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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
SOCIETY JAJJ
Human
Richard Gavin Pamela Tim M.
distribution
Valerie Olson3, Ding5,
C. Rasmussen6, Blackburn4,
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
distribution
to what
of extinction
extent areas
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.
a full global
geographical for species impact. individual extinction on in
relative
continental pattern
best
gradients
extinction
human for on
secondary which
The
is true influence
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
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
geographical available
quality
previously
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
is thought
speciation
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
individual regional
or globally. The within individual impact 2005 2127 ? 2006 and The
to other regions
electronic
ecological Royal
incomplete representation, of global variation in human as well as the distinct gradients, Society
Received Accepted
November
2128
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
Fund
The
policy-making
Palaearctic,
Afrotrop
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
and Australasian; for analyses since realms. area shelf, markedly threatening with et al. omitted
operating and
falling within not data were islands, from these the are of
oceanic 200 km
located
database ranges
distribution continents at
since
islands
equal-area
a resolution
comparable
We impact skewing ance of tested and
to Io latitude X longitude
equal ecological numbers condition of
locations
predictors so as to
Duncan
St Blackburn
so as to avoid to the
bias
in terms
analyses predictors
in favour from
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
predictors to the
above) equal-area
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
density, and
NDVI, range
elevation
analysis. resolutions are as follows: for from 1995 human units, have at of the eight selected population resolution census on data
raw
productive Vegetation
variability cover types
population the UN
that estimated
habitat heterogeneity.
spurious variables
To minimize
analyses, we
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
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
1993 at 30 arc-s
Survey (USGS minimum
methodology
models
major biogeographic
realms
(Olson et ah 2001).
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
taxonomy
range vector a Behrmann This grid
(Sibley StMonroe
maps were converted at projection cell size
is equivalent
and
environmental resolution
available the
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
as Critical, risk
land-area
with
in lower or other
categories
Evaluated; we
analyses with not spatial autocorrelation distributed, linear and analyses mixed a
converted (2000)
response were
taxonomy
1990),
and calculated
the
based
Proc. R. Soc. B
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
Poisson
error
model
Spatial
GLMMs
structure
in spatial
parameter autocorrelation
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
estimating
errors
models
predictor
result from
of
the
threatened
influence of
species
richness
on
globally
the
may their
relevant All
separately entered
topography to
occurrence
realm.
of restricted-range
inherent associated
species
(Jetz et ah 2004)
population
and
model, within
vulnerability
decline
observations
(Stattersfield
in some cases contractions leaving
et ah 1998; Manne
this from occurrence human-impacted populations
GLMMs
the pseudo-likelihood 1993) of not the that scale obtains parameter a true
(PL)
procedure
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
Criterion, were
quadratic in of models
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
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
activity positive
explored levels
among St Keough
rather development
than
tolerance were
conservation
sufficiently
(i.e.
greater
Quinn St Keough
and Neotropical between land-area observed urban
2002)
realms human (for
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
material). that
significance could
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.
threatened
ahead
predictor
for
the Nearctic
global
distribution
marked
of
threatened
spatial
avian the
species Indo
coastal
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,
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
species However,
whereas elevation
proportion
of species
and does of not
threatened
simply either threatened 1c). areas
(figure
mirror
impact These
deforestation
numbers
large-scale
productivity. consequences underlie for under contemporary between the realm of for realms pre one or
Palaearctic,
Madagascar,
the differences and the richness either those combination observed to other of
for the
species
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
variation
numbers the of
in human
threatened
impacts
species realm shows
(figure
(figure the
2a)
2b). For maximum species
and/or
example,
the
global richness
factors
human might primary remaining since we areas
Accounting
such be as
average
of
threatened
non-threatened
population
than
density,
in any other does
these being
realm. not In even
elevation
inversely of
density since
the MAM
variation
in human
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)
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
(superscripts log10-transformed.
estimate
95%
interval
expl.
deviance
GDP2 NDVI2
land-area intercept
41.4
Table
2. Minimum
(Abbreviations
birds
within
biogeographical
realms.
realm =
predictor
estimate
95%
conf.
interval
F-value
expl.
deviance
Australasian
(d.f.
932)
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.
NDVI
land-area
GDP
non-threat intercept Indo-Malayan (d.f. = 881) non-threat temperature temperature2 spp. spp.2
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
66.1
26.6
51.1
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
59.4
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
patterns, less
regional
having
considerably
power
2132
R. G. Davies
and others
104
global
thank
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,
(Oxford University),
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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