Novel methods improve prediction of species’ distributions from occurence data

Abstract

Prediction of species’ distributions is central to diverse applications in ecology, evolution and conservation science. There is increasing electronic access to vast sets of occurrence records in museums and herbaria, yet little effective guidance on how best to use this information in the context of numerous approaches for modelling distributions. To meet this need, we compared 16 modelling methods over 226 species from 6 regions of the world, creating the most comprehensive set of model comparisons to date. We used presence-only data to fit models, and independent presence-absence data to evaluate the predictions. Along with well-established modelling methods such as generalised additive models and GARP and BIOCLIM, we explored methods that either have been developed recently or have rarely been applied to modelling species’ distributions. These include machine-learning methods and community models, both of which have features that may make them particularly well suited to noisy or sparse information, as is typical of species’ occurrence data. Presence-only data were effective for modelling species’ distributions for many species and regions. The novel methods consistently outperformed more established methods. The results of our analysis are promising for the use of data from museums and herbaria, especially as methods suited to the noise inherent in such data improve.

Full text

Novel methods improve prediction of species’ distributions from
occurrence data
Jane Elith*, Catherine H. Graham*, Robert P. Anderson, Miroslav Dudı
´k, Simon Ferrier, Antoine Guisan,
Robert J. Hijmans, Falk Huettmann, John R. Leathwick, Anthony Lehmann, Jin Li, Lucia G. Lohmann,
Bette A. Loiselle, Glenn Manion, Craig Moritz, Miguel Nakamura, Yoshinori Nakazawa, Jacob McC. Overton,
A. Townsend Peterson, Steven J. Phillips, Karen Richardson, Ricardo Scachetti-Pereira, Robert E. Schapire,
Jorge Sobero
´n, Stephen Williams, Mary S. Wisz and Niklaus E. Zimmermann
Elith, J., Graham, C. H., Anderson, R. P., Dudı´k, M., Ferrier, S., Guisan, A., Hijmans, R. J.,
Huettmann, F., Leathwick, J. R., Lehmann, A., Li, J., Lohmann, L. G., Loiselle, B. A., Manion, G.,
Moritz, C., Nakamura, M., Nakazawa, Y., Overton, J. McC., Peterson, A. T., Phillips, S. J.,
Richardson, K. S., Scachetti-Pereira, R., Schapire, R. E., Sobero´n, J., Williams, S., Wisz, M. S. and
Zimmermann, N. E. 2006. Novel methods improve prediction of species’ distributions from
occurrence data. /Ecography 29: 129 /151.
Prediction of species’ distributions is central to diverse applications in ecology, evolution and
conservation science. There is increasing electronic access to vast sets of occurrence records in
museums and herbaria, yet little effective guidance on how best to use this information in the
context of numerous approaches for modelling distributions. To meet this need, we compared 16
modelling methods over 226 species from 6 regions of the world, creating the most comprehensive
set of model comparisons to date. We used presence-only data to fit models, and independent
presence-absence data to evaluate the predictions. Along with well-established modelling methods
such as generalised additive models and GARP and BIOCLIM, we explored methods that either
have been developed recently or have rarely been applied to modelling species’ distributions. These
include machine-learning methods and community models, both of which have features that may
make them particularly well suited to noisy or sparse information, as is typical of species’
occurrence data. Presence-only data were effective for modelling species’ distributions for many
species and regions. The novel methods consistently outperformed more established methods. The
results of our analysis are promising for the use of data from museums and herbaria, especially as
methods suited to the noise inherent in such data improve.
J. Elith (j.elith@unimelb.edu.au), School of Botany, Univ. of Melbourne, Parkville, Victoria, 3010
Australia. /C. H.Graham, Dept of Ecology and Evolution, 650 Life Sciences Building, Stony Brook
Univ., NY 11794, USA. /R. P. Anderson, City College of the City Univ. of New York, NY, USA. /
M. Dudı´k, Princeton Univ., Princeton, NJ, USA. /S. Ferrier, Dept of Environment and Conservation
Armidale, NSW, Australia. /A. Guisan, Univ. of Lausanne, Switzerland. /R. J. Hijmans, The Univ.
of California, Berkeley, CA, USA. /F. Huettmann, Univ. of Alaska Fairbanks, AK, USA. /J. R.
Leathwick, NIWA, Hamilton, NZ. /A. Lehmann, Swiss Centre for Faunal Cartography (CSCF),
Neuchaˆ tel, Switzerland. /J. Li, CSIRO Atherton, Queensland, Australia. /L. Lohmann, Univ.de Sa
˜o
Paulo, Brasil. /B. A. Loiselle, Univ. of Missouri, St. Louis, USA. /G. Manion, Dept of Environment
and Conservation, NSW, Australia. /C. Moritz, The Univ. of California, Berkeley, USA. /M.
Nakamura, Centro de Invest. en Matema´ ticas (CIMAT), Me´xico. /Y. Nakazawa, The Univ.of
Kansas, Lawrence, KS, USA. /J. McC. Overton, Landcare Research, Hamilton, NZ. /A. T.
Peterson, The Univ. of Kansas, Lawrence, KS, USA. /S. J. Phillips, AT&T Labs-Research, Florham
Park, NJ, USA. /K. S. Richardson, McGill Univ., QC, Canada. /R. Scachetti-Pereira, Centro de
Refereˆncia em Informac
¸a
˜o Ambiental, Brazil. /R. E. Schapire, Princeton Univ., Princeton, NJ, USA.
/J. Sobero´ n, The Univ. of Kansas, Lawrence, KA, USA. /S. E.Williams, James Cook Univ.,
Queensland, Australia. /M. S. Wisz, National Environmental Research Inst., Denmark. /N. E.
Zimmermann, Swiss Federal Research Inst. WSL, Birmensdorf, Switzerland.
*The first two authors have contributed equally to this paper.
Accepted 25 January 2006
Copyright #ECOGRAPHY 2006
ECOGRAPHY 29: 129 /151, 2006
ECOGRAPHY 29:2 (2006) 129

Detailed knowledge of species’ ecological and geo-
graphic distributions is fundamental for conservation
planning and forecasting (Ferrier 2002, Funk and
Richardson 2002, Rushton et al. 2004), and for under-
standing ecological and evolutionary determinants of
spatial patterns of biodiversity (Rosenzweg 1995, Brown
and Lomolino 1998, Ricklefs 2004, Graham et al. 2006).
However, occurrence data for the vast majority of species
are sparse, resulting in information about species dis-
tributions that is inadequate for many applications.
Species distribution models attempt to provide detailed
predictions of distributions by relating presence or
abundance of species to environmental predictors. As
such, distribution models have provided researchers with
an innovative tool to explore diverse questions in
ecology, evolution, and conservation. For example,
they have been used to study relationships between
environmental parameters and species richness (Mac
Nally and Fleishman 2004), characteristics and spatial
configuration of habitats that allow persistence of
species in landscapes (Arau
´jo and Williams 2000, Ferrier
et al. 2002a, Scotts and Drielsma 2003), invasive
potential of non-native species (Peterson 2003, Goolsby
2004), species’ distributions in past (Hugall et al. 2002,
Peterson et al. 2004) or future climates (Bakkenes et al.
2002, Skov and Svenning 2004, Arau
´jo et al. 2004,
Thomas et al. 2004, Thuiller et al. 2005), and ecological
and geographic differentiation of the distributions of
closely-related species (Cicero 2004, Graham et al.
2004b).
Most research on development of distribution model-
ling techniques has focused on creating models using
presence/absence or abundance data, where regions of
interest have been sampled systematically (Austin and
Cunningham 1981, Hirzel and Guisan 2002, Cawsey et
al. 2002). However, occurrence data for most species
have been recorded without planned sampling schemes,
and the great majority of these data consist of presence-
only records from museum or herbarium collections that
are increasingly accessible electronically (Graham et al.
2004a, Huettmann 2005, Sobero´n and Peterson 2005).
The main problem with such occurrence data is that the
intent and methods of collecting are rarely known, so
that absences cannot be inferred with certainty. These
data also have errors and biases associated with them,
reflecting the frequently haphazard manner in which
samples were accumulated (Hijmans et al. 2000, Reese et
al. 2005). Thus, the considerable potential of occurrence
data for analysis of biodiversity patterns will only be
realised if we can use them critically. Simultaneous with
increasing accessibility of species’ occurrence data,
environmental data layers of high spatial resolution,
such as those derived from satellite images (Turner et al.
2003) and through sophisticated interpolation of climate
data (Thornton et al. 1997, Hijmans et al. 2005), are
now much more abundant and available. In parallel,
development of methods for efficient exploration and
summary of patterns in large databases has accelerated
in other disciplines (Hastie et al. 2001), but only a few of
these have been applied in ecological studies. Given the
widespread use of distribution modelling, and the
synergy of advances in data availability and modelling
methods, a clear need exists for broad synthetic analyses
of the predictive ability and accuracy of species’ dis-
tribution modelling methods for presence-only data.
There is now a plethora of methods for modelling
species’ distributions that vary in how they model the
distribution of the response, select relevant predictor
variables, define fitted functions for each variable, weight
variable contributions, allow for interactions, and pre-
dict geographic patterns of occurrence (Guisan and
Zimmerman 2000, Burgman et al. 2005, Wintle and
Bardos in press). Initial attempts to analyze presence-
only data used methods developed specifically for that
purpose, based either on calculations of envelopes or
distance-based measures (Go´ mez Pompa and Nevling
1970, Rapoport 1982, Silverman 1986, Busby 1991,
Walker and Cocks 1991, Carpenter et al. 1993). Atten-
tion then turned to adapting presence-absence methods
(i.e. those that model a binomial response) to model
presence-only data, using samples of the background
environment (random points throughout the study area),
or of areas designated as ‘‘non-use’’ or ‘‘pseudo-
absence’’ (Stockwell and Peters 1999, Boyce et al. 2002,
Ferrier et al. 2002a, Zaniewski et al. 2002, Keating and
Cherry 2004, Pearce and Boyce in press).
More recently several novel modelling methods have
been proposed that have foundations in ecological and/
or statistical research, that may perform well for
distribution modelling with noisy data, such as pre-
sence-only records. Some of these methods use informa-
tion on the presences of other members of the
community to supplement information for the species
being modelled. Community methods are promising,
especially for rare species, because the additional in-
formation carried by the wider community may help
to inform the modelled relationships. Further, exten-
sive research in the machine-learning and statistical
disciplines has produced methods that are able to
capture complex responses, even with noisy input data.
These have received very little exposure in distribution
modelling, though the work that has been done is
promising (Phillips et al. 2006, Leathwick et al. in press.).
Regardless of the modelling method chosen, a major
problem in evaluation is that species’ distributions
are not known exactly. In many instances evaluation
focuses on predictive performance, some known occur-
rences are withheld from model development (either by
splitting the data set, k-fold partitioning, or bootstrap-
ping; Fielding and Bell 1997, Hastie et al. 2001, Arau
´jo
et al. 2005), and accuracy is assessed based on how
well models predict the withheld data (Boyce et al. 2002,
130 ECOGRAPHY 29:2 (2006)

