Australasian Plant Pathol.
(2013) 42:43–51
DOI 10.1007/s13313-012-0178-7
Taxonomic uncertainty and decision making for biosecurity:
spatial models for myrtle/guava rust
J. Elith & J. Simpson & M. Hirsch & M. A. Burgman
Received: 22 March 2012 / Accepted: 14 November 2012 / Published online: 16 December 2012
# Australasian Plant Pathology Society Inc. 2012
Abstract The causal agent of myrtle rust, initially described
as Uredo rangelii, was recorded in Australia for the first
time in 2010. Much of the monitoring effort in Australia and
elsewhere is driven by existing understanding of Puccinia
psidii sensu lato (guava rust), because U. rangelii is part of
the guava rust complex. Bioclimatic analyses for guava rust
in Australia indicate highest risk along the eastern coast,
from northern Queensland to the south coast of NSW. These
analyses rely on native and invaded range records for
Puccinia psidii sensu lato. However, models that are instead
fitted to records representing U. rangelii emphasise different
risk areas, with less focus on northern coastal Queensland
and new predictions into NSW tablelands and north-eastern
Victoria. These differences have important implications for
biosecurity containment and monitoring efforts. Here we
use this example as a case study to explore the implications
of taxonomic uncertainty for predictions of the potential
distribution of a species of biosecurity concern in
Australia. The important message is that the different pat-
terns implied by different taxonomic assumptions may have
very different management implications and taxonomic un-
certainty should be evaluated alongside other sources of
Electronic supplementary material The online version of this article
(doi:10.1007/s13313-012-0178-7) contains supplementary material,
which is available to authorized users.
J. Elith (*) : M. A. Burgman
Australian Centre of Excellence for Risk Analysis,
School of Botany, University of Melbourne,
Parkville 3010, Australia
e-mail: j.elith@unimelb.edu.au
J. Simpson
15 Sprent St,
Narrabundah 2604, Australia
M. Hirsch
CSIRO, Canberra, Australia
uncertainty. We could find no published example where
taxonomic uncertainty was evaluated in biosecurity plan-
ning, despite the fact that taxonomic uncertainties are com-
mon. We outline how to model such uncertainties and
discuss methods for exploring them and their impacts on
predicted distributions.
Keywords Biosecurity . Eucalyptus rust . Guava rust .
Species distribution models
Introduction
The rust species Puccinia psidii (Guava rust) was described
from Psidium guajava, or guava, in Brazil in 1884 and
reported on non-native eucalypts in the same region by
Joffily (1944). It was restricted to South and Central
America and the Caribbean until its appearance in Florida in
1977 (Marlatt and Kimbrough 1979). Apart from two unusual
reports in Taiwan and South Africa (detailed in Glen et al.
2007) and unconfirmed reports from Indonesia of a Puccinia
species on eucalypts (Hardiyanto and Tridasa 2000), there was
no evidence of spread to other countries until it was discov-
ered in Hawaii in 2005 (Killgore and Heu 2007). Once there, it
spread quickly (Loope and La Rosa 2008).
Puccinia psidii has a large and varied host range
(Carnegie and Lidbetter 2012; Morin et al. 2012) and some-
times has substantial impacts on infected species (Glen et al.
2007; Carnegie and Cooper 2011; Morin et al. 2012). Guava
rust has long been viewed as a biosecurity threat to Australia
(Grgurinovic et al. 2006; Simpson et al. 2006; Glen et al.
2007; Old 2007) because of its potential impacts on a broad
range on Myrtaceae, and the diversity and importance of
these species in Australian ecosystems. The rust has been
considered a significant risk for entry and establishment in
Australia because of its rapid spread in Hawaii, and because,
44
like other Puccinia species that have established and spread
(Wellings et al. 1987), it has adherent urediniospores that
may attach to the clothing of travellers, traded commodities
or containers (Glen et al. 2007). Most spatial models for the
species’ potential distribution in Australia emphasised hab-
itat centred in coastal Queensland, extending from Cape
York to coastal New South Wales, near Sydney (Booth et
al. 2000; Magarey et al. 2007; Glen et al. 2007; Booth and
Jovanovic 2012).
There are a number of synonyms for P. psidii and its
anamorph Uredo psidii (Simpson et al. 2006), and some
complexity in the taxonomy of related anamorphs. Simpson
et al. (2006) distinguished U. rangelii from P. psidii and its
anamorph, U. psidii. The specific status of U. rangelii is based
on the presence of a tonsure and differences in size, shape and
wall thickness of the urediniospores (Simpson et al. 2006).
Langrell et al. (2008) described a molecular diagnostic tech-
nique for P. psidii and Zhong et al. (2008) developed micro-
satellite markers. Molecular phylogenetic analyses of U.
rangelii and other taxa in the P. psidii complex have provided
a different viewpoint to the morphometric analyses, placing U.
rangelii within the P. psidii complex (Carnegie et al. 2010a).
Uredo rangelii was recorded in Australia for the first time in
2010 (Carnegie et al. 2010b). Teliospores of the introduced
