A Journal of Conservation Biogeography Diversity and Distributions, (Diversity Distrib.

) (2010) 16, 109–118

BIODIVERSITY Predicting the impact of climate change on

RESEARCH
Australia’s most endangered snake,
Hoplocephalus bungaroides
Trent D. Penman1*, David A. Pike2, Jonathan K. Webb2 and Richard Shine2

1
Forest Resources Research, NSW Department ABSTRACT
of Primary Industries, PO Box 100, Beecroft,
Aim To predict how the bioclimatic envelope of the broad-headed snake (BHS)
NSW 2119 Australia, 2School of Biological
Sciences A08, University of Sydney, NSW
(Hoplocephalus bungaroides) may be redistributed under future climate warming
2006, Australia scenarios.
Location South-eastern New South Wales, Australia.

Methods We used 159 independent locations for the species and 35 climatic
variables to model the bioclimatic envelope for the BHS using two modelling
approaches – Bioclim and Maxent. Predictions were made under current
climatic conditions and we also predicted the species distribution under low and
high climate change scenarios for 2030 and 2070.
Results Broad-headed snakes currently encompass their entire bioclimatic
envelope. Both modelling approaches predict that suitable climate space for
BHS will be lost to varying degrees under both climate warming scenarios, and
under the worst case, only 14% of known snake populations may persist.
Main conclusions Areas of higher elevation within the current range will be most
important for persistence of this species because they will remain relatively moist
and cool even under climate change and will match the current climate envelope.
Diversity and Distributions

*Correspondence: Trent D. Penman, Forest
Resources Research, NSW Department of
Primary Industries, PO Box 100, Beecroft, NSW
2119 Australia.
E-mail: trent.penman@industry.nsw.gov.au
Conservation efforts should focus on areas where suitable climate space may
persist under climate warming scenarios. Long-term monitoring programs should
be established both in these areas and where populations are predicted to become
extirpated, so that we can accurately determine changes in the distribution of this
species throughout its range.
Keywords
BIOCLIM, bioclimatic envelope modelling, geographical distribution, Maxent,
range contraction.

climate change may be mediated via habitat modification

INTRODUCTION
Predicted changes to global and regional climates (Pausas,
2004; Hennessy et al., 2005; Suppiah et al., 2007) are likely to
have direct and indirect effects upon biodiversity (Theurillat
& Guisan, 2001; Hughes, 2003; Meynecke, 2004). Shifts in
environmental conditions can alter critical biological traits
such as the timing and magnitude of reproductive output
(Forchhammer et al., 1998; Crick & Sparks, 1999; Winkler
et al., 2002; Pike et al., 2006), sex ratios of offspring (Janzen,
1994; Hays et al., 2003; Mitchell et al., 2008), resource
availability (Visser & Both, 2005), habitat use (including nest
site selection; Telemeco et al., 2009) and rates of survival
(Chamaille-Jammes et al., 2006). The greatest impacts of
(Meynecke, 2004). Some species are likely to benefit from the
changes, by extending their ranges into areas previously
unoccupied; but many other taxa are expected to experi-
ence dramatic reductions in the spatial extent and connec-
tivity of areas that provide conditions suitable for their
existence.
The taxa most vulnerable to any climatically imposed
reductions in suitable habitat are likely to be those that have:
(1) highly specialized habitat requirements, (2) ‘slow’ life-
histories (delayed maturation, low reproductive rate), thereby
reducing their ability to recover from population reductions,
(3) low dispersal rates, making it difficult to colonize suitable
areas outside the existing range, and (4) are already under

DOI: 10.1111/j.1472-4642.2009.00619.x
ª 2009 Blackwell Publishing Ltd www.blackwellpublishing.com/ddi 109
T. D. Penman et al.