Hirzel et al. unpubl.). In presence-only modelling,
such withheld data are unlikely to provide a general
test of model accuracy in predicting species’ distribu-
tions, because the occurrence records often have biases in
both geographic and environmental space (Bojo´ rquez et
al. 1995, Hijmans et al. 2000, Soberon et al. 2000,
Kadmon et al. 2004) and such biases will persist in
common resampling designs. More importantly, with-
held data are presence-only, which limits the options
for, and power of, statistical evaluations of predictive
performance. One step towards improving evaluation of
model performance in predicting distributions of species
is to use independent, well structured presence-absence
datasets for validation. Such datasets have rarely
been used to evaluate predictions from presence-
only models (but see Ferrier and Watson (1997) for an
earlier example of this type of evaluation). Further
possibilities include modelling artificial data and asses-
sing whether responses are correctly predicted (Austin et
al. 1995), or modelling with both presence-only and
presence-absence data and comparing the fitted func-
tions.
The primary aim of this research was to evaluate the
capacity of presence-only occurrence data for predicting
species’ distributions. We chose to focus on evaluation at
independent sites, using the performance of different
modelling methods averaged over many species. Detailed
evaluation of the ecological realism of all models was not
practical. We tested the performance of a representative
selection of modelling methods for presence-only data,
using data sets typical of the types of species and
environmental data that are commonly employed. This
exercise also provides insights into whether and how
these increasingly available data can be used for improv-
ing knowledge of species’ ranges. We compiled data only
for regions in which independent presence-absence data
were available for evaluation and invited participation
from researchers experienced in a range of distribution
modelling methods, including several novel methods that
have not been used widely in ecology. Our model
comparison is very broad, applying 16 methods to
modelling the distributions of 226 species from 6 regions
around the globe.
Materials and methods
Data for modelling and evaluation
Our intention in selecting data was to collect represen-
tative examples of the types of data and species that are
commonly used in species distribution modelling. Spe-
cies locality data were assembled for six regions of the
world (Tables 1 and 2): birds and plants of the
Australian Wet Tropics (AWT); birds of Ontario, Ca-
nada (CAN); plants, birds, mammals and reptiles of
north-east New South Wales, Australia (NSW), plants of
New Zealand (NZ), plants from five countries (see
footnote, Table 1) of South America (SA), and plants
of Switzerland (SWI). The individual data sets had up to
54 species with a range of geographic extents and
rarities. For each region, two sets of data were available:
1) presence-only (PO) data, from unplanned surveys or
incidental records, including those from museums and
herbaria; and 2) independent presence-absence (PA) data
from planned surveys, with accurate location records.
The independence of these two data sets for each region
is an important underlying assumption of this study, and
the totally different data collection methods for the two
sets give us a high level of confidence that this assump-
tion is well founded. Table 2 summarises the key
characteristics of the data. One feature is that the
accuracy and sample size of the modelling data vary
between regions: mostly they were small-to-moderate
(10
1
/10
2
occurrences), but were larger (10
1
/10
3
)for
SWI. Evaluation (PA) sets were larger for CAN, NZ and
SWI than the other regions. Further, the PA data sets for
AWT, NSW, NZ and SWI all provide better environ-
mental and geographic coverage than those for CAN
and SA.
The environmental data used for each region were
selected for their relevance to the species being modelled
(Austin 2002), as determined by the data provider
(Tables 1 and 3). Eleven to 13 predictors were supplied
per region, with grid cell sizes ranging from ca 100 by
100 m (AWT, NSW, NZ, SWI) to 1000 by 1000 m (CAN,
SA). Several regions had data sets where previous
research informed the development of ecologically
relevant predictors (NSW, NZ, SWI). Other regions
simply used variables typical of those used in distribu-
tion modelling, with emphasis on climatic data (CAN,
SA, AWT). None of the exact data sets used in our study
have previously been used for modelling, but some
published studies (Ferrier and Watson 1997, Guisan et
al. 1998, Venier et al. 2001, Leathwick 2002) use subsets
of these or closely related data.
The data as supplied required considerable grooming
and manipulation to generate datasets of consistent
quality both within and between regions. For each
region, environmental predictor variables were manipu-
lated so that projections, grid cell size and alignment,
and spatial extent were consistent across all layers. Some
species data (particularly in the PO datasets) had many
records per grid cell because of either repeat observa-
tions over years, or sites in close proximity to each other.
We reduced both the PO and PA datasets to one record
per grid cell. For the PA data, if any record in the grid
cell was a presence, presence was assigned to the one
record that was kept regardless of whether there was also
an absence in the same grid cell. This was not a common
feature of the data, but it occurred at least once in all
data sets except SA. In a small number of cases both a
PO and PA record occurred in the same grid cell; in these
ECOGRAPHY 29:2 (2006) 131

the PA record was usually deleted in order to ensure that
there was no spatial overlap between the modelling and
evaluation data sets. The exception was for the AWT
data, where the PO record was deleted due to limited PA
data.
As some of the modelling methods required data akin
to absences, background samples (sometimes also re-
ferred to as ‘‘pseudo-absences’’; Ferrier et al. 2002a)
were generated by drawing a random sample of 10 000
sites for each region. These were intended as a sample of
the whole region, and it is possible that a background
sample coincided with a presence record. In another
study we address alternative strategies for creating such
data (and see Zaniewski et al. 2002, Graham et al.
2004a). It is important to note that the community
methods (MARS-COMM and GDM) described below
use the species data differently to single species methods
(see Appendix, Text S1). In particular, they define a
model using all sites available for all the species of the
relevant biological group, and assume absences at sites if
presence is not recorded, effectively treating all presence
sites for a group of species as indicating an absence for
any species not recorded at a particular site. To preserve
as much consistency as possible between the single
species and community modelling approaches, we used
background samples in addition to the community data
for fitting the community models. Nevertheless, the
inclusion of the community data sets these apart from
the single-species methods.
Modelling methods
Eleven distinct modelling methods were used, but a
number of these were implemented in more than
one way, resulting in the 16 approaches presented here
(Table 4 and Appendix, Text S1). The methods form two
broad groups based on the type of data they use: those
that only use presence records (BIOCLIM, DOMAIN,
LIVES), and those that characterise the background
with a sample (all other methods). Among the techni-
ques that characterise the background, a critical distinc-
tion exists between those that use only the presence
records for the modelled species vs those that use data
describing the presences of other species, i.e. community-
based techniques. Details and key references for each
method are in Appendix, Text S1 and Table 4, and the
following briefly introduces the methods, with more
detail for lesser-known techniques.
The first group of methods (those that only use
presence records) includes one envelope-style method
(BIOCLIM) that characterises sites that are located
within the environmental hyper-space occupied by a
species, and two distance-based methods (DOMAIN
and LIVES) that assess new sites in terms of their
environmental similarity to sites of known presence. The
two distance-based methods differ both in their theore-
tical assumptions and the procedures used for calculat-
ing similarities.
The second group of methods includes several regres-
sion approaches (Table 4). Generalised linear models
Table 1. Summary of data available for modelling and evaluation.
Region Species records Predictor variables
Groups
(number species)
PO
1
:
number (mean)
PA
1
: number sites;
mean
Broad class Cell size (m) and
extent (km
2
/10
6
)
Australian Wet Tropics (AWT) birds (20) 155 340; 97 10 climate, 80
/80 m
plants (20) 35 102; 30 3 topography 0.024
Ontario, Canada (CAN) birds (20) 255 14571; 1282 6 climate,
3 topography,
1 distance,
1 vegetation
1
/1km
1.088
New South Wales, birds (10) 162 mean
3
920; 74 5 climate, 2 soil, 100/100 m
Australia (NSW) plants (29) 22 mean
3
1333; 214 1 site moisture, 0.089
mammals (7) 27 570; 76 3 topography,
reptiles (8) 84 1008; 62 1 disturbance,
1 vegetation
New Zealand (NZ) plants (52) 18 /211 19120; 1801 8 climate, 100 /100 m
2 substrate, 0.265
3 topography
South America
2
(SA) plants (30) 17 /216 152; 12 11 climate 1 /1km
14.654
Switzerland (SWI) plants (30) 36 /5822 10013; 810 7 climate, 100/100 m
2 substrate, 0.041
2 topography,
2 vegetation
1
PO is the presence-only modelling data and PA, the presence-absence evaluation data.
2
Five countries: continental Brazil, Ecuador, Colombia, Bolivia, and Peru.
3
Means of subsets of sites appropriate to different groups of plants and birds (Table 2).
132 ECOGRAPHY 29:2 (2006)

