Ecography 39: 001–009, 2016

doi: 10.1111/ecog.01961
 2015 The Authors. Ecography  2015 Nordic Society Oikos
Subject Editor: Brody Sandel. Editor-in-Chief: Miguel Araújo. Accepted 7 December 2015

NicheA: creating virtual species and ecological niches in

multivariate environmental scenarios

Huijie Qiao, A. Townsend Peterson, Lindsay P. Campbell, Jorge Soberón, Liqiang Ji

and Luis E. Escobar
H. Qiao and L. Ji, Key Laboratory of Animal Ecology and Conservation Biology, Inst. of Zoology, Chinese Academy of Sciences, Beijing, 100101
China. – A. T. Peterson (http://orcid.org/0000-0003-0243-2379), L. P. Campbell and J. Soberón, Biodiversity Inst., Univ. of Kansas, Lawrence,
KS 66045, USA. – L. E. Escobar (lescobar@umn.edu), Minnesota Aquatic Invasive Species Research Center and Dept of Veterinary Population
Medicine, Univ. of Minnesota, St Paul, MN 55108, USA.

Robust methods by which to generate virtual species are needed urgently in the emerging field of distributional ecology
to evaluate performance of techniques for modeling ecological niches and species distributions and to generate new
questions in biogeography. Virtual species provide the opportunity to test hypotheses and methods based on known and
unbiased distributions. We present Niche Analyst (NicheA), a toolkit developed to generate virtual species following the
Hutchinsonian approach of an n-multidimensional space occupied by the species. Ecological niche models are generated,
analyzed, and visualized in an environmental space, and then projected to the geographic space in the form of continuous
or binary species distribution models. NicheA is implemented in a stable and user-friendly Java platform. The software,
online manual, and user support are freely available at < http://nichea.sourceforge.net >.

The ecological niche has been a central concept in mod-
ern ecology (Hutchinson 1957, Leibold 1995, Holt 2009).
G. Evelyn Hutchinson presented a formalization of niche
concepts more than 50 yr ago (Hutchinson 1978), propos-
ing relationships between ecological niches and geographic
spaces (which he called ‘biotopes’; Soberón and Nakamura
2009); this linkage is known as the Hutchinsonian dual-
ity (Pulliam 2000, Colwell and Rangel 2009). However,
tools for visualizing, exploring, and analyzing distributions
of species in these linked spaces have remained limited
(but see Duan et al. 2015 and Leroy et al. 2015). Recent
years have seen massive increases in availability of data on
occurrences of species and important environmental dimen-
sions, and researchers have developed correlational algo-
rithms by which to estimate ecological niches and explore
potential distributional areas (Peterson et al. 2011). These
correlational models have been referred to as species distri-
bution models (Austin 2007, Pearson et al. 2007, Pearson
2010), habitat models (Hirzel and Arlettaz 2003, Guisan
and Thuiller 2005), or ecological niche models (Soberón
and Peterson 2005, Peterson 2006). Soberón and Peterson
(2005) proposed the biological-abiotic-mobility (BAM)
scheme, a heuristic framework linking niche concepts and
distributional areas. BAM highlights three important causal
factors of species’ distributions: the geographic distribution
of suitable abiotic environmental conditions, the geographic
distribution of suitable biotic conditions, and the potential
to reach areas by dispersal in relevant time periods.
Basic ideas for measurement of niche characteristics
were first proposed more than four decades ago (Colwell
and Futuyma 1971). Initial correlational niche modeling
approaches were outlined beginning in the 1970s (Green
1971, Hutchinson 1978), with enormous interest and
advancement in the 1990s and 2000s (Peterson et al. 2011).
More recently, techniques and toolkits have been developed
to make these approaches more tractable (Abrams 1980,
Rödder and Engler 2011); in particular, these approaches
have been integrated in tools for model calibration (Phillips
and Dudík, 2008) and for comparative studies of eco-
logical niche models in ENMTools (Warren et al. 2008,
2010, Glor and Warren 2011) and NichePy (Bentlage and
Shcheglovitova 2012) software packages.
The interplay of different factors in geographic (G) and
environmental (E) spaces can be explored via so-called ‘virtual
species’. Virtual species are made by software in the form of
simulated data that resemble real species, the researcher char-
acterizes a priori a limited and known set of causal factors to
create a ‘species’ of known niche characteristics. Niches and
distributions of virtual species can then be explored using
modeling techniques; numerous recent studies used virtual
species to explore complicated questions in distributional
ecology (Hirzel et al. 2001, Elith and Graham 2009, Godsoe
2010, Rödder and Engler 2011, Barbet-Massin et al. 2012,
Saupe et al. 2012, Owens et al. 2013, Meynard et al. 2013,
Miller 2014, Moudrý 2015). Virtual species allow research-
ers to avoid problems such as biases related to choice of

Early View (EV): 1-EV

species (Barbet-Massin et al. 2012), biases and error in real-
world data sets (Saupe et al. 2012), and unknown true niche
dimensions (Broennimann et al. 2012).
Two software packages, SDMvspecies and virtualspecies
(Duan et al. 2015 and Leroy et al. 2015), were presented
recently to permit researchers to design virtual species by
several methods. SDMvspecies software can create virtual
species based on ideas from four previous reports: niche
syntheses (Hirzel et al. 2001), pick mean method (Jiménez-
Valverde and Lobo 2007), pick median method (Lobo and
Tognelli 2011), and artificial bell-shaped response method
(Varela et al. 2014). In virtualspecies, users can generate
virtual species by two approaches – defining response func-
tions for each environmental variable, and defining suitabil-
ity from a principal components analysis (PCA; Leroy et al.
2015). Both, virtualspecies and SDMvspecies, were written
as R packages (R Development Core Team).
In this paper, we introduced a platform from which to
explore Hutchinson’s duality in the context of the BAM
framework and other niche-related concepts, called Niche
Analyst (NicheA). This freely available program allows users
to create virtual environmental spaces and species, and to
analyze ecological niches in both multivariate environmental
and geographic spaces, effectively linking views of niche and
distribution.
Description
NicheA is a cross-platform application released under a
GNU Public License (GPL). NicheA is written in Java,
combining several other libraries from R (R Development
Core Team), Weka (Mark Hall et al. 2009), JAMA (Hicklin
et al. 2012), GDAL (GDAL Development Team 2011),
and QuickHull3D (Lloyd 2012; Supplementary material
Appendix 1). The platform is a window-based, user-friendly
application that executes on most common operating sys-
tems (Fig. 1), including Microsoft Windows, Mac OS X,
and some Linux releