland/grassland to standar In order land-area overlaid across with 1993) Raw-data of and raw

is equivalent

and

cropland/woodland definition datasets, terrestrial to the of

at 30? latitude N/S

Behrmann grid cells present that the therefore at latitudes in a grid breeding species projection

(l/360th of the width of the globe under a

using 360 than if any range were but Least of fell those not Concern) the WGS84 by 152 87.13?. the cells, datum). omitting were Species sources cell The global as the partial scored indicated boundaries. Endan categories (Data 2000). used in avian

environmental resolution

a high prior cells, land-area the latter each to or

contains higher cell

(ESRI gird. this

available the

within classified those

re-sampling outside of cells, falling portions were from excluded re-sampling were the by weighted cell. raw-data remaining

Io Behrmann

definition

calculations, associated

Threatened gered (Near Deficient Where BirdLife

as Critical, risk

land-area

with

and Vulnerable, Threatened, and Not necessary, International

in lower or other

categories

Evaluated; we

BirdLife the back to

International taxonomy the standard

(c) Statistical To deal

analyses with not spatial autocorrelation distributed, linear and analyses mixed a

converted (2000)

simultaneously variable on that was a Poisson

response were

taxonomy

(Sibley & Monroe

(2006)

1990),

and calculated

the

based

normally errors generalized

Proc. R. Soc. B

Global distribution of extinction risk

R. G. Davies

and others

2129

(a)

'%*>-#

31 ?psp??PF'f y

i?

(
0.185

???

fpppi

(c)

943 llppl

W)

3.21

-5.00 Figure species threat which 1. Geographical distribution of avian threatened richness and of human (a) Threatened species population density, over areas with richness, of species that are threatened those (b) The (with a blue mask proportion high proportional use of a scaling to 0.1) to enable than or equal and low species richness for illustrative purposes (less than 30), (greater are otherwise reveals details of the map which richness, obscured), (c) Non-threatened (d) Logi0 (human general species density).

population

modelling
which with an

(GLMM) method
exponential and Proc spatial latitudinal Glimmix

(SAS, Littell
covariance cell v. 1.0 centroid add-in other of

et al
values in SAS spatial

1996)
is fitted as spatial v. 9.1.3.

in

non-spatial took realms geographic degrees, account

Poisson

error

model

residuals. among by major estimating

Spatial

GLMMs

structure

of the differences autocorrelation or range

biogeographical the maximum (p), measured in equivalent to occur. in

longitudinal using of was

in spatial

variables The choice

distance over which errors

parameter autocorrelation

the exponential, based on

over inspection

covariance of

spatial model

structures,

semi-variograms

independent

residuals

was

observed

Proc. R. Soc. B

(2006)

2130
This residuals the

R. G. Davies
involved

and others
p Poisson from

Global distribution of extinction risk

the semi-variogram that included for as with the each spatial spatial same of However, the importance of elevation range as a positive

estimating

of non-spatial combination six estimates

errors

models

predictor
result from

of
the

threatened
influence of

species

richness
on

globally
the

may their

relevant All

of predictors, of in for p were the global then

separately entered

topography to

occurrence

realm.

of restricted-range
inherent associated

species

(Jetz et ah 2004)
population

and

covariance autocorrelation realm.

parameters assumed used

model, within

vulnerability

decline

observations

(Stattersfield
in some cases contractions leaving

et ah 1998; Manne
this from occurrence human-impacted populations

et al. 1999). Alternatively,

may result lower in more from elevation mountainous range areas

GLMMs

the pseudo-likelihood 1993) of not the that scale obtains parameter a true

(PL)

procedure

(Wolfinger likelihood-like et al 1996).

St O'Connell estimate PL use does of model

a maximum (cp) (Littell

remnant

compute selection

log-likelihood, based stepwise determine terms nonlinear cannot so we be used non areas variables Tolerance following for on

precluding Akaike's model-building MAMs. predictors relationships. derived percentage spatial modelled, using levels models we The

procedures and forward to linear for

influence regions. Similarly, the positive be linked to contraction of the geographic

threatened species to remaining areas

of NDVI may distribution of

of high plant

Information procedures fit was of

Criterion, were

quadratic in of models

employed as as well to allow

also included the signifi productivity. The global MAM cant negative influence of GDP indicating that, having
accounted development for other are factors, coincident areas with of high economic numbers of lower

tested Estimates

order variance that explained For

explained use PL,

from of

spatial total as an

threatened
local contractions (including influence investment In contrast

species. This
from urban of and resulting areas areas),

is more
in of highest

likely to result
species economic from on any

from
range

deviance

from all

equivalent

extinction

threatened

indication. collinearity (Quinn high

geographical predictor 2002). than 0.1,

activity positive

explored levels

among St Keough

rather development

than

tolerance were

economic effectiveness. to the global

conservation

sufficiently

(i.e.