Table 2. Species data.
Region Species information PO data PA data
AWT 20 birds and 20 plants. Birds from incidental surveys (accurate locations
1
); plants from
herbarium records within last 40 yr (9
/3 km) #presence records:
Birds mean 155, range 32/265; 78% occupied cells have /1
species. Plants: mean 35, range 9 /74; 37% occupied cells have /1
species.
Birds from planned field surveys at 340 sites (accurate
locations); plants from planned surveys over 20 yr at 102 sites
(accurate locations) #presence records: birds mean 97, range
32 /265; plants: mean 30, range 13 /214.
CAN 20 birds. Ontario Nest Records, Royal Ontario Museum (ROM).
Temporal Span 1870 /2002 (mostly 1960 /2001). Coordinates
derived from map by ROM; some locations with GPS. #presence
records: mean: 255, range: 16 /749; 26% occupied cells have /1
species.
Breeding Bird Atlas (BBA) for Ontario. 14571 sites #presence
records: mean: 1282, range: 24/4512.
NSW 54 species: 7 bats (ba); 8 diurnal birds (db);
2 nocturnal birds (nb); 8 open-forest trees
(ot); 8 open-forest
understorey plants (ou); 7 rainforest trees
(rt); 6 rainforest understorey plants (ru); 8
small reptiles (sr).
Fauna incidental records from Atlas of NSW Wildlife; flora from
herbaria. Across all groups, #presence records: mean: 62, range:
2/426; 21% occupied cells have /1 species.
From designed surveys (Ferrier and Watson 1996, Pearce et al.
2001). Accurate locations. #of sites: ba 570, db 702, nb 1137, ot
2076, ou 1309, rt 1036, ru 909, sr 1008. Across all groups,
#presence records: mean: 148, range: 4 /693.
NZ 52 plants, mostly trees and shrubs. Herbarium data. #presence records: mean: 59, range
18 /211; 16% occupied cells have /1 species.
Designed surveys; 19 120 sites. Accurate locations. #presence
records: mean: 1801, range 20/10 581.
SA 30 plants, family Bignoniaceae. Herbarium data. #presence records: mean: 74, range
17 /216; 36% occupied cells have /1 species.
Al Gentry’s data (Missouri Botanic Gardens); 152 sites.
#presence records: mean: 12, range 7 /29.
SWI 30 trees. Forest vegetation data from non-systematic surveys. 1913 /1998;
75% post-1940. #presence records: mean: 1170, range 36/5822;
80% occupied cells have 
/1 species.
Forest inventory plots on regular lattice; 10 013 sites. Accurate
locations. #presence records: mean: 810, range: 19 /6953.
1
Locations are considered accurate if location error is estimated to be 5/100 m. Where accuracy data are not presented, no information was provided.
ECOGRAPHY 29:2 (2006) 133

(GLMs) and generalised additive models (GAMs) are
used extensively in species’ distribution modelling
because of their strong statistical foundation and ability
to realistically model ecological relationships (Austin
2002). GAMs use non-parametric, data-defined smooth-
ers to fit non-linear functions, whereas GLMs fit
parametric terms, usually some combination of
linear, quadratic and/or cubic terms. Because of their
greater flexibility, GAMs are more capable of modelling
complex ecological response shapes than GLMs (Yee
and Mitchell 1991). BRUTO provides a rapid method
to identify both the variables to include and the degree
of smoothing to be applied in a GAM model, but has
only recently been used in ecological applications
(Leathwick et al. unpubl.). Multivariate adaptive regres-
sion splines (MARS) provide an alternative regression-
based method for fitting non-linear responses, using
piecewise linear fits rather than smooth functions.
They are much faster to implement than GAMs, and
simpler to use in GIS applications when making maps of
predictions. An added feature that we investigate here is
their ability to analyse community data (Leathwick et al.
2005), i.e. to simultaneously relate variation in the
occurrence of all species to the environmental predictors
in one analysis, and then estimate individual model
coefficients for each species (MARS-COMM). Most of
our implementations of the regression methods did not
attempt to model interactions, with the exception of the
MARS models, where we allowed the fitting of first-
order interactions in single species models (MARS-
INT).
We implemented two versions of GARP: the desktop
version that has been used widely for modelling data
from natural history collections (DK-GARP), and a new
open modeller implementation (OM-GARP), that has
updated algorithms for developing rule sets. These both
use a genetic algorithm to select a set of rules (e.g.
adaptations of regression and range specifications) that
best predicts the species distribution (Stockwell and
Peters 1999).
Two methods have been developed within the machine
learning community: maximum entropy models
(MAXENT and MAXENT-T) and boosted regression
trees (BRT, also called stochastic gradient boosting).
MAXENT estimates species’ distributions by finding
the distribution of maximum entropy (i.e. closest to
uniform) subject to the constraint that the expected
value of each environmental variable (or its transform
and/or interactions) under this estimated distribution
matches its empirical average (Phillips et al. 2006). In
the MAXENT application for modelling presence-
only species’ data, choices can be made about the
complexity of the fitted functions. BRT combines
two algorithms: the boosting algorithm iteratively calls
the regression-tree algorithm to construct a combina-
tion or ‘‘ensemble’’ of trees. Regression trees are
Table 3. Environmental data.
Region Grid cell
size
Projection Variables Correlated pairs (pearson r
/0.85)
AWT 80 m UTM 10 climate from BIOCLIM (1: annual mean temp, 2: temp seasonality, 3: max temp of warmest month,
4: min temp of coolest month, 5: annual precipitation, 6: precip seasonality, 7: precip driest quarter, 8: annual
mean radiaton, 9: moisture index (MI) seasonality, 10: mean MI of lowest quarter MI), 11: slope,
12: topographic position, 13: terrain ruggedness index.
1 with 3,4;5 and 8 with 7;6 and 8 with
9;10 with 6,7,8,9.
CAN 1 km unprojected 6 climate from WORLDCLIM (1: annual mean temp, 2: april temp, 3: temperature seasonality, 4: annual
precipitation, 5: precip seasonality, 6: precip driest quarter), 7: altitude, 8: aspect (northness), 9: slope,
10: distance from Hudson Bay, 11: vegetation class (5 classes).
1 with 2,5,6,10;2 with 3,5,6,10;3 with
5,6;4 with 5,6; 5 with 6.
NSW 100 m unprojected 5 climate (mean annual rainfall, mean rainfall of driest quarter, annual mean temperature, minimum
temperature of the coldest month, annual mean solar radiation), moisture index, soil fertility, soil depth,
ruggedness, topographic position, compound topographic index, disturbance, vegetation class (9 classes).
min temp with mean temp.
NZ 100 m NZ map
grid
8 climate (mean October vapour pressure deficit at 09:00 h, vapour pressure deficit, mean annual solar
radiation, mean annual temperature, temperature seasonality, average monthly ratio of rainfall to potential
evaporation(r2pet), annual precipitation, solar radiation seasonality), age of bedrock, toxic cations in soil,
altitude, slope, hillshade.
r2pet with annual precip; mean annual
temp with altitude.
SA 1 km unprojected 11 climate variables from WORLDCLIM (1: annual mean temperature, 2: mean diurnal range,
3: temperature seasonality, 4: max temp of warmest month, 5: min temp of coldest month, 6: temp annual
range, 7: mean temp of wettest quarter, 8: annual precipitation, 9: precip seasonality, 10: precip driest quarter,
11: precip warmest quarter).
1 with 4,5,7;2 with 6,4 and 5 with 7.
SWI 100 m UTM 7 climate (1: growing days above freezing, 2: average temp coldest month, 3: days of summer frost, 4: annual
precip, 5: days rain 
/1 mm, 6: site water balance, 7: potential yearly global radiation), slope, topographic
position, soil nutrient index, calcareous bedrock, broadleaf cover, conifer cover.
1 with 2.
134 ECOGRAPHY 29:2 (2006)