pressure from other anthropogenic processes (Hughes, 2003;
Meynecke, 2004; Araújo et al., 2006). One taxon that meets all
of these criteria is the broad-headed snake (BHS) (Hoploceph-
alus bungaroides), Australia’s most endangered snake species
(Cogger, 2000). This small, brightly coloured nocturnal snake
is restricted to sandstone habitats within 250 km of Sydney, in
south-eastern New South Wales (Cogger, 2000). BHS have
specific habitat requirements and during the cooler months,
they thermoregulate in crevices and under rocks in exposed
locations (Webb & Shine, 1998; Pringle et al., 2003; Croak
et al., 2008). The snakes grow slowly and mature at 5 years of
age and females produce small litters (3–7 young) infrequently
(every 2–3 years, Webb & Shine, 1998; Webb et al., 2002a,b).
Radiotelemetry (Webb & Shine, 1997) and long-term mark-
recapture studies (Webb et al., 2002a,b) show that adults
display high site fidelity and low rates of dispersal between rock
outcrops in close proximity (< 3 km). Populations of BHS
have declined dramatically throughout their geographical
range in the 200 years since European settlement because of
urbanization, removal of bush-rock for gardens and the
removal of snakes for the pet trade (Shine et al., 1998; Webb
et al., 2002a,b; Newell & Goldingay, 2005). In combination,
these factors appear likely to make the BHS susceptible to
changes in climate.
In this study, we modelled the bioclimatic envelope of this
endangered snake species under current climatic conditions to
identify climatic factors that may limit the species’ current
distribution. We also predicted the climatic envelope for the
species under the 2030 and 2070, low and high climate change
scenarios (Suppiah et al., 2007). On the basis of results of these
analyses, we determined the risk of future extinctions for this
species and identified sites within the current range that
deserve high priority for conservation.
3. High warming scenario for 2070, where temperatures
increase by 4 C and rainfall is reduced by 5% (hereafter,
2070 high scenario).
Grids representing the climate change scenario were gener-
ated within the Bioclim program using inbuilt functions.
A total of 159 independent sites for H. bungaroides were
used to predict the climatic envelope (Fig. 1). These sites were
derived from the New South Wales National Parks and
Wildlife Service Wildlife Atlas (accessed May 2008) and
supplemented by our own surveys. To ensure that our records
represented independent locations, the final dataset of 159
records comprised only of records that were more than 3 km
from all other records or (if < 3 km from another record)
occurred on a separate ridgeline, i.e. those ridges which occur
in close proximity, but are separated by deep gullies that
appear to form barriers to movement (based on Webb & Shine,
1997, 1998). All records were of extant populations or recently
extinct populations, which are known to have gone extinct
because of collection or urban development, i.e. not because of
climatic changes.
The study area for the analysis was the Sydney Basin
Bioregion (as defined by Environment Australia 2000) plus
a 100-km buffer zone around it (Fig. 1). Almost all known
records for the species lie within the Sydney Basin Bioregion
and available genetic data suggest that this low-vagility
species is unlikely to migrate > 100 km in the < 70-year
period being considered under the climate change sce-
narios (Sumner et al., 2009). Topographic data for the

METHODS
We predicted climatic envelopes for H. bungaroides for current
climatic conditions and under future climate change scenarios
using two modelling approaches – Bioclim (Nix, 1986) and
Maxent (Phillips et al., 2006). Climatic envelopes were
predicted for four separate climate scenarios. The first is based
on current climatic conditions; successful prediction of the
current range from climatic variables would suggest that
climatic conditions somehow limit the present distribution of
the species (Araújo & Pearson, 2005) and hence provide a
robust basis for predicting effects of climate change. The
remaining three scenarios were based on the predicted climatic
changes suggested by Suppiah et al. (2007) for the Sydney
region. These are:
1. Low warming scenario for 2030, where temperatures
increase by 0.5 C and rainfall is reduced by 5% (hereafter,
2030 low scenario);
2. High warming scenario for 2030, where temperatures Figure 1 Map of the 159 independent broad-headed snake
increase by 1.5 C and rainfall is reduced by 5% (hereafter, (Hoplocephalus bungaroides) sites used in our model. Grey line
2030 high scenario). This scenario is equivalent to a low represents the study area, i.e., the Sydney Basin Bioregion plus a
warming scenario for 2070; and 100 km buffer zone.

110 Diversity and Distributions, 16, 109–118, ª 2009 Blackwell Publishing Ltd
 Impact of climate change on an endangered snake

study area were extracted from the Geoscience Australia
9 second (250 m) Digital Elevation Model of Australia,
version 2.1.
Statistical analysis
The Bioclim program has been discussed extensively in the
literature (Nix, 1986; Busby, 1991; Lindenmayer et al., 1991)
and we describe it only briefly here. Bioclim derives 35
climatic surfaces from long-term meteorological data and a
digital elevation model. The climatic surfaces are categorized
broadly into those relating to temperature, radiation,
precipitation or moisture index (Table 1). A profile for the
species is developed based on known localities and elevation
at each location. Through homocline matching, Bioclim
determines the locations of climatically suitable areas, which

Table 1 Bioclimatic profile for the broad-headed snake (Hoplocephalus bungaroides), based on 35 climatic variables sampled at 159
independent snake locations.