greater

Quinn St Keough
and Neotropical between land-area observed urban

2002)
realms human (for

in all cases except for the Nearctic

where population values see some redundancy density, table GDP was and

model,

our

analyses

of

threat

within human
importance (table

that realms generally suggested biogeographic lower in variables ranked relatively impact
compared The only with exception 2). ecological to this was predictors Australasia,

tolerance Hence,

1 in electronic minor in

supplementary consequence the Neotropical of the other

material). that

this had of human

the relatively population from

significance could

density the effect

which
primary even of latter species was

showed

human

population
species predictor

density

to be

the
the

MAM two

not

be

separated

socio-economic

predictors.

of predictor non-threatened the primary for

threatened

species richness. of numbers realms, and

richness, However, of was and

ahead

threatened the Palaearctic, only

3. RESULTS AND DISCUSSION

The
richness

predictor

all remaining to enter MAMs

for

the Nearctic

global

distribution
marked

of

threatened
spatial

avian the

species Indo
coastal

the two higher

numbers realms, rank being of subsequent orders, the with second

latitude realms with

species. For entered predictors NDVI,

the lowest absolute

the major MAMs and tropical in different temperature for realms.

exhibits

large-scale

heterogeneity

threatened

(figure Malayan
areas forests. distribution extent, (Kerr constant absolute threatened continental portional and New of

across much of la), being highest realm and parts of the Neotropics,
the Andes, work threatened on 1995; that McKee Amazonia has shown species of et al and the that Atlantic the Previous of

including

elevation

range,

strongest

predictors, and

respectively,

richness overall 2003).

geographical is, to some richness the

Afrotropical,

Neotropical,

Indo-Malayan

Likely
elevation global NDVI the

reasons
range or MAM,

for
are

the positive
the the range same as importance

slopes
those of

for NDVI
proposed temperature is indicative on for

and
the over of the

dependent & Currie

species However,

whereas elevation

proportion

of species
and does of not

threatened
simply either threatened 1c). areas

(figure
mirror

lb) is far from

the for patterns la) or non the respect richest to pro

in Indo-Malaya lowland of plant

impact These

of widespread distribution results the have

deforestation

numbers

(figure Instead, with

large-scale

productivity. consequences underlie for under contemporary between the realm of for realms pre one or

(figure species or larger-island threat Zealand. are the southern

important that instance, analyses that

standing extinction the results

mechanisms For global indicate

Palaearctic,

Madagascar,

processes. of analyses of cannot threatened be our

the differences and the richness either those combination observed to other of

Our global model

effect of spatial

revealed that, after controlling

variation in non-threatened

for the
species

specific dictors realm

species

richness, human population density (figure Id) was the primary global driver of geographical patterns of numbers of followed bird species, of threatened by extent
agricultural (table role, with 1). activity Environmental elevation range as a secondary factors and NDVI played entering human a more influence minor

assumed

to apply

globally. One
realms often

reason for this appears to be that individual

contain relatively limited geographical

variation
numbers the of

in human
threatened

impacts
species realm shows

(figure
(figure the

2a)
2b). For maximum species

and/or
example,

the

as subsidiary we and to the

Indo-Malayan levels both

global richness

factors
human might primary remaining since we areas

in the global MAM.

factors expect and gradients to

Accounting
such be as

for the effects of

avian richness, range related threatened species

average

of

threatened

non-threatened

(figure 2b) and of human

an order of of magnitude this, human for spite enter

population
than

density,
in any other does

these being
realm. not In even

elevation

higher population Indo-Malaya

productivity variation could expect

inversely of

density since

in numbers fewer with

species, in pristine impact.

the MAM

variation

in human

threatened areas of lower

that

coincide

human

population density within the realm is relatively limited at the weak this spatial resolution (figure 2a). Hence,

Proc. R. Soc. B

(2006)

Global distribution of extinction risk R. G. Davies

Table model 1. Global was minimum using adequate forward in the number of threatened variation model for geographic is the overall % expl. deviance percentage stepwise procedures. as spatial GLMMs. test All other combinations of predictors are Difference GDP as follows: birds.

and others

2131

obtained

the same models using non-spatial to spatial Abbreviations of slope, and F-value) refer GLMM results. interval non-threatened domestic Normalized NDVI, gross GDP, product; species; to quadratic elevation range, terms). density, Population agricultural-area, ***p<0.001; **0.001<p<0.01; *0.01 <p<0.05.) conf.