used because they are good at selecting relevant variables
and can model interactions; boosting is used to
overcome the inaccuracies inherent in a single tree
model. The regression trees are fitted sequentially on
weighted versions of the data set, where the weights
continuously adjust to take account of observations that
are poorly fitted by the preceding models. Boosting can
be seen as a method for developing a regression model in
a forward stage-wise fashion, at each step adding small
modifications in parts of the model space to fit the data
better (Friedman et al. 2000). When using BRT, we
avoided overfitting by using cross-validation to progres-
sively grow models while testing predictive accuracy on
withheld portions of the data.
Finally, generalised dissimilarity models (GDM)
model spatial turnover in community composition (or
‘‘compositional dissimilarity’’) between pairs of sites as a
function of environmental differences between these
sites. The approach combines elements of matrix regres-
sion and generalised linear modelling, thereby allowing
it to model non-linear responses to the environment that
capture ecologically realistic relationships between dis-
similarity and ecological distance (Ferrier 2002, Ferrier
et al. 2002b). For predicting species’ distributions,
an additional kernel regression algorithm (Lowe 1995)
is applied within the transformed environmental space
generated by GDM, to estimate likelihoods of occur-
rence of a given species at all sites. Two versions of this
approach were applied in the current study: 1) ‘‘GDM’’
in which a single GDM was fitted to the combined
data for all species in a given biological group, such
that the output from this GDM was then used as a
common basis for all of the subsequent kernel regression
analyses; and 2) ‘‘GDM-SS’’ in which a separate GDM
was fitted to the data for each species alone, such that
kernel regression analysis for each species was based on
the output from a GDM tailored specifically to that
species.
As the manner in which a particular method is
implemented can have substantial effects on model
performance, we deliberately used experienced analysts
to develop all models. We also implemented batch
processing of methods in order to run the large number
of models presented here, using settings judged by the
modellers to provide a robust and reliable implementa-
tion of the method. Details are contained in Appendix,
Text S1 and Table S1.
Experimental design
All analyses were carried out by modellers (Appendix,
Table S2) blind to the evaluation data, which was not
examined until after all modelling was complete. We
provided modellers with the presence-only (PO) loca-
tions for each species and random background locations
(one set for each of the 6 regions). For each region,
environmental data were presented in two formats: either
as a table that included the environmental variables for
each PO and random locality, or as environmental grids.
Table 4. Modelling methods implemented.
Method Class of model,
and explanation
Data
1
Software Std
errors?
2
Contact
person
BIOCLIM envelope model p DIVA-GIS no CG, RH
BRT boosted decision trees pa R, gbm package no JE
BRUTO regression, a fast implementation
of a gam
pa R and Splus, mda package yes JE
DK-GARP rule sets from genetic algorithms;
desktop version
pa DesktopGarp no ATP
DOMAIN multivariate distance p DIVA-GIS no CG, RH
GAM regression: generalised additive model pa S-Plus, GRASP add-on yes AG,AL,JE
GDM generalised dissimilarity modelling;
uses community data
pacomm Specialized program not general released;
uses Arcview and Splus
no SF
GDM-SS generalised dissimilarity modelling;
implementation for single species
pa as for GDM no SF
GLM regression; generalised linear model pa S-Plus, GRASP add-on yes AG,AL,JE
LIVES multivariate distance p Specialized program not general released no JLi
MARS regression; multivariate adaptive
regression splines
pa R, mda package plus new code to handle
binomial responses
yes JE, FH
MARS-
COMM
as for MARS, but implemented with
community data
pacomm as for MARS yes JE
MARS-INT as or MARS; interactions allowed pa as for MARS yes JE
MAXENT maximum entropy pa Maxent no SP
MAXENT-T maximum entropy with threshold
features
pa Maxent no SP
OM-GARP rule sets derived with genetic
algorithms; open modeller version
pa new version of GARP not yet available no ATP
1
p/only presence data used; pa/presence and some form of absence required /e.g. a background sample; comm/community
data contribute to model fitting.
2
any method can have an uncertainty estimate derived from bootstrapping the modelling; these data refer to estimates that are
available as a statistical part of the method.
ECOGRAPHY 29:2 (2006) 135

The modellers could use the modelling data in any way
they chose, to decide how to develop their models (e.g.
they could run cross-validation testing on the PO data).
Based on these decisions, modellers made predictions for
each species using a set of evaluation data, one set of
which was prepared for each region. This consisted of a
table of environmental data for the evaluation sites, but
it contained no species records, i.e., predictions were
made from the PO data with no knowledge of the
pattern of the species as described by the PA dataset.
Modellers who applied more than one method to the
data (Appendix, Table S2) approached each method as a
new situation and aimed to implement the method in the
best way possible for a multi-species analysis (Appendix,
Text S1).
Evaluation
The evaluation focussed on predictive performance at
sites. We used three statistics, the area under the Receiver
Operating Characteristic curve (AUC), correlation
(COR) and Kappa, to assess the agreement between
the presence-absence records and the predictions. AUC
has been used extensively in the species’ distribution
modelling literature, and measures the ability of a model
to discriminate between sites where a species is present,
versus those where it is absent (Hanley and McNeil
1982). This provides an indication of the usefulness of
the models for prioritising areas in terms of their relative
importance as habitat for the particular species. The
AUC ranges from 0 to 1, where a score of 1 indicates
perfect discrimination, a score of 0.5 implies predictive
discrimination that is no better than a random guess,
and values B
/0.5 indicate performance worse than
random. ‘‘Worse than random’’ can occur because a
model may fit the modelling data but predict badly, and
we tested predictive performance on independent data
rather than model fit. AUC values can be interpreted as
indicating the probability that, when a presence site and
an absence site are drawn at random from the popula-
tion, the first will have a higher predicted value than the
second. It is closely related to a Mann-Whitney U
statistic, and it is in this context it is seen to be a rank-
based statistic /the prediction at the presence site can be
higher than the prediction at the absence site by a small
or large amount, and the value of the statistic will be the
same. Standards errors were calculated with the methods
of Hanley and McNeil (1982).
The correlation, COR, between the observation in the
PA dataset (a dichotomous variable) and the prediction,
is known as the point biserial correlation, and can be
calculated as a Pearson correlation coefficient (COR;
(Zheng and Agresti 2000)). It is similar to AUC, but
carries with it extra information: instead of being rank-
based, it takes into account how far the prediction varies
from the observation. This gives further insight into the
distribution of the predictions, and in the technical
evaluation framework of Murphy and Winkler (1992)
further informs the user about the model’s discrimination.
Kappa (Cohen 1960), which is a chance-corrected
measure of agreement, is commonly used in ecological
studies with presence-absence data. It requires a thresh-
old to be applied to the predictions, to convert them to
presence-absence predictions. Kappa provides an index
that considers both omission and commission errors. We
calculated a maximum kappa (KAPPA) for each model
by calculating kappa at all possible thresholds for each
species-specific set of predictions and identifying both
the maximum kappa and the threshold at which this
occurred. This method used information not available to
the modeller /i.e. information from the evaluation data
set. It returns the best possible KAPPA score for each
method. Liu et al. (2005) have demonstrated that other
methods are more reliable for selecting thresholds from
the training data (i.e. that used in model development),
but in this case we wanted to characterise the best
possible KAPPA value that could be attained on these
evaluation data, so that threshold selection did not
confound the results.
Variation in AUC and COR values were analysed
using Generalized Linear Mixed Models, with the
statistic (e.g. AUC) as the response and modelling
method fitted as a fixed effect. Both species and an
interaction between modelling method and region were
fitted as random effects, the interaction term allowing
for differing performance of methods across regions.
Analyses were performed using WinBUGS (Spiegelhal-
ter et al. 2003a), which fits a Bayesian model. We
assumed uninformative priors for all parameters, result-
ing in a GLMM that is equivalent to one fitted using
maximum likelihood. Comparisons of the sixteen meth-
ods were summarized from 50 000 Monte Carlo itera-
tions after a burn-in period of 10 000. The performance
of modelling method was summarized as the mean and
standard deviation of the posterior distributions. The
percentage of runs where the response (e.g. AUC) for
method X is greater than that for method Y estimates the
probability that the true difference between the methods
is greater than zero. This is a 2-tailed test, and values
close to 1 mean that method X’s response is greater than
that of method Y, and vice-versa for values close to
zero. The importance of each term in the GLMM
was assessed by change in the Deviance Information
Criterion (DIC, Spiegelhalter et al. 2003b) for the full
model compared with subsets where each term was
excluded from the model. The DIC is the Bayesian
equivalent of Akaike’s Information Criterion, and rules
of thumb suggest that changes in DIC of 
/10 units
indicate that the excluded term had an important effect
(Burnham and Anderson 2002, McCarthy and Masters
2005).
136 ECOGRAPHY 29:2 (2006)