Bioclim variable Mean SD Minimum Maximum P-value Classification

Temperature values are in C

Annual mean temperature 14.9 1.64 10.4 17.2 0.019** Restricted
Mean diurnal range [mean (period max–min)] 11 1.11 9 13.7 0.035** Lower
Isothermality 0.48 0.01 0.45 0.51 0.229
Temperature seasonality (C of V) 1.45 0.14 1.22 1.83 0.216
Max temperature of warmest period 26.1 1.35 22.3 29.7 0.173
Min temperature of coldest period 3.3 1.87 )0.2 6.6 0.048** Higher
Temperature annual range 22.7 2.13 18.9 28.1 0.033** Lower
Mean Temperature of wettest quarter 18.6 2.47 11.6 22.4 0.105
Mean temperature of driest quarter 10.6 1.97 5.6 13.8 0.003** Higher
Mean temperature of warmest quarter 20.1 1.38 15.9 22.6 0.036** Lower
Mean temperature of coldest quarter 9.5 1.93 4.7 12.4 0.17
Precipitation – values in mm
Annual precipitation 1079 170.75 655 1621 0.024** Higher
Precipitation of wettest period 32 5.64 17 48 0.158
Precipitation of driest period 11 4.97 0 16 < 0.001** Higher
Precipitation seasonality (C of V) 27 5.1 14 39 0.077* Restricted
Precipitation of wettest quarter 360 62.48 197 553 0.078* Higher
Precipitation of driest quarter 183 29.59 111 246 0.006** Higher
Precipitation of warmest quarter 334 55.51 196 500 0.086* Higher
Precipitation of coldest quarter 232 48.71 127 325 0.004** Higher
Radiation – values in MJ m)2 day)1
Annual mean radiation 16.4 0.47 15.3 17.8 0.085* Lower
Highest period radiation 24.4 0.51 22.9 25.7 0.234
Lowest period radiation 7.8 0.45 7 8.8 0.495
Radiation seasonality (C of V) 34 1.85 32 37 0.281
Radiation of wettest quarter 17.8 3.43 9.5 24.5 0.004** Lower
Radiation of driest quarter 13.3 0.67 10.2 14.5 0.001** Restricted
Radiation of warmest quarter 21.8 0.73 20 24.1 0.095* Lower
Radiation of coldest quarter 10 0.41 9.1 10.7 0.073* Lower
Moisture index – values range from 0 (dry) to 1 (saturated)
Annual mean moisture index 0.76 0.09 0.5 0.94 0.265
Highest period moisture index 0.97 0.06 0.72 1 0.347
Lowest period moisture index 0.41 0.08 0.26 0.68 0.006** Higher
Moisture index seasonality (C of V) 24 5.62 10 37 0.364
Mean moisture index of high Qtr. MI 0.96 0.07 0.66 1 0.158
Mean moisture index of low Qtr. MI 0.51 0.1 0.32 0.79 0.023** Higher
Mean moisture index of warm Qtr. MI 0.56 0.1 0.32 0.81 0.077* Higher
Mean moisture index of cold Qtr. MI 0.95 0.07 0.66 1 0.364

Shown are mean values, standard deviations and minimum and maximum values for each variable. Period refers to a week, e.g., the wettest period is
the wettest week of the year. Quarter refers to the 3-month period to the nearest week, e.g., the wettest quarter represents the wettest 13-week period
within the year. For further details regarding variable definitions, see http://fennerschool.anu.edu.au/publications/software/anuclim/doc/params.html.
Also shown is the outcome of the chi-squared analyses comparing the BHS sites and the random sample with the P-value of the analysis and the
assigned classification. C of V = coefficient of variation. *denotes a marginally significant result; **denotes a significant result at the P = 0.05 level.