rust have been reported, prompting questions about the appro-
priateness of the rust’s placement in the genus Uredo
(Carnegie and Cooper 2011). Indeed, Carnegie and Cooper
(2011) recommend that the common name myrtle rust be
discontinued for the introduced rust in Australia, and it be
recognized as a member of the eucalypt/guava rust complex
instead. We will refer to it throughout this paper as myrtle/
guava rust primarily as a reminder of the taxonomic uncer-
tainty for which this system provides a useful case study.
Taxonomic disagreements based on interpretations of
morphometric and genetic data are commonplace. Both
genetic and morphological approaches may fail to correctly
discriminate species, or may separate entities that turn out
not to be justified (Schlick-Steiner et al. 2010). Species
delimitations are essentially theoretical constructs that re-
flect ecology, morphology or geneaology (Bock 1992;
Mallet 2007), the interpretations of which are subjective
(Schlick-Steiner et al. 2010). Even for sexually reproducing
species, reproductive isolation is an appropriate definition of
species boundaries for only a subset of recognised taxonom-
ic entities. Many species of the genus Eucalyptus, for in-
stance, readily form fertile hybrids. Species definitions are
subject to change, revision and amalgamation as new evi-
dence accumulates. This form of uncertainty is essentially
linguistic, reflecting the indeterminacy of theoretical con-
cepts in science generally (Regan et al. 2002).
Ideally, taxonomic uncertainty is reconciled by the accu-
mulation of evidence, the revision of theories about species
and phylogenies and the synthesis of evidence from genetic,
J. Elith et al.
morphological and ecological sources (Schlick-Steiner et al.
2010). However, it can have urgent practical importance, as
exemplified in the case of the recent incursion by myrtle/
guava rust into Australia (Carnegie et al. 2010b). Spatial
predictions for the species guide the deployment of resour-
ces and the development of containment and surveillance
strategies. Most of these rely, at least in part, on known
locations of the species in its native range (Venette et al.
2010). We use this incursion as a case study, and propose
alternative scenarios that illustrate how scientists might deal
with taxonomic uncertainty when managing invasive taxa. If
the circumscription of U. rangelii is correct in the sense that
it is informative about the pathogens’ ecology and habitat,
then the data used to predict the species distribution in
Australia should be a subset of the data for the P. psidii
complex. Alternatively, if U. rangelii does not have unique
genetic composition and ecological behaviour, then the full
data set for P. psidii is a more appropriate source for a
model. Since most data on the P. psidii complex does not
differentiate between component species, and because infor-
mation on the specific impacts of U. rangelii are unavail-
able, to date most monitoring and surveillance advice has
relied on knowledge about P. psidii sensu lato, the broad
circumscription that subsumes U. rangelii.
In this example of decision making under uncertainty,
managers need to decide whether to assume P. psidii sensu
lato is an appropriate taxonomy, or to discriminate the taxa
and assume U. rangelii has entered Australia. There are a
number of ways of solving this problem. Decision makers
may take the most likely, or the most broadly supported
option. Alternatively, they may select the option that max-
imises expected return. This can be found by combining the
probabilities of each of several possible states of the world
with the outcomes that would result if the world turns out to
be in that state (e.g., Costello and Polasky 2008). If decision
makers choose either of these options, they expose them-
selves to failure if the alternative turns out to be true. A third
decision option is to make decisions that result in satisfac-
tory outcomes, no matter which state of the world turns out
to be true. Simon et al. (1958) and Simon (1991) defined a
robust strategy as a series of decisions that result in out-
comes that are good enough, rather than optimal, but give a
satisfactory solution, no matter which possible state of the
world is true.
The purpose of this study is to explore the implications of
taxonomic uncertainty for the management of a new inva-
sive pathogen. We use the Australian incursion of myrtle/
guava rust as an example, not to argue which taxonomic
interpretation, data set or model is correct, but to highlight
the impact of taxonomic belief on modelled predictions. We
present spatial models based on bioclimatic variables that
distinguish between the species concepts from a number of
perspectives. We suggest strategies that can deal with
Taxonomic uncertainty and decision making for biosecurity
uncertainty, and discuss approaches that are robust to this
uncertainty.
Methods
Models used to predict the potential distribution of an inva-
sive species in a new region usually rely—at least in part—
on records of that species in its native (and sometimes its
invaded) range. For instance, the suite of methods termed
species distribution models (SDMs), bioclimatic models or