To further explore the results, we calculated a series of
metrics that define the distances between sites, and the
area occupied, in both environmental and geographic
space. These are useful for assessing to what extent a
species ‘‘fills’’ the regional space. The nearest neighbour
(NN, also called p-median) summarises the distances
between points in multidimensional environmental
space, using a Manhattan distance (Sneath and Sokal
1973). We used the 10 000 random points and the
presences in the evaluation data set and calculated the
median of the minimum distances between any one
random point and all the presence points. In each
dimension, the distance was scaled by the range of the
random points. NN values are larger for species that
occupy only part of the environmental space of the
region. The range overlap method used the same sets of
points but summarised the mean overlap in ranges over
all environmental dimensions; high values indicate
species that span most of the environmental space in
the regions. In contrast to those two methods, maximum
distance and area of convex polygons are measures in
geographic space, calculated on presence only data but
presented in relation to the maximum measures in the
presence absence data for each biological group in each
region. Small distances or areas indicate species that are
restricted geographically.
Finally, we recorded time taken for modelling and
made comments on potential improvements, and these
are presented in Appendix, Table S2.
Results
To display mapped model results, we show distributions
predicted with several modelling methods for species in
NSW (Fig. 1 and Appendix, Fig. S1). These maps
illustrate variation in model predictions among techni-
ques. There is considerable agreement between some
methods. The most obvious differences are in the
proportion of the region that appears to be predicted
most suitable for the species, and although these might
stem from predictions that are scaled in different ways,
the variation in AUC suggests actual differences between
methods.
Evaluating the results at independent sites across all
species and methods, we found clear indications that
presence-only data can provide the basis for accurate
predictions, but also marked variation in modelling
success (Fig. 2 and Appendix, Table S3). For example,
although AUC varied from 0.07 to 0.97, 40% of models
had an AUC 
/0.75 (a useful amount of discrimination;
see methods), and 90% of methods performed better
than random (AUC 
/0.5). The following analyses
explore trends and sources of variation among methods,
regions and species.
A) Broad trends across regions and species
Assessments of modelling success using AUC and COR
indicate that methods can be analysed in three groups
(Fig. 3 and Appendix, Fig. S2). The first and highest-
performing group (above-right of the solid black line in
Fig. 3) included MARS community (MARS-COMM),
boosted regression trees (BRT), generalised dissimilarity
(GDM and GDM-SS) and maximum entropy (MAX-
ENT and MAXENT-T) models, all of which performed
relatively well according to each of the evaluation
measures (Figs 3 and 4). A second group of methods
(between the solid and dashed diagonal black lines, Fig.
3) showed intermediate performance for AUC and COR.
It included most of the standard regression methods
(generalised additive models /GAM and BRUTO;
generalised linear models /GLM; individual multi-
variate adaptive regression splines /MARS), and OM-
GARP. Models in the third group all performed
relatively poorly (lower left of the dashed black line,
Fig. 3) and included the 3 methods that use only
presence data (BIOCLIM, LIVES, DOMAIN) with no
inferred absences, along with DK-GARP, and the
MARS individual models fitted with interactions
(MARS-INT). This group also deviated from the gen-
erally linear relationship between AUC and COR results,
i.e. assessment of their predictive success depends on
which measure is used. DOMAIN was close to the
middle group in relation to AUC but not COR, and DK-
GARP was close for COR but less successful with AUC.
DK-GARP results are not completely comparable to the
others because this method could not be applied to for
one region (NZ) because of computational constraints.
The GLMMs confirmed our graphical interpretation
and indicated that differences in the performance of
modelling methods were statistically important, because
the removal of the modelling method term from the full
model resulted in a change in the Deviance Information
Criterion of 472 units (Table 5). These results are for
AUC, and those for COR are comparable. Results from
the pair-wise comparison of methods (Table 6) indicate
that those methods occurring to the top right in Fig. 3
provide significantly better performance that those to the
lower left, and also to several methods located in the
central part of this figure. The error bars in Fig. 3 are
those estimated from the model, and bars that do not
overlap coincide with high probabilities that the methods
are different (Fig. 3, Table 6).
The same general trends are evident also for KAPPA
(Fig. 4), although methods are not so clearly separated
because KAPPA is a less sensitive measure, estimating
performance at a single prediction threshold. Never-
theless, the highest-performing methods as assessed by
AUC and COR also had the highest KAPPA scores (to
the right in Fig. 4) and the presence-only methods also
ranked among the lowest.
ECOGRAPHY 29:2 (2006) 137

B) Comparison of regional results
Predictive success varied markedly across regions (Fig. 5,
Appendix, Figs S3 and S4; Table 7 and Appendix, Table
S3). For example, AUC scores were generally high for
SWI and SA (mean values of 0.77 and 0.76, Table 7, and
details in Appendix, Table S3), intermediate for NSW,
NZ and AWT (mean AUC 0.69, 0.71, 0.67 respectively)
and mostly poor for CAN (mean AUC 0.58). Regional
differences as assessed using COR (Table 7, Appendix,
Table S3) and KAPPA (Appendix, Tables S3 and S4)
indicated a slightly different ordering of regions: SA and
AWT were better modelled than the next group (SWI,
NSW, NZ), but again predictions for CAN species were
generally poor. The relative success of different methods
did not differ greatly across the three test statistics,
therefore we focus on AUC statistics for the remainder of
this section.
In some regions there were relatively large differences
in mean AUC among methods, whereas in others
differences were more muted, as shown by the vertical
spread of the lines in Fig. 5 (and see Table 7). When data
are analysed within regions there is less power to detect
differences, so error bars were relatively larger (compare
Fig. 3 and Appendix, Fig. S5) and fewer pairwise
differences between methods were significant. Our
Fig. 1. Maps for two species from NSW for each of three selected techniques. Details: ousp6, Poa sieberiana (53 records for
modelling and 512 presence/797 absence for evaluation); srsp6 Ophioscincus truncatus (79 model, 74/932 eval). The first column
shows modelling sites (grey triangles) and evaluation sites: presence
/black circle, absence/black cross.
138 ECOGRAPHY 29:2 (2006)

ability to clearly distinguish between methods was partly
related to the amount of evaluation data; SWI, NZ, and
CAN had more data (i.e. number of records per species
is to the right of Appendix, Fig. S6, leading to lower
standard errors, Appendix, Fig. S7), and more records
provide more opportunity to find differences in these
regions. For example, Table 8 presents pairwise differ-
ences for SA and SWI, and demonstrates more distinc-
tions between methods in SWI. Nevertheless, relative
rankings of methods were broadly consistent across
regions (Fig. 5, Table 7, and see Appendix, Table S3,
Fig. S5) and the group of highest-performing methods
identified in Fig. 3 was generally reliable across all
regions, but with some variation depending on the
evaluation statistic (Appendix, Fig. S5).
Some interesting patterns and exceptions to model
performance by region were apparent. Performance of
methods in NSW, NZ, SA, and SWI was generally
similar, with BRT, MAXENT and MAXENT-T,
MARS-COMM, and GDM and GDM-SS usually
performing well. In NZ, presence-only methods (BIO-
CLIM, DOMAIN and LIVES) performed particularly
poorly. In most regions DOMAIN and LIVES had
lower COR values in relation to AUC than the average
method (i.e. they sit below the line fitted to the means in
Appendix, Fig. S5).
The importance of the interaction term (method 
/
region) in the GLMM (Table 5) indicated anomalies in
the relative performance of methods in particular
regions. These tended to occur in regions with lower
overall performance /i.e. to the right in Fig. 5 (see also
Table 7 and Appendix, S3), and with highest uncertainty
in estimates (see standard errors, Appendix, Fig. S5). For
example, in AWT, OM-GARP ranked with the better
methods (GDM-SS and MAXENT-T), whereas it gen-
erally had only intermediate overall performance in all
other regions (Fig. 5, Table 7). However, AUC only
varied from 0.64 to 0.70 in AWT and many differences
were not statistically important (Appendix, Fig. S5).
Canada had the lowest AUC, COR and KAPPA scores
of any region, and many methods performed poorly,
with evaluation statistics only marginally better than
random (Table 7 and Appendix, Table S3; Figs S3, S4
and S5). Of the two methods with the highest AUC
scores in CAN, one (MARS-COMM) ranked consis-
Fig. 2. Distribution of AUC for all species and from all
methods. The solid curve is a density plot, and the y axis is
scaled to show the relative density of points; for the histogram
bars the sum of the area below all bars is 1. The grey vertical
solid line shows random predictions, and grey dashed indicates
reasonable predictive performance.
Fig. 3. Mean AUC vs mean
correlation (COR) for modelling
methods, summarised across all
species. The grey bars are standard
errors estimated in the GLMM (see
Appendix), reflecting variation for
an average species in an average
region. The labels are broad
classifications of the methods: grey
underlined
/only use presence
data, black capitals
/use presence
and background samples, black
lower case italics
/community
methods.
ECOGRAPHY 29:2 (2006) 139

tently among the best across regions, whereas the other
(BIOCLIM) tended to be among the lowest.
C) Results at the species level
The greatest variation in the performance of different
methods was apparent at the species level, but with
similar trends: methods shown above to perform well
when averaged across species and regions (i.e. MARS-
COMM, BRT, MAXENT/MAXENT-T and GDM/
GDM-SS) also tended to perform well when ranked
against other methods on an individual species’ basis
(Table 9 and Appendix, Fig. S2). Figure 6a shows results
from SA that are typical for most regions, in that there is
marked variation in which methods perform best for
different species (i.e. lines cross in the graphs). Most
methods occasionally failed badly, although the better
methods tended to have more stable performance. The
exception to this general pattern was SWI, in which we
observed clear and reasonably consistent separation
between methods (Fig. 6b), probably reflecting the larger
amounts of accurately located data for both modelling
and evaluation. Results for SWI using both AUC and
COR indicate that the highest-performing methods for
most species were MARS-COMM, BRT and MAXENT/
MAXENT-T, while the lowest were LIVES, BIOCLIM,
and DK-GARP (Appendix, Fig. S5). Across all species
in all regions, and considering the best method only, 78%
of species had AUC scores of 0.70 or more, and 64% had
scores of 0.75 or more (Appendix, Table S3). In some
cases a species was modelled well (or poorly) by most
methods, whereas in others there was considerable
variation in predictive accuracy depending on the
method used. Mean AUC scores per species varied
from 0.36 to 0.97, with coefficients of variation (cv)
ranging from 2 to 47% (Appendix, Table S3). The
correlation between mean AUC and cv was weakly
negative (Pearson r
//0.39), indicating a slight trend
for more variation across methods for species with low
mean AUC scores.
Predictive performance did not vary consistently
with number of presence records available for modelling
(Table 10, Fig. 7, Appendix, Table S5). Species that
are rarer because they are environmentally or geogra-
phically restricted appeared to be modelled with greater
accuracy than more common and generalist species
(Table 10). We note here and discuss later that this
result depends on the spatial extent of analysis and the
type of evaluation.
Discussion
The model comparison developed herein is unique for
its broad geographic scope, application of numerous
modelling methods (including several new techniques) to
Table 5. Analysis of importance of factors affecting predictive
performance, from Generalized Linear Mixed Model.
DIC
1
Full model:AUC /Method/Method /Region