Diversity and Distributions, 16, 109–118, ª 2009 Blackwell Publishing Ltd 111
T. D. Penman et al.

collectively define the climatic envelope. Terminology to

0.00 0.05 0.10 0.15 0.20 0.25 0.30 0.35

(a) Lower
define the climatic envelope varies between publications.
Here, we follow the terminology of Penman et al. (2005)

Proportion of sites
with the ‘range’ being defined as the 100th percentile of the
climatic envelope, ‘intermediate’ being defined as the 95th
percentile of the climatic envelope and ‘core range’ being
defined as the 90th percentile of the climatic envelope. The
percentiles refer to the ranges of each of the parameters. For
19 20 21 22 23 24
example, the 90th percentile or core comprises those grid Radiation of warmest quarter
cells where the values of all 35 parameters fall within

0.00 0.05 0.10 0.15 0.20 0.25 0.30

(b) Higher
the 90% range for each of the 35 climatic variables
considered.

Proportion of sites
Maximum entropy is a machine learning approach
considered to have high predictive powers for modelling
species distributions based on presence-only data (Elith
et al., 2006). Models were developed using the Maximum
Entropy Species Distribution Modelling Software, version
3.3.1 (as described in Phillips et al., 2006). By comparing the 600 800 1000 1200 1400
Annual precipitation
1600 1800 2000

presence sites with the background data (10,000 random

0.20
(c) Restricted
points), predictions for the probability of occurrence are
made for the species distribution based on environmental

0.15
Proportion of sites
layers. Models were built on 70% of the data with the
0.10
remaining 30% used as a test sample. Models were built
solely on the 35 bioclimatic parameters derived from the
0.05

Bioclim program to allow for a direct comparison of the

predictions of the two methods.
0.00

Model fit for the two methods was measured using the
area under the curve (AUC) from the receiver operating
characteristic curve (ROC). A ROC curve is the relationship
between the sensitivity and the false positive fraction
(Woodward, 1999). AUC is the area under the ROC curve,
with a value of 0.5 representing a random model, values
between 0.8 and 0.9 representing models with good fit
and those over 0.9 as being an excellent fit (Thuiller et al.,
2003).
To determine how BHS localities differed from random sites
within the broader study region, we compared the climatic
profile for the BHS to 10,000 random sites within the study
area (i.e. a background sample sensu Wintle et al., 2005).
Random sites were generated within the study area using
ArcGIS (ESRI, USA). Histograms were then prepared for each
of the Bioclim variables for both the BHS and random sites
using the R-package v 2.7.2 (R-Development Core Team
2007). We compared the resulting distributions using a chi-
squared test (Sokal & Rohlf, 1995). For this analysis, the
observed values were the counts from the BHS records and
expected values were calculated from the density function of
the random sites. Where significant (P < 0.05) or marginal
(0.05 < P < 0.10) differences were recorded, we assigned the
difference to one of three categories: lower, higher or
restricted. The lower and higher categories reflect cases where
the BHS localities averaged lower or higher for a particular
variable than did the random sample. The ‘restricted’ category
reflected a situation where the BHS locations had a similar
mean to that of the random sample, but a lower range of
variation (Fig. 2).
8 10 12 14 16 18
Annual mean temperature
Figure 2 Examples of the classification of distributional differ-
ences between the broad-headed snake (Hoplocephalus bungaro-
ides) and random sites: (a) lower, i.e., BHS distribution is
significantly lower than the background sample, (b) higher, i.e.,
BHS distribution is significantly higher than the background
sample and (c) restricted, i.e., BHS distribution has a range lower
than the background sample that creates a higher peak within the
distribution. Solid lines represent random sites and dashed lines
represent BHS locations.
RESULTS
Broad-headed snakes are presently distributed throughout
most of the predicted bioclimatic envelope (Figs 3a & 4a).
Based on the Bioclim predictions, only 2 of the 159 known
sites (1%) were not predicted as climatically suitable for the
species, with 26% of sites predicted as ‘range’ (within the 100%
percentile of the climatic envelope), 26% predicted as ‘inter-
mediate’ (within the 95% percentile) and 48% predicted as
‘core’ habitat (within the 90% percentile: see Fig. 5). Both
approaches predicted two broad areas within the species range
where BHS have not been recorded – the Cumberland Plain
and the Southern Highlands (Figs 3a & 4a). An area of
approximately 300 km2 to the west of Myall Lakes was the only
area predicted that lies outside the known range of the species.
Model fit was considered ‘good’ for the Bioclim model
(AUC = 0.89) and excellent for Maxent (AUC = 0.973
training, AUC = 0.960 testing).