(The minimum adequate of total deviance by explained statistics 95% confidence (slope, non-threat spp., number of refer

Index Vegetation were all and NDVI

(superscripts log10-transformed.

predictor non-threat non-threat population

estimate

95%

interval

Fu 560.63** 201.55** 43.50** 20.77** 19.75** 13.74** 8.92** 3.38*

expl.

deviance

spp. spp.2 density2

0.0051 -0.0000003 0.0052 0.0049 0.0090 -0.0158 1.6600 0.1090 -0.5110

agricultural-area2 elevation range2

GDP2 NDVI2
land-area intercept

?0.0004 ?0.0000 ?0.0016 ?0.0021 ?0.0040 ?0.0084 ?1.0894 ?0.1162 ?0.4690

variation in the number of threatened

41.4

Table

2. Minimum

(Abbreviations

models adequate as and methodology

for geographic in table 1.)

birds

within

biogeographical

realms.

realm =

predictor

estimate

95%

conf.

interval

F-value

expl.

deviance

Australasian

(d.f.

932)

population elevation non-threat temperature temperature2

density2 range2 spp.

0.0128

0.0322 0.000003
-0.2783 0.0025 1.2802 -0.0859 8.4088

NDVI
land-area intercept Afrotropical (d.f. = 2313) non-threat spp.

0.0044 1.5221 0.4889 0.0655

NDVI
land-area

GDP
non-threat intercept Indo-Malayan (d.f. = 881) non-threat temperature temperature2 spp. spp.2

-0.000002 -1.6776 0.0014 0.1484 -0.0013 0.8523 0.0262 -0.0442 -2.1804

NDVI
agricultural-area land-area intercept Nearctic (d.f. = 2061) non-threat non-threat land-area intercept Neotropical (d.f. = 2039) non-threat non-threat elevation population spp. spp.2 range2 density2 spp. spp.2

0.0270 -0.0001 0.1586 -3.7346 0.0037 -0.000002 0.0228 0.0052 0.0215 0.5051 -0.0746 -0.0008 0.0654 -2.6566

?0.0053 ?0.0140 ?0.0000 ?0.1634 ?0.0016 ?1.0929 ?0.2509 ?4.3179 ?0.0014 ?0.6795 ?0.2777 ?0.0419 ?0.0000 ?1.3679 ?0.0003 ?0.0685 ?0.0007 ?0.4784 ?0.0196 ?0.1226 ?1.7534 ?0.0121 ?0.0000
?0.6768

21.93*** 20.25*** 18.67*** 11.14*** 9.36** 5.27* 0.45

66.1

38.92*** 19.28*** 11.90*** 9.38** 4.72*

26.6

105.94*** 18.04*** 14.42*** 12.19*** 6.89** 0.50

51.1

19.18*** 10.03** 0.21

21.4

agricultural-area2 land-area agricultural-area temperature2 temperature intercept Palaearctic (d.f. = 5603) non-threat land-area intercept spp.

?3.0745 ?0.0008 ?0.0000 ?0.0083 ?0.0026 ?0.0113 ?0.2740 ?0.0484 ?0.0005 ?0.0523 ?1.9331 ?0.0004 ?0.1905 ?0.7752
with in intra-realm comparison models variation to the

78.14*** 50.28*** 28.86*** 14.80*** 13.93*** 13.06*** 9.13** 8.35** 6.01*

59.4

0.0041 0.0271 0.2982

356.29*** 0.08

28.3

Indo predictive strength of population density within Malaya belies the fact that this variable is largely driving
the peaks in global avian same pattern is repeated Proc. R. Soc. B (2006) threatened across several species of richness. the other The realms,

typically global

being

relatively resulting

minor in than the the

patterns, less

regional

having

considerably

power

full global analysis.

2132

R. G. Davies

and others

Global distribution of extinction risk

risk for the one on best-known of vertebrate taxon, analyses currently our findings of human available to

104

represent impacts inform We

the highest-resolution biodiversity policy.

global

conservation T. Allnutt, Fry,

thank

H. J. Cromie, C. Perrins, R. C. Robertson, M.

B. Beehler, T B. Coates, Brooks, P. Higgins, D. McNicol, D. Mehlman, H. R. S. Ridgely, Porter, Pratt, N. Redman, A. M. M. Silcocks, Unwin, Strange,

M. Whitby, P. Williams, B. Young, D. Wynn, Weston, A. & C. Black, Academic BirdGuides J. Zook, Press, Ltd, Birds Conservation Inter Australia, Helm, Christopher the Ornitho Oxford national, NatureServe, Press, University logical Society for access Press of New to data; Zealand L. Birch, and R. Princeton Prys-Jones, University B. Sheldon,

the Alexander Library

(Oxford University),

and the Natural

to libraries; M. Museum for access History (Tring) Burgess, for technical F. Eigenbrod and N. three assistance; Pickup for comments; and O. Schabenberger reviewers anonymous for analytical advice. This work was funded by The Natural

Environment Research Council (grant nos NER/O/S/2001/ and 01258, NER/O/S/2001/01257, NER/O/S/2001/01230, NER/O/S/2001/01259).

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