ecological niche models (Elith and Leathwick 2009) use
some type of statistical model to estimate the relationship
between records of the species and the environments in
which it has been recorded. These models can then be used
to predict distributions across continuous geographic
regions, including areas in which the species is currently
unknown. SDMs have been used to predict the potential
distribution of guava rust (e.g. Booth et al. 2000). Related
models that rely on a mixture of expert knowledge, labora-
tory experiments and species locations (e.g. NAPPFAST,
CLIMEX, and expert models tailored to available knowl-
edge) have also been used for this species (Magarey et al.
2007; Kriticos and Leriche 2008; Booth and Jovanovic
2012). Whilst there is ongoing debate about the most ap-
propriate modelling methods for invasive species (Venette et
al. 2010), here we use only one method and focus instead on
the question of how taxonomic uncertainty can impact pre-
dictions, and how decision makers may seek robust alter-
natives. This approach will apply in principle to all methods
for species distribution modelling.
Predictor variables
Urediniospore germination, infection and disease progres-
sion are most likely to occur under conditions when tem-
perature is about 15–25 °C, humidity and leaf wetness are
high and light intensity is low, and these conditions persist
for several hours (Ruiz et al. 1989; Glen et al. 2007; Zauza
et al. 2010). Our approach to selecting the candidate varia-
bles involved finding relevant available variables with glob-
al coverage (more precisely, those with data in all relevant
regions), and choosing a subset consistent with ecological
knowledge for this taxon. We then tested pairwise correla-
tions between the variables on random samples of locations
in the regions on which the models were trained (see next
section). Based on our experience using bioclimatic varia-
bles for modeling, we preferred to avoid those variables
involving combinations of temperature and precipitation
because the way they are constructed—relying on the iden-
tification of say a warmest month or quarter—can make it
difficult to interpret results when predicting to new times
and places. However we did retain one such variable in the
45
candidate set (mean temperature of the wettest quarter)
because of its likely ecological relevance. This process led
to the selection of seven variables with pairwise Pearson’s
correlations less than 0.8 in the model training region shown
in Fig. 1 (Table 2, Online Appendix 2). All variables were
unprojected (i.e., latitude and longitude in degrees, for in-
stance WorldClim data—http://www.worldclim.org/), with
grid cell resolution of 2.5 min (approximately 5 km by
5 km).
Species data
Our presence records were gathered from vouchered speci-
mens and records of colleagues (detailed in Online
Appendix 1). For most records, features that might separate
U. rangelii from Puccinia records were not documented, so it
is not possible to distinguish U. rangelii from P. psidii unam-
biguously. We created five datasets that reflect five different,
plausible interpretations of the distribution data (Table 1). In
doing so, we recognise that the basis for these groupings could
be debated and that future evidence might prove them wrong.
Our aim here is not to argue that the specific choices are
indisputable, but to create five distinct datasets with reasoning
behind each, and to use these to model and predict. Our aim is
to use these data as a test case to explore the effect of uncer-
tainty on modelled predictions. The first dataset, Uredo_10,
uses only records in which the discriminating tonsure has been
identified (Simpson et al. 2006; Pérez, pers.comm. 2010). The
second dataset, Uredo_27, assumes an additional 17 records
(none of which have been examined for the presence/absence
of a tonsure) from the same general geographic region and
similar environments are in fact Uredo. The “similar environ-
ments” were identified by sorting all records on all available
climatic variables (one at a time) and finding that variable for
which the Uredo_10 records were least spread through the
sorted list. Mean temperature of the wettest quarter (tmean_-
wetq, Table 2) achieved this and displayed a range of 13.9 °C
to 23.5 °C across the Uredo_10 records. The additional 17
records were within this range, and from Brazil (14),
Argentina (2) and Uruguay (1). It is plausible that the spread
south into Uruguay and the cooler areas of Argentina origi-
nated from these areas of Brazil, and in relatively recent times.
The third dataset, Puccinia_94 assumes all verified records
are from a single taxonomic entity, Puccinia psidii. The
remaining two datasets, Puccina_84 and Puccinia_67, make
more restrictive assumptions about the identity of records
accorded to Puccinia; these are simply Puccunia_94 without
Uredo_10 or Uredo_27 records. Therefore within this setup,
the total of 94 records are regarded as: all Puccinia
(Puccinia_94), divided on known attributes (Uredo_10 vs
Puccinia_84), or divided on informed guesses of attributes
(Uredo_27 vs Puccinia_67). This arrangement of the records
allows us to construct a series of predictive maps, each of
46 J. Elith et al.