112 Diversity and Distributions, 16, 109–118, ª 2009 Blackwell Publishing Ltd
 Impact of climate change on an endangered snake

(a) (b)

(c) (d)

Figure 3 Predictions of the distribution of the broad-headed snake (Hoplocephalus bungaroides) under different climate change scenarios
using Bioclim: (a) current climatic conditions, (b) a low warming scenario for 2030, (c) a high warming scenario for 2030 (equivalent to a
low warming scenario for 2070) and (d) a high warming scenario for 2070. Points represent the known locations of broad-headed snakes
(n = 159). Categories were defined after Penman et al. (2005) and are as follows: light grey areas are the ‘range’ or the 100th percentile of the
climatic envelope, mid range grey areas are the ‘intermediate’ or the 95th percentile of the climatic envelope and dark grey represents the
‘core’ or the 90th percentile of the climatic envelope. The boundaries of the study area occur beyond the boundaries of the mapped area.

Comparisons between BHS localities and the random annual temperatures (majority from 13 to 17 C) and low
sample revealed statistically significant differences in 14 of mean diurnal ranges. In the driest and coldest quarters, BHS
the 35 Bioclim variables and marginal differences in a sites experience higher temperatures than do the random
further seven (Table 1). BHS sites have a small range of sites, whereas in the wettest quarter, BHS sites experience

Diversity and Distributions, 16, 109–118, ª 2009 Blackwell Publishing Ltd 113
T. D. Penman et al.

(a) (b)

(c) (d)

Figure 4 Predictions of the distribution of the broad-headed snake (Hoplocephalus bungaroides) under different climate change scenarios
using Maxent: (a) current climatic conditions, (b) a low warming scenario for 2030, (c) a high warming scenario for 2030 (equivalent to a
low warming scenario for 2070) and (d) a high warming scenario for 2070. Points represent the known locations of broad-headed snakes
(n = 159).

lower temperatures compared with random sites. BHS sites
also have a lower annual temperature range than do random
sites. Both precipitation and moisture indices indicate
that the BHS sites are wetter, on average, than are the
random sites. Radiation values average lower at BHS sites
compared with random sites for the annual mean, wettest
quarter and warmest quarter. There is a restricted range of
solar radiation at the BHS sites in the driest quarter, with
95% of the sites experiencing between 12 and 15
MJ m)2 day)1. These variables were consistent with the

114 Diversity and Distributions, 16, 109–118, ª 2009 Blackwell Publishing Ltd
 Impact of climate change on an endangered snake

100

1.0
Not predicted
Range
Intermediate

0.8
Mean probability of occurrence
80

Core
Number of sites

0.6
60

0.4
40

0.2
20

0.0
0

Current Low 2030 High 2030 High 2070 Current Low 2030 High 2030 High 2070
Climate scenario
Figure 6 The mean probability of occupancy of known broad-
Figure 5 The proportion of known broad-headed snake headed snake (Hoplocephalus bungaroides) localities using Maxent
(Hoplocephalus bungaroides) locations (n = 159) predicted to for the four climate scenarios considered.
remain within each habitat category (range, intermediate, core, or
not predicted in the above habitats and thus outside of the climate
envelope) both under current conditions and under low and high low warming scenario (which change in temperature and
climate warming scenarios. See Fig. 2 for terminology of the rainfall by the same amount) suggest a far greater negative
classifications. impact (Figs 3c & 4c). Under this scenario, Bioclim predicts
46% of all known sites rendered climatically unsuitable for the
Table 2 The percentage contribution of the main variables in the species (Fig. 5), whereas Maxent predicts a reduction in
Maxent model. the probability of occurrence to 0.36 ± 0.02 (Fig. 6). Most of
the sites in the north-east of the species’ range and those
Variable Percent contribution immediately to the south of Sydney would be lost under this
scenario. In addition to these potential losses in range, both of
Radiation of warmest quarter 21.2 the 2030 scenarios predicted a small increase in the south-
Lowest period radiation 19.3 western extent of the climatic envelope (Figs 3c & 4c).
Radiation of coldest quarter 10.6
Under the 2070 high warming scenario, significant reduc-
Lowest period moisture index 9.5
tions in climatic suitability are predicted. Bioclim predicts that
Precipitation of wettest period 6
only 14% of known sites will remain climatically suitable for
Radiation seasonality (C of V) 5.2
Precipitation seasonality (C of V) 4.9 BHS (Fig. 5), whereas Maxent predicts a mean probability of
Annual mean radiation 3.9 occurrence of only 0.11 ± 0.01 (Fig. 6), which represents an
Annual mean temperature 3.4 83% reduction from the predictions of the current climate.
Within the known distribution, only three relatively small
All 35 bioclim variables were included in the analysis. Variables not patches continue to be climatically suitable under this scenario
presented in this table contributed < 2% to the composition of the
(Figs 3d & 4d). These patches comprise one large area in the
final model. C of V = coefficient of variation.
north-west of the range (in the Blue Mountains World
Heritage Area/Wollemi Wilderness), plus two smaller areas
variables contributing to the final model in the Maxent further south. Although both of these southern areas lie within
models (Table 2). the species’ current range, only the southernmost patch is
Predictions for each of the climate change scenarios were known to support BHS (Figs 3d & 4d).
consistent between the two modelling approaches we tested.
Changing climates are predicted to reduce the size and spatial
DISCUSSION
distribution of the climatic envelope (Figs 3–6). Bioclim
predictions for the 2030 low scenario excluded 13% of known We modelled the bioclimatic envelope of the BHS, a
sites, predominantly from the Sydney city area (Figs 3b & 4b). geographically restricted species that is threatened by habitat
Under this model, most of the species range would still be degradation (Schlesinger & Shine, 1994; Shine et al., 1998;
climatically suitable. Similarly, Maxent predicts a drop in the Pringle et al., 2009) and collection of individuals from the wild
average probability of occurrence from 0.65 ± 0.02 (SE) (Webb et al., 2002a,b). The ability of these models to predict
(current) to 0.55 ± 0.02 (SE) (low 2030) (Fig. 6). The model the current range accurately suggests that climate is an
predictions for the 2030 high warming scenario and the 2070 important determinant of the species’ distribution (Linden-