/Species

/8996
Without Method 
/8524
Without Interaction (Method
/Region) /8803
Without Species 
/4071
1
Deviance Information Criterion. Changes in DIC /10 are
important.
Fig. 4. Performance measured by
maximum kappa and its variance
across all species. Variance axis
reversed so low is higher on plot; it
is desirable to have high kappa and
low variance (i.e. upper right in
plot), for consistent and good
performance. Labels as for Fig. 3.
140 ECOGRAPHY 29:2 (2006)

the analysis of presence-only data, and incorporation of
extensive presence/absence data to enable well-informed
evaluations of predictive performance. We interpret
results in the context of the feasibility of using such
techniques to predict accurately species’ distributions in
situations in which only presence data are available. Our
evaluation also explores the utility of the vast storehouse
of occurrence information in resources such as world
natural history museums.
We can draw two major conclusions from our results.
First, presence-only data are useful for modelling
species’ distributions. This result bolsters the recent
movement to capitalize on the growing availability of
both species’ occurrence records (Soberon et al. 2000,
Graham et al. 2004a) and high-resolution spatial envir-
onmental data. Not all species were predicted well
according to our evaluation data, but we found that
64% of the best models for each species had AUCs

/0.75 and an additional 14% had AUCs between 0.7
and 0.75. These AUC scores indicate that predictions
based on presence-only data can be sufficiently accurate
to be used in conservation planning (Pearce and Ferrier
2000a) and in numerous other applications in which
estimates of species’ distribution are relevant. Second,
new modelling methods that have only recently been
applied to the challenge of modelling species’ distribu-
tions generally outperformed established methods. Some
of these new methods originated in other disciplines and
have had little exposure in ecological analyses. These
methods appear to offer considerable promise across a
much broader range of ecological applications, providing
an exciting avenue for future research. The other strong
performers were community methods, and these also
deserve further scrutiny, particularly where data for the
species in question are sparse.
Broad comparison of methods
We demonstrated differences in predictive performance
among modelling methods, despite substantial variation
at both regional and species levels. Within the suite of
relatively commonly used methods, those that character-
ise the background environment and that can differen-
tially weight variables outperform those that use
presence data alone (BIOCLIM, LIVES, and /for
some measures /DOMAIN). These results give no
support to using methods that do not attempt to
characterise the distribution of a species relative to the
background environment in which it occurs. The various
regression-based methods are largely indistinguishable
from one another in terms of predictive performance.
The new version of GARP (OM-GARP), first imple-
mented for this study, is comparable to, but slower than,
the regression methods and outperforms the widely used
desktop version.
Table 6. Probability that the method in the column gives a higher AUC than the method in a row. Low values indicate that the method in the row tends to give higher AUCs than the
method in the column. Values outside the arbitrary limits p
/(0.025, 0.975) are highlighted in bold; for this two-tailed test.
/
BIOCLIM
/
BRT
/
BRUTO
/
DOMAIN
/
GAM
/
GLM
/
DK-GARP
/
OM-GARP
/
GDM
/
GDM-SS
/
LIVES
/
MAXENT
/
MAXENT-T
/
MARS
/
MARS-INT
/
MARS-COMM
BIOCLIM
BRT 0.000
BRUTO 0.019 0.988
DOMAIN 0.013 0.979 0.422
GAM 0.011 0.976 0.413 0.495
GLM 0.012 0.982 0.447 0.527 0.534
DK-GARP 0.200 0.999 0.882 0.914 0.916 0.909
OM-GARP 0.010 0.976 0.400 0.475 0.487 0.449 0.078
GDM 0.000 0.757 0.061 0.089 0.093 0.076 0.004 0.095
GDM-SS 0.000 0.791 0.076 0.107 0.110 0.095 0.006 0.119 0.549
LIVES 0.124 0.999 0.832 0.877 0.877 0.864 0.394 0.888 0.993 0.992
MAXENT 0.000 0.733 0.052 0.075 0.081 0.065 0.004 0.085 0.469 0.424 0.007
MAXENT-T 0.000 0.658 0.032 0.049 0.050 0.043 0.002 0.054 0.381 0.337 0.003 0.411
MARS 0.017 0.986 0.479 0.552 0.560 0.530 0.111 0.580 0.933 0.918 0.151 0.942 0.964
MARS-INT 0.169 0.999 0.877 0.914 0.916 0.900 0.465 0.924 0.996 0.995 0.576 0.997 0.998 0.888
MARS-COMM 0.000 0.425 0.009 0.015 0.016 0.012 0.000 0.016 0.190 0.157 0.000 0.213 0.279 0.010 0.000
ECOGRAPHY 29:2 (2006) 141

Results for the more common approaches are con-
sistent with previous studies of presence-absence model-
ling methods, and with the relatively few comparisons of
methods used to model presence-only data. Studies of
presence-absence modelling methods suggest that several
non-linear techniques (e.g. GAMs, artificial neural net-
works, and MARS) are comparable in terms of pre-
dictive ability, and are often superior to methods such as
traditional single decision trees (Ferrier and Watson
1997, Elith and Burgman 2002, Moisen and Frescino
2002, Mun
˜oz and Fellicı´simo 2004, Segurado and
Araujo 2004). Comparisons of methods using pre-
sence-only records are less common, but tell a similar
story: GLMs and GAMs generally outperform simpler
methods (Ferrier and Watson 1997, Brotons et al. 2004),
MAXENT outperforms GARP (Phillips et al. 2006) and
some presence-only methods (e.g. DOMAIN, ENFA,
(Hirzel et al. 2002)) have advantages over BIOCLIM
(Loiselle et al. 2003). Most of these studies, however,
have focused on single geographic regions and/or smaller
numbers of species. Several have been evaluated on the
same data sets as were used for model development, and
this makes it difficult to generalise results, and to discern
whether models have good predictive performance or
whether they are simply overfit (Leathwick et al.
unpubl.). Our use of independent presence/absence test
Table 7. Regional data: mean AUC and mean COR per method, per region.
Method Mean AUC Mean COR
AWT CAN NSW NZ SA SWI AWT CAN NSW NZ SA SWI
BIOCLIM 0.65 0.63 0.63 0.61 0.75 0.71 0.22 0.08 0.12 0.08 0.29 0.15
BRT 0.68 0.60 0.71 0.73 0.80 0.81 0.24 0.08 0.19 0.18 0.32 0.24
BRUTO 0.64 0.55 0.68 0.72 0.75 0.79 0.20 0.04 0.15 0.17 0.22 0.20
DK-GARP 0.68 0.56 0.66 na
1
0.75 0.70 0.28 0.05 0.15 na
1
0.21 0.13
DOMAIN 0.67 0.60 0.70 0.69 0.77 0.73 0.22 0.05 0.15 0.10 0.21 0.14
GAM 0.65 0.55 0.68 0.73 0.75 0.80 0.22 0.04 0.15 0.17 0.23 0.21
GDM 0.67 0.57 0.73 0.74 0.79 0.78 0.24 0.06 0.21 0.16 0.30 0.19
GDM-SS 0.70 0.56 0.70 0.73 0.79 0.79 0.29 0.04 0.17 0.15 0.28 0.20
GLM 0.66 0.57 0.68 0.71 0.74 0.78 0.24 0.06 0.16 0.16 0.22 0.19
LIVES 0.66 0.61 0.66 0.66 0.77 0.72 0.23 0.06 0.12 0.08 0.21 0.13
MARS 0.66 0.55 0.67 0.72 0.75 0.79 0.23 0.04 0.15 0.17 0.24 0.21
MARS-COMM 0.67 0.64 0.73 0.74 0.77 0.82 0.20 0.11 0.19 0.18 0.26 0.26
MARS-INT 0.65 0.54 0.64 0.70 0.73 0.78 0.22 0.05 0.14 0.16 0.24 0.21
MAXENT 0.68 0.58 0.71 0.74 0.78 0.80 0.23 0.05 0.18 0.18 0.27 0.24
MAXENT-T 0.69 0.58 0.71 0.73 0.77 0.80 0.24 0.06 0.18 0.18 0.26 0.25
OM-GARP 0.69 0.55 0.68 0.70 0.77 0.77 0.29 0.04 0.15 0.13 0.25 0.19
mean 0.67 0.58 0.69 0.71 0.76 0.77 0.24 0.06 0.16 0.15 0.25 0.20
1
DK-GARP could not be run for NZ; the large number of grid cells could not be accomodated by the available computers.
Fig. 5. Predictive success across
regions, for 10 methods. Regions
are sorted by the mean AUC
across all 16 methods and all
species per region.
142 ECOGRAPHY 29:2 (2006)