Fig. 1 Location of records for

the species complex used in
these analyses. The main figure
shows locations for
Puccinia_84 excepting one in
Taiwan (Table 1). The inset
panel shows all records for
Uredo_27 as white circles with
black outlines; solid black
circles are superimposed on
these for the 10 tonsured
Uredo_10 records

which accords with a different interpretation of evidence,
thereby displaying the implications of taxonomic uncertainty
for the management of the invasion in Australia.
Modelling
We used MaxEnt (Phillips et al. 2006; Elith et al. 2011)
because it is appropriate for presence-only records such as
these, it is widely used for modelling, and has settings that
can be adjusted according to specific modelling problems
and data needs. For invasive species, SDM methods can fail
if the climates in the area of interest (here, Australia) are
substantially different to those in the native and current
invaded ranges. We first ascertained, using the multivariate
environmental similarity surfaces (MESS maps) in MaxEnt
(Elith et al. 2010), that most of Australia is climatically
similar to the countries from which we had data (Online
Appendix 4). Hence, we have evidence that the models do
not need to extrapolate in our regions of interest, which
enhances confidence in their predictions.
Since MaxEnt is well described elsewhere, it is sufficient
here to say that this method uses information from the
presence records, a landscape (“background”) selected by
the user, and environmental covariates, and fits a model that
minimises the relative entropy between the probability den-
sity estimated for the presence records and that for the