Diversity and Distributions, 16, 109–118, ª 2009 Blackwell Publishing Ltd 115
T. D. Penman et al.
mayer et al., 1991; Pearson & Dawson, 2003). Unlike many
reptiles that only inhabit a small portion of areas with
potentially suitable climates (Araújo & Pearson, 2005; Araújo
et al., 2006), BHS are distributed throughout most of their
bioclimatic envelope. However, this does not necessarily mean
that the species will be able to track future changes in climate.
Indeed, recent phylogenetic analyses have revealed that
H. bungaroides comprises two genetically divergent clades that
correspond to distinct regions north and south of Wollongong
(Sumner et al., 2009). These northern and southern popula-
tions were isolated from each other during the late Pleistocene,
approximately 755,000–850,000 years ago (Sumner et al.,
2009), when the climate in eastern Australia oscillated from
warm, wet periods during interglacials to cold dry conditions
during glacial periods (Byrne, 2008). Climate change during
the Pleistocene constrained dispersal and gene flow in
H. bungaroides over relatively short (100 km) distances, which
suggests that the species may be unable to track future changes
in climate, if similar barriers occur.
Our analyses predict that the geographical distribution of
BHS will shrink under predicted levels of climate warming.
Each of the scenarios we examined predicted significant range
contractions using both modelling approaches, rather than
expansion into new areas (e.g., Figs 3 & 4). Much of the area
that is currently suitable climatically will become too hot and
dry, and only the areas that are highest in elevation (and thus,
that are coolest) will remain. As BHS currently exist in most of
the areas that are predicted to remain suitable over longer
timescales (Figs 3 & 4), these areas are likely to support snakes
in the future. However, in the most extreme case (high
warming scenario in 2070; Figs 3d & 4d), most (87%) of the
currently known populations would be lost. Although much of
the area predicted to remain suitable is legally protected,
legislative protection may do little to eliminate anthropogenic
threats, such as illegal theft of animals (Webb et al., 2002a,b),
destruction of habitat (i.e. bush-rock removal; Schlesinger &
Shine, 1994; Shine et al., 1998) and overgrowing vegetation,
which shades critical habitat features (Pringle et al., 2009).
Thus, current levels of habitat protection in these areas may
not be sufficient and other actions (such as limiting human
access to these areas and implementing strategies to maintain
open habitats) may be required. High priority should be given
towards more effective protection of snake habitat throughout
the current range, but especially in the areas predicted to
experience suitable climate space over longer timescales.
Most research on BHS has focused on a small part of the
species’ range (e.g. Webb et al., 2002a,b; Pringle et al., 2003,
2009) and many gaps in knowledge remain for other areas.
Surveys for this species are needed, especially in areas where
the species is predicted to occur but has not been reported
(Figs 3 & 4) and in areas that are predicted to remain
climatically suitable under climate change (Figs 3 & 4). Such
surveys could evaluate whether BHS occur in these areas and
whether or not such sites contain suitable microhabitats (Shine
et al., 1998; Pringle et al., 2003; Webb et al., 2005). Artificial
habitat restoration or manipulation (Croak et al., 2009)
116 Diversity and Distributions, 16, 109–118, ª 2009 Blackwell Publishing Ltd
followed by translocation of snakes might be feasible for
conserving this species in areas where habitats are unsuitable or
where populations cannot be located. Such studies will require
careful consideration to avoid problems associated with the
introduction of diseases and the loss of the genetic integrity
within wild populations (e.g. Greer, 1996; Brown et al., 2007).
We acknowledge that predictions based on broad scale
climatic variables have limited power, particularly when the