Table 8. Probability that the method in the column gives a higher AUC than the method in a row for (a) SA and (b) SWI.Format follows Table 6.
/
BIOCLIM
/
BRT
/
BRUTO
/
DOMAIN
/
GAM
/
GLM
/
DK-GARP
/
OM-GARP
/
GDM
/
GDM-SS
/
LIVES
/
MAXENT
/
MAXENT-T
/
MARS
/
MARS-INT
/
MARS-COMM
(a)
BIOCLIM
BRT 0.002
RUTO 0.653 0.999
DOMAIN 0.098 0.940 0.045
GAM 0.521 0.998 0.367 0.912
GLM 0.740 1.000 0.600 0.973 0.723
DK-GARP 0.649 0.999 0.495 0.953 0.628 0.395
OM-GARP 0.102 0.944 0.049 0.513 0.093 0.028 0.049
GDM 0.011 0.706 0.003 0.155 0.009 0.002 0.003 0.147
GDM-SS 0.024 0.814 0.009 0.254 0.022 0.004 0.010 0.243 0.638
LIVES 0.223 0.982 0.124 0.706 0.206 0.081 0.126 0.694 0.939 0.886
MAXENT 0.083 0.931 0.038 0.467 0.074 0.022 0.039 0.456 0.826 0.721 0.268
MAXENT-T 0.160 0.969 0.083 0.618 0.145 0.051 0.085 0.609 0.907 0.834 0.408 0.650
MARS 0.598 0.999 0.444 0.939 0.577 0.347 0.448 0.936 0.995 0.987 0.844 0.949 0.893
MARS-INT 0.906 1.000 0.824 0.996 0.897 0.748 0.825 0.995 1.000 1.000 0.980 0.997 0.989 0.857
MARS-COMM 0.193 0.976 0.104 0.667 0.178 0.065 0.105 0.657 0.927 0.863 0.459 0.696 0.552 0.133 0.014
(b)
BIOCLIM
BRT 0.000
BRUTO 0.000 0.995
DOMAIN 0.006 1.000 1.000
GAM 0.000 0.954 0.193 0.000
GLM 0.000 0.999 0.719 0.000 0.925
DK-GARP 0.809 1.000 1.000 1.000 1.000 1.000
OM-GARP 0.000 1.000 0.996 0.000 1.000 0.981 0.000
GDM 0.000 1.000 0.943 0.000 0.993 0.841 0.000 0.140
GDM-SS 0.000 0.992 0.439 0.000 0.762 0.231 0.000 0.002 0.041
LIVES 0.142 1.000 1.000 0.919 1.000 1.000 0.026 1.000 1.000 1.000
MAXENT 0.000 0.900 0.102 0.000 0.342 0.033 0.000 0.000 0.002 0.132 0.000
MAXENT-T 0.000 0.840 0.060 0.000 0.243 0.016 0.000 0.000 0.001 0.081 0.000 0.387
MARS 0.000 0.988 0.391 0.000 0.719 0.194 0.000 0.002 0.032 0.448 0.000 0.839 0.899
MARS-INT 0.000 1.000 0.853 0.000 0.971 0.676 0.000 0.053 0.295 0.886 0.000 0.989 0.995 0.907
MARS-COMM 0.000 0.251 0.000 0.000 0.009 0.000 0.000 0.000 0.000 0.001 0.000 0.025 0.048 0.002 0.000
ECOGRAPHY 29:2 (2006) 143

data across multiple geographic regions provides a
broader basis for comparisons.
A novel aspect of our work is the inclusion of newer
modelling methods that have had little exposure in
previous comparative studies and few applications in
ecology in general. These novel methods outperform the
established methods, and this observation should pro-
voke attention and scrutiny. Several of the novel
methods have been developed and tested in fields other
than species’ distribution modelling, and have been
shown to handle noisy data and complex analytical
challenges successfully. For example, boosted regression
trees have been a focus of attention in the machine-
learning and statistical fields for a number of years
(Ridgeway 1999, Hastie et al. 2001), but the present
paper and a companion application to New Zealand fish
(Leathwick et al. in press) are among the first in ecology.
Similarly, maximum entropy methods are well known in
other fields (Jaynes 1982) but only recently developed for
questions of species’ distributions (Phillips et al. 2006).
One question of interest is whether our ‘‘best’’
methods share certain characteristics that set them apart
from the others? One feature that they all share in
common is a high level of flexibility in fitting complex
responses. As a consequence they all have what we term
here ‘‘expressiveness’’ /a well-developed ability to
express or demonstrate the complex relationships that
exist in the data. In several methods this includes
effective mechanisms for modelling interactions among
variables. However, expressiveness needs to be controlled
so that models are not overfit, and to that end several
methods use ‘‘regularization’’ techniques (Hastie et al.
2001) to achieve balance between complexity and
parsimony.
These methods achieve those goals in different ways.
BRT achieves expressiveness by combining the strengths
of regression trees, namely omission of irrelevant vari-
ables and ability to model interactions, with those of
boosting, that is, the building of an ensemble of models
that approximate the true response surface more accu-
rately than a single model by overcoming the misclassi-
fication problems inherent in single tree models. Both the
model building procedure (a penalized forward stepwise
search) and our cross-validation methods for finding
optimal numbers of trees help to control overfitting. The
application of maximum entropy methods to distribu-
tion modelling was developed specifically for use with
presence-only occurrence data (Phillips et al. 2006). In
MAXENT, strong focus has been placed on the role of
penalty functions (i.e. regularization) in parameter
estimation. Regularization has most impact when sam-
ple sizes are small, so the MAXENT modellers tuned
their regularization in relation to sample size (Phillips et
al. 2004). MAXENT can also fit complex functions
between response and predictor variables, and can
include interaction terms but to a more limited extent
than BRT. GDM-SS models are single-species versions
of the community-based GDM models. They are para-
meterized on data for individual species, not including
community data. These models are developed in a 2-step
fashion, which together achieves controlled expressive-
ness. The first step operates effectively like a GAM,
fitting additive smoothing functions (albeit to dissim-
ilarities rather than raw observations). The second step,
a kernel regression, incorporates interactions by model-
ling distances and densities within a truly multivariate
predictor space, with no assumption of additivity. The
success of the kernel regression step depends on the first
step accurately transforming the predictor space, thereby
addressing the ‘‘curse of dimensionality’’ normally
associated with kernel regression type techniques
(Lowe 1995).
The success of these new methods suggests that
predictive performance of some more common methods
such as GAMs might be improved substantially if better
tradeoffs between expressiveness and complexity could
be incorporated into the model fitting process. In the
case of GAMs the issue is not whether it can fit complex
responses /it can /but whether the model building
methods commonly used are optimal for species dis-
tribution modelling. One aspect is modelling interac-
tions, which is possible in regression but rarely
implemented when many species are being modelled.
Another is model selection: alternative techniques al-
ready exist, but are rarely implemented in ecology. For
example, the ‘‘lasso,’’ used for regularization in MAX-
ENT, can be applied to variable selection and coefficient
estimation in regression (Hastie et al. 2001), and has
been shown to perform better than stepwise selection for
determining model complexity (Tibshirani 1996).
Ferrier et al. (2002b) developed and used GDM for
reserve selection and survey design. The impetus for
using GDM and other community modelling methods
(e.g. MARS-COMM) in distribution modelling is based
Table 9. Comparison of performance at a species level and over
all species. Rank of 1 
/highest to 16/lowest.
Method
(sorted by column 2)
Mean AUC
rank per species
Rank of mean
AUC over all species
MARS-COMM 6.15 1
BRT 6.20 2
MAXENT-T 6.42 3
MAXENT 6.69 5
GDM-SS 7.38 6
GDM 7.53 4
GAM 8.26 7
GLM 8.64 10
DOMAIN 8.70 9
BRUTO 8.79 12
MARS 8.92 8
OM-GARP 8.92 11
MARS-INT 9.72 13
LIVES 10.22 14
DK-GARP 10.47 15
BIOCLIM 10.85 16
144 ECOGRAPHY 29:2 (2006)

on the understanding that important but subtle environ-
mental trends may only be apparent in the response of
multiple species, and, more pragmatically, that rare
species are difficult to model with standard statistical
methods and, hence, community signals may improve
model performance. Community models use information
from suites of species to inform variable selection and
modelling. Relevant predictors are included because of
their strong community signal, whereas that signal might
be insufficient to trigger inclusion in single species
models (Leathwick et al. unpubl.). GDM is the more
ecologically sophisticated approach to community mod-
elling, but the simpler MARS-COMM models per-
formed at least as well in this study. The comparison
of MARS single species models with the MARS com-
munity models shows that the community models
Fig. 6. Predictive performance
for the 30 South American
species (6a) and the 30 Swiss
species (6b), for 10 methods.
Species are sorted by the mean
AUC across all 16 methods.
ECOGRAPHY 29:2 (2006) 145

perform strongly in this trial. In contrast, a recent
application of MARS and MARS-COMM to modelling
a large presence-absence freshwater fish data set from
New Zealand indicated no consistent advantage for
community models (Leathwick et al. unpubl.). This
difference probably reflects the advantage in using
community data to infer absences for presence-only
data (see Materials and methods), the importance of
the community signal in the more biased and noisier
data analyzed here, and the small amounts of data
available for fitting some species. Further comparisons
of alternative community modelling approaches would
be useful to identify the factors influencing success of the
various techniques. It is likely that the inference of
absences is important, and could be adopted as an
alternative to random background samples for single
species models.
In summary, we suggest that the good performance of
the novel methods result from their ability to fit complex
responses (often including interactions) and select a
relevant set of variables. We do not expect that the fitted
models are too complex /for example, MAXENT as
used here is similar in expressiveness to a GLM without
interactions (for species with B
/80 presences) or a GAM
with pairwise interactions (Appendix, Text S1). How-
ever, for any method an ability to fit complex responses
is not useful in itself, and has to be balanced against the
requirement for model features to be ecologically
realistic (Austin 2002). Systematic assessment of whether
individual fitted responses were realistic was beyond the
scope of the current research as our aim was to test
models across numerous species and regions. None-
theless, this would be a fruitful avenue for research and
model assessment.
Table 10. Pearson correlation coefficients between data characteristics and the maximum AUC achieved per species, summarised at
a regional level.
Sample sizes Environmental space Geographic space
P in PO Prevalence Nearest neighbour Range overlap Relative maximum distance Relative area of convex polygon
AWT 
/0.340* /0.276 0.589* /0.302 /0.456* /0.421*
CAN 0.048* 
/0.318 0.621* /0.589* /0.313 /0.304
NSW 
/0.045 /0.654* 0.675* /0.770* /0.230 /0.359*
NZ 0.028 
/0.298* 0.713* /0.669* /0.491* /0.300*
SA 
/0.601* /0.714* 0.541* /0.922* /0.667* /0.708*
SWI 
/0.186 /0.165 0.619* /0.520* /0.534* /0.620*
Asterisks indicate significant values (pB
/0.05). Prevalence is the frequency of occurrence records in the PA data. The measures are
detailed in the methods.
Fig. 7. The maximum AUC (over
all methods) vs sample size in
modelling data. Each point
represents a species.
146 ECOGRAPHY 29:2 (2006)