Table 1 Five interpretations of presence records for U. rangelii and P. psidii

Database label Number of records Details

Uredo_10 10 The 10 records in which there is confirmation of a tonsure, the diagnostic characteristic of
U. rangelii—records are mostly restricted to Uruguay, with one record from a persistent
(well established) invasive population in Jamaica (Fig. 1).
Uredo_27 27 The records in Uredo_10 plus 17 records currently attributed to Puccinia from within the
same general geographic area as the Uredo records, and in similar environments (inset panel).
The environments were assessed by sorting all Uredo and Puccinia records on each of the
available predictor variables (see below), then selecting that variable (mean temperature of
the wettest quarter) for which the Uredo_10 records were most “clumped”—i.e. for which
they spanned the smallest proportion of that variable’s range. Any Brazilian, Uruguayan or
Argentinean records within the range spanned by Uredo were then included in the
Uredo_27 dataset.
Puccinia_94 94 All 94 Puccinia and Uredo records, in both native and invaded ranges. All but the Taiwanese
invasive Puccinia record are shown in Fig. 1 (across the map and the inset), and the locations
are tabulated in Online Appendix 1. This grouping is loosely referred to as Puccinia psidii sensu lato.
Puccinia_84 84 The data for Puccinia_94 omitting records in Uredo_10.
Puccinia_67 67 The data for Puccinia_94 omitting records in Uredo_27.
Taxonomic uncertainty and decision making for biosecurity
Table 2 Candidate predictor variables for the MaxEnt models
Name Explanation
tmax_max Maximum temperature of the warmest month
tmin_min Minimum temperature of the coldest month
tmean_wetq Mean temperature of the wettest quarter
prec_max Precipitation of the wettest month
prec_min Precipitation of the driest month
humidity Mean annual relative humidity. Data originally
at 0.5 °—resampled to match resolution of others
aridity Aridity index0mean annual precipitation/mean annual potential
evapotranspiration. Data originally at 30 arcsec and resampled to 2.5 min
landscape (Elith et al. 2011). The model can be made more
or less complex by choice of feature types (similar to data
transformations used in regression models—e.g. for model-
ling quadratic responses), and by changing the regularisa-
tion (smoothing) parameters for the model. The results
presented here are based on the following selections: (i) species
records from native and invaded ranges (Table 1) to give the
best possible representation of suitable environments for the
species, (ii) background from the countries or states in which
the species was recorded (Fig. 1); 10,000 samples placed
randomly but proportional to cell area (Elith et al. 2011); (iii)
only linear and quadratic features, because the smallest data set
(10 records) would not support more complexity. These deci-
sions are based on experience using MaxEnt and will affect the
outcome, so Online Appendix 6 explores the effect of the
decisions about feature classes and candidate variables.
Results
We present and discuss predictions for Australasia and its
northern neighbours (Fig. 2), but include world-wide results
in online appendices. The most restricted data set for U.
rangelii, Uredo_10, displays the model for those specimens
for which a diagnostic tonsure is recorded. It indicates a
preferred habitat centered on NSW and extending into
southern Queensland, and with significant potential habitat
in the north island of New Zealand.
Uredo_27 includes records from similar locations and envi-
ronments as Uredo_10, but generates a model for the rust
species’ habitat that extends further north in Australia and
expands high predictions in the highlands of Papua New
Guinea and Indonesia. With the selected settings, the only
models predicting high values in the south of Western
Australia are Uredo_27 and, to a lesser extent, Puccinia_94,
though that result varies with other choices of candidate
variables and feature classes (Online Appendix 6). Models
for Puccinia (Puccinia 94, 84 and 67) are roughly consistent
47
Source
Worldclim (http://www.worldclim.org/)
CRU, East Anglia, UK
(http://www.cru.uea.ac.uk/cru/data/hrg/)
CGIAR (http://www.cgiar-csi.org/)
with one another. They indicate reduced suitability of the more
southerly regions in southern mainland Australia, Tasmania
and New Zealand but high suitability in parts of Indonesia,
New Guinea and New Caledonia.
The differences in these mapped predictions are driven
by differences in the variables selected as important in
determining the distributions, and the shapes of the fitted
functions (Table 3 and Online Appendix 5). Whilst the
results should be interpreted carefully for those datasets with
very few presence records (Uredo_10 and Uredo_27) and
because some predictors are correlated with others (Online
Appendix 2), a general pattern is that minimum temper-
atures and minimum precipitation are most consistently
chosen as important predictors. As the species data move
away from the Uredo or Uredo-like records (i.e.
Puccinia_67) maximum temperatures and mean tempera-
tures of the wettest quarter become more prominent.
Figure 3 shows the modelled responses for relationships
when only one variable, minimum monthly temperature, is in
the model. It clearly distinguishes different responses derived
from the Uredo_10 and the three Puccinia data sets drawn
from significantly warmer temperatures. In situ experiments
may provide the information to evaluate whether this difference
is a sampling effect, or is a real ecological phenomenon.
The models in Fig. 2 were created from records from
outside of Australia. The incursion provides an opportunity
(albeit so far, a limited one) to evaluate the model predic-
tions. Details of two predictions are provided in Fig. 4,
together with the locations of all records from within
Australia up until April 2011. We note that many of these
are from artificial environments such as nurseries and home
gardens. Since then it has extended as far north as
Rockhampton, Queensland, and in 2012 has been found in
Victoria, mostly around Port Phillip Bay (DPI 2012). Both
models provide reasonable predictions for the invasive path-
ogen, at least up until the point at which records were
available for these analyses. The limitation to this evaluation
is that, without knowledge of how the nurseries and
48 J. Elith et al.

Fig. 2 MaxEnt predictions for Uredo_10 Puccinia_94

the habitat of the Uredo/
Puccinia species complex in
Australia under five different
assumptions concerning the
observation data. The datasets
are detailed in Table 1

Uredo_27 Puccinia_84

Puccinia_67

Legend:

gardeners modify their environments, it is difficult to assess Discussion

the true environments in those locations. The anomalies in
Fig. 4 (the two more inland records) are both records from
nurseries to which the pathogen was transported by road, in
consignments of infected host plants. The most robust eval-
uation will be from established invasions in unmanaged
vegetation; in fact, a complete evaluation is not possible
until the species has spread to all suitable environments in
Australia (Elith et al. 2010).
The potential consequences of this disease for the flora of
Australia are substantial. It will have significant impacts on
new growth of a very large number of indigenous species,
including many rare, endangered and geographically re-
stricted taxa (Carnegie and Lidbetter 2012). All the spatial
models predict extensive potential habitat, no matter which
interpretation of the data is applied. Repeated damage to