analysis is based only on known sites of occurrence (Elith et al.,
2006) and caution must be used when interpreting the output.
Within the predicted climatic envelope, there will be local scale
factors that will further limit the distribution of the species
under future climates, primarily the presence of suitable rock
habitat (Shine et al., 1998; Webb et al., 2002a,b) or important
prey species (e.g. velvet geckos, Oedura lesueurii), which were
not considered in this study. Similarly, wildfire is thought to
play an important role in maintaining habitats for the species
(Webb et al., 2005); however, the model cannot consider the
changes in wildfire regimes that may occur under the predicted
climate change scenarios (Cary & Banks, 1999; Cary, 2002;
Bradstock et al., 2008). If these factors were considered in a
model, they would be likely to reduce further the area of
suitable habitat under predicted climate change scenarios,
rather than increase it. The congruence between the two
modelling approaches provides strong support that the broad
predicted trends in climatic suitability cannot be ignored.
Results of this study are transferable to a number of other
threatened species and endangered ecological communities,
which have similar geographical distributions to the BHS.
Examples include Pseudophyrne australis (red-crowned toad-
let), the northern populations of Heleioporus australiacus
(giant burrowing frog), Zieria involucrate (no common name),
Acacia prominens (Gosford wattle) and Acacia pubescens
(downy wattle). Although the shape and size of the climatic
envelopes will probably vary between these species, the broader
impact of climatic change is likely to be the same, resulting in a
significant reduction in the extent of climatically suitable areas.
Further research is needed to assess the impact of climate
change on such species.
In summary, our analyses suggest that broad-head snakes,
already endangered through a series of anthropogenic threats,
are likely to be vulnerable to future changes in climate. Under
rapid climate change, most of the current geographical range of
the species will no longer provide suitable climatic conditions
for viable populations within a few decades. If we are to
prevent extinction of this species, we need to initiate extensive
surveys of hitherto-unexplored regions within the species’
current distribution, and develop intensive conservation and
protection programs in the few sites predicted to remain
climatically suitable for the species in the future.
ACKNOWLEDGEMENTS
We thank researchers who provided locality records, partic-
ularly Phil Craven, Gary Daly, Mark Fitzgerald, Rachel
Melrose and Jai Thomas. We thank Meagan Ewings for

providing access to the New South Wales National Parks and
Wildlife Service locality database. Frank Lemckert, Betsy
Roznik and Alison Towerton provided encouragement, advice
and logistic support. Rod Kavanagh and Reid Tingley
provided comments on an early draft of this manuscript.
Financial support was provided by the Australian Reptile
Park, Australian Research Council, Forests New South Wales,
the Hawkesbury-Nepean Catchment Authority, New South
Wales Department of Environment and Climate Change and
Zoos Victoria.
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BIOSKETCHES
The project team is examining the conservation biology of the
broad-headed Snake specifically considering the roles of fire
and climate change.
Trent Penman is a fire ecologist with Industry and Invest-
ment NSW examining the impacts of long-term fire exclusion
on community structure and function (http://www.dpi.nsw.-
gov.au/research/staff/trent-penman).
David Pike, Jonathan Webb and Richard Shine are ecologists
at the University of Sydney with a broad range of research
interests including reptile life-history strategies and conserva-
tion management (http://www.bio.usyd.edu.au/Shinelab/).
Author contributions: T.P., D.P., J.W., and R.S. conceived the
ideas; T.P., D.P. and J.W. collated the data; T.P. analysed the
data; and T.P. and D.P. led the writing.
Editor: Chris Thomas