Regional patterns
The trend for good performance of the novel methods in
our experiment (BRT, MAXENT, GDM and MARS-
COMM) was also apparent at the regional level. In
regions such as SWI, where overall performance was
high, the ‘‘best’’ methods performed well, but most
methods were able to produce reasonable quality models
for many of the species. Exceptions to the usual ranking
of methods sometimes appeared in regions in which
overall predictive performance was generally poor
(AWT and CAN), with no clearly consistent cause. In
these regions, the good methods usually still showed
moderate success in prediction, and often were not
impacted as severely as other methods. The fact
that some regions had relatively poor average perfor-
mance and even the best methods could not do well
suggests that in such regions, modelling might most
benefit from attempts to improve the training data set.
This is particularly apparent in the CAN bird data set, in
which the data display many of the biases possible in
presence-only data (Anderson 2003). Sampling is se-
verely biased towards the southern extreme of the study
area, species’ ranges are only partially represented, and
environmental data may not capture the important
conditions affecting bird distribution, especially those
with large ranges. The effects of the partial sampling of
ranges may be similar to those demonstrated by Randin
et al. (in press) in their study of transferability of models.
One would expect the sampling bias to be particularly
problematic in cases, like CAN, where the gradient in
sampling efforts corresponds to strong environmental
gradients. As a consequence, only 4 of 20 species had a
mean ROC
/0.7, and the best methods (MARS-COMM
and BIOCLIM) had mean ROC scores of only 0.63
across species. With poor modelling success, the data
and ecology of the species need to be further investigated
to identify how to supplement or otherwise improve
both the species’ occurrence records and the predictor
variable set.
Species-level patterns
Though not a primary focus of our analysis, we can
highlight two results to guide those needing to under-
stand what species’ attributes might affect model per-
formance. Firstly, sample size (i.e. number of occurrence
records) was not consistently related to modelling
success. While other studies indicate that small sample
size negatively influences modelling success (Pearce and
Ferrier 2000a, Harrell 2001, Stockwell and Peterson
2002, Kadmon et al. 2003), this was not evident here,
perhaps because in general we had adequate localities for
modelling (mean number of modelling records 
/233;
range
/2/5822)
The other pattern that emerged is that species judged
to be specialists by the distribution of records in
environmental or geographic space tended to have higher
AUC scores than generalists, an effect also observed
elsewhere (Guisan and Hofer 2003, Segurado and
Arau
´jo 2004, Thuiller et al. 2004, Luoto et al. 2005).
However, this result requires further scrutiny because it
may well be largely a function of spatial extent of the
analysis. If the extent is fixed (say, to a region) and the
evaluation data are a constant set of sites, specialists by
definition are those that exist in only a subset of that
space. Therefore, the evaluation data set will have many
zero records for such species, and any model that can
restrict its non-zero predictions to the zone that the
species occupies will have a good AUC score, due to the
comparison with the many absence sites. The question
then becomes one of whether a constant extent of
analysis is appropriate for all species in relation to the
purpose of the predictions. This consideration needs to
be defined by the end-use of the predictions, and merits
attention in any evaluation of modelling success.
Limitations
Whereas the present study points to both the potential
informativeness of presence-only data and to improved
methods for predicting species distributions from them,
we also need to emphasize what this doesn’t tell us. Our
evaluation strategy is specific to the question of how best
to predict species distribution under current environ-
mental conditions, which has broad relevance to con-
servation and ecological or macro-ecological studies
(Ferrier 2002, Funk and Richardson 2002, Rushton et
al. 2004). Our approach, however, does not inform
selection of methods for predicting potential ranges or
extrapolation from the current to alternative climates.
Modelling of potential ranges (Sobero´n and Peterson
2005) has applications to predicting expansions of
invasive species and investigating speciation processes
(Hugall et al. 2002, Peterson 2003). However, as the true
potential range may differ from the realized range
because of dispersal limitation, competition or other
factors (Van Horne 1983, Hanski 1994, Tyre et al. 2001,
Anderson et al. 2002), evaluating model performance is
a complex task and use of observed absences may be
misleading.
Projection of modelled relationships to alternative
climates (past or future) is an increasingly popular
application of distributional modelling, e.g. in relation
to potential effects of global warming. However, strong
performance of a particular method in the present
climate does not guarantee similar performance under
different climates (Thuiller 2004), particularly where this
requires prediction outside the range of environments on
which the original model was based (Arau
´jo et al. 2005).
ECOGRAPHY 29:2 (2006) 147

Community-based models could fail if patterns of
species co-occurrence change due to idiosyncratic re-
sponses by individual species to climate change. Similar
caution should also be exercised in making predictions
for sites distant from the geographic domain from which
the modelling data were drawn, given the potential for
accurate prediction to be confounded either by unrecog-
nised environmental factors or by large-scale geographic
variation in disturbance regimes (Randin et al. in press).
Clearly, different models may be necessary, and different
approaches needed to evaluate relative performance of
modelling methods under these scenarios. Arau
´jo et al.
(2005) argue that whilst evaluation of models to new
climates and new regions is particularly difficult, evalua-
tions of models even within one region are prone to over-
optimism. This occurs when modelling and evaluation
data sets are not sufficiently independent. Because the
data in this study were sourced from collections with
substantially different survey protocols, the evaluation is
unlikely to suffer from such problems. Nevertheless,
further explorations of the ability of the novel methods
to predict reliably to unsampled areas /for example, by
spatially stratifying the evaluation data /would be
useful.
A further qualification needs to be made. Whilst these
models have been shown to produce predictions that are
useful in their ability to rank sites for relative suitability
for many species, we do not suggest that models built
with presence-only data will be well calibrated. That is,
they do not accurately predict probability of presence
because they do not have access to reliable information
on the prevalence (frequency of occurrence) of the
species in the region. Rather, they provide relative indices
of suitability. Our implementations of most of the
methods that take into account background were simple
applications of the methods, not attempting to make
sophisticated adjustments at might improve this aspect.
Alternative approaches such as case control methods for
regression (Keating and Cherry 2004, Pearce and Boyce
in press), and Bayesian methods (Gelfand et al. 2006)
continue to be developed and have potentially important
contributions because they deal more appropriately with
the presence-only paradigm.
This is by no means a complete analysis, and
important questions remain. To advance our under-
standing of the strengths and weaknesses of methods
and the differences between them requires theoretical
investigations of the techniques (Austin 2002), testing on
simulated data (Austin et al. 1995), analysis of the
modelled response shapes (e.g. Austin et al. 1994, Bio
2000, Leathwick 2002) and evaluation of the spatial
trends in errors (Fielding and Bell 1997, Barry and Elith
in press). Our method of evaluation relies on the
evaluation site data and summarises over these, but
this might mask effects that can only be clarified with
these more detailed evaluations. Operator expertise will
affect model performance, and tests of the sensitivity of
the methods to how they are applied would be informa-
tive. These were beyond the scope of this paper but are
necessary for a deeper understanding of the models and
their predictive capacities and limitations.
Finally, we stress that modelling can never provide a
complete substitute for detailed, ongoing collection
of field data, including data on species’ distribution,
demography, abundance, and interactions (Guisan
and Thuiller 2005). Modelling approaches which attempt
to integrate such information include Bayesian
approaches (Gelfand et al. 2006), investigation of
competitors (Leathwick and Austin 2001, Anderson et
al. 2002), and studies of connectivity (Ferrier et al.
2002a, Moilanen et al. 2005). To date, few models
have been validated via collection of new data (but see
Ferrier and Watson 1997, Elith and Burgman 2002,
Raxworthy et al. 2003, Peterson 2005). Likewise, colla-
borative efforts between modellers and users such
as conservation managers are rare (Pielke Jr 2003).
Ideally, models should be developed and tested in
iterative cycles that take account of the desired uses of
model, investigate the ecological rationality of the
modelled responses and explore errors in predictions
(Burgman et al. 2005, Barry and Elith in press). We hope
that our model comparisons will stimulate more research
into both modelling methods, further development of
efficient user interfaces for the more successful methods,
and greater integration among modellers and end-users.
Acknowledgements /Data were kindly provided by
organizations for whom a number of authors worked. We also
thank Mark and George Peck, Royal Ontario Museum, for
access to Ontario nest record data (B
/http://
www.birdsontario.org/onrs/onrsmain.html
/); Mike Cadman,
Bird Studies Canada, Canadian Wildlife Service of
Environment Canada, for access to BBS data; Missouri
Botanical Garden, especially Robert Magill and Trisha
Consiglio, for access to TROPICOS and Gentry transect
databases; T. Wohlgemuth and U. Braendi from WSL
Switzerland for access to the NFI and forest plots data, and
Andrew Ford, CSIRO Atherton, for AWT PA plant records.
Michael McCarthy helped with the GLMM analysis and
WinBUGs programming. The comments of Mike Austin and
Miguel Arau
´jo improved the manuscript substantially. J. E. was
funded by ARC Grant DP0209303. This research was initiated
in a working group at the National Center for Ecological
Analysis and Synthesis (NCEAS), Santa Barbara, USA:
‘‘Testing Alternative Methodologies for Modelling Species’
Ecological Niches and Predicting Geographic Distributions’’,
conceived of and led by Peterson and Moritz.
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