Table 3 Percent contribution (permutation importance) of each of the described in its accompanying tutorial. The two most important vari-
explanatory variables (rows) for each of the five models (columns; see ables in each model are in bold type, as identified by a relatively high
text for details). Values calculated within the MaxEnt program, as value for both measures

Uredo_10 Uredo_27 Puccinia_94 Puccinia_84 Puccinia_67

prec_min 50 (80) 11 (1) 25 (13) 22 (11) 17 (5)

prec_max 0 (0) 1 (0) 7 (24) 8 (19) 11 (18)
tmin_min 38 (19) 70 (82) 20 (14) 21 (16) 14 (13)
tmax_max 6 (1) 1 (1) 11 (18) 12 (32) 13 (26)
tmean_wetq 0 (0) 7 (2) 8 (8) 8 (11) 14 (30)
aridity 5 (0) 5 (12) 6 (23) 3 (13) 7 (8)
humidity 1 (0) 7 (2) 24 (1) 26 (0) 25 (0)
Taxonomic uncertainty and decision making for biosecurity
Fig. 3 Modelled responses to minimum monthly temperature for the
five data sets
flushes of new growth over several growing periods may
kill seedlings, saplings and trees, or greatly reduce crop
yield, as has been observed in Brazil, Florida and Hawaii
(MacLachlan 1938; Coutinho et al. 1998; Rayachhetry et al.
2001; Loope 2010; Martins et al. 2011). While plantation
managers in some of these areas have been able to breed
disease resistant varieties (Junghans et al. 2003; Alfenas et
al. 2004), the resource demands to develop resistant varie-
ties of all affected Australian natives would be overwhelm-
ing. Considerable loss of biodiversity seems possible; rare,
susceptible, geographically restricted members of the
Myrtaceae seem to be particularly at risk.
a Uredo_27 b Puccinia_94
Rockhampton
Port Phillip Bay
Fig. 4 Details of MaxEnt predictions for eastern Australia, from models
based on 2 different datasets (Uredo_27, left, and Puccinia_94, right).
Black dots are observations from the incursion in 2010 and 2011; legend
as for Fig. 2. The more inland observations outside of predicted high risk
areas are both from nurseries
49
While the predictions of the MaxEnt model are consistent
with the incursion data to date, the predictions in Figs. 2 and
4 do not encompass the full range of uncertainty. Other
sources of uncertainty could be considered and decisions
could be made robust to them. Model predictions are likely
to be affected by the choice of candidate variables, the
selection of feature types, and the constraints imposed on
model functions. Maps produced by models that make al-
ternative assumptions are shown in Online Appendix 6. This
sort of uncertainty is not unique to MaxEnt—all model
fitting requires decisions about data and settings which
affect the end product; few modelling exercises make these
uncertainties explicit (but see, for instance, Lawson et al.
2010 for an example of an uncertainty analysis with another
modelling package, CLIMEX).
The qualitative predictions of the two taxonomic theories
and the contingent interpretation of the data are starkly
different. Recognition of P. psidii sensu lato (Puccinia_94)
would lead managers to place lower priority on surveillance
and containment in Western Australia, and to increase the
focus of activities in Australia’s northern and eastern neigh-
bours (e.g. New Caledonia). Recognition of U. rangelii (e.g.
Uredo_27) would lead managers to increase the priority for
these activities in New Zealand, Tasmania and Western
Australia (Fig. 2). Both theories are consistent with the data
on the distribution of the invasion documented to date
(Fig. 4).
The prediction from the most restricted dataset, Uredo_10, is
the least consistent with the reported locations, because it fails to
predict high risk areas in Queensland. There are several possible
interpretations of this result. First, the narrow circumscription to
those records with known tonsures may have restricted the
modelling dataset too severely, to only a subset of the true
locations for the species. Second, the small number of records
(10) substantially limits what the model is able to distinguish, so
the idea might be correct, but the analysis too underpowered to
form a useful prediction. Third, our evaluation dataset may be
only of limited value or even misleading because it contains so
many records in artificial environments (nurseries, gardens etc.).
Records in natural environments are needed to validate the
alternative theories (though they are not desirable).
It may be that the disease behaves differently in Australia
than in other places it has been detected, because of the
presence or absence of competitors, unique combinations of
climatic or microclimatic conditions, and the large suite of
native Myrtaceae. Taxonomy is a rough guide to ecological
behaviour, at best. However, given the spatial implications
of taxonomic uncertainty for this taxon, a robust decision
(sensu Simon 1991) should consider that either interpreta-
tion is possible. This leads to strategies for containment and
surveillance that focus on far northern Australia and Papua
New Guinea, as well as on Victoria, Tasmania, Western
Australia, New Zealand and New Caledonia.
50
In the interim, it would be worthwhile to devote signifi-
cant resources to resolving the taxonomic uncertainty. This
should involve compiling and synthesising genetic, morpho-
metric and ecological information (Schlick-Steiner et al.
2010). The taxonomic effort should be supplemented by
controlled environmental experiments, and studies that im-
prove the extent and reliability of presence records. The
latter may have several elements, including evaluating the
provenance of all existing records, and adding to the data
base by acquiring new records of the disease in its native
range and in newly invaded areas.
Our modelling is an example of situations where there is
taxonomic uncertainty (or related uncertainties such as dif-
ferent strains and pathogenicities of organisms originating
from different regions), and demonstrates how managers
may deal with the effect of such uncertainty on predictions.
While it is self-evident that different sets of species records
will produce different predictions, the important result is
that managers need to be mindful of exactly how and where
predictions will change as datasets change, because of the
interplay between geographic locations and environmental
conditions. How these may play out in a region of interest is
quite complex (Colwell and Rangel 2009). For instance, in
our case study the predictions in south-eastern Australia
from the Puccinia_94 model are more focussed (cover less
area) than the predictions from the Uredo_10 model, yet the
Puccinia_94 data includes those in Uredo_10. This is related
to the weight of evidence—with more data, and more exam-
ples across a range of environments, the model has more
capacity to identify optimal conditions for the “species” of
interest. The results also reflect the vagueness inherent in
small samples (Uredo_10). The exact nature of such mod-
elled relationships is not easy to guess beforehand, and we
believe that the best way to explore uncertainty is to actually
test it with alternative interpretations of real or simulated
data. If real datasets were not available, simulated ones
could be constructed based on expert opinion about the main
uncertainties involved. Tools for understanding predictions
are also useful: for instance, MaxEnt has a technique—the
“explain” tool—to explore exactly what it is about the con-
ditions at any given site that lead to a high or low prediction
(Elith et al. 2010). We note that, in a related body of
literature, ecologists have suggested that SDMs be fit to
sub-taxon level data when predicting effects of climate
change on species (or sub-species) (Pearman et al. 2010).
This is consistent with the idea and motivation behind our
study outlined here, that subsets of data might be both
ecologically realistic and illuminating.
Once the impact of uncertainties is quantified, some deci-
sion needs to be made regarding the most appropriate response.
Whilst it is often tempting to try to reduce uncertainties by
choosing one option, or to continue discussions until some
consensus is achieved, the reality is that uncertainty is real, and
J. Elith et al.
consensus does not always provide a result closer to the truth.
When the range of predictions that emanate from alternative
theories are large enough to alter decisions, managers are
confronted by a decision under uncertainty. We have illustrated
here one general and effective approach—termed robust deci-
sion making—which finds satisfactory rather than optimal
solutions for the alternative, plausible models. In this way, a
decision maker provides outcomes that people can live with,
no matter which interpretation of the data turns out to be
correct. The monitoring strategy that includes both Western
Australia and the tropical north illustrates such a robust alter-
native, and could be continued until the taxonomic uncertainty
were reconciled by the results of field experiments that exam-
ined the ecological behaviour of the different morphotypes.
Climate-based modelling systems are commonly used to
predict potential range limits for pest species (Venette et al.
2010). Our results here show a significant effect of taxo-
nomic concepts on spatial models with implications for their
use in predicting potential distribution and geographic
range. They show the importance of the origin of the inoc-
ulum for the likelihood of its establishment and spread in a
new environment in which there are native potential hosts.
Acknowledgements Jane Elith was supported by Australian Re-
search Council grant FT0991640. Many people contributed to the
location data for the species records—we thank them for this important
resource and list sources in Online Appendix 1. The manuscript was
substantially improved by helpful comments and suggestions from two
anonymous reviewers and the editor, Dr Angus Carnegie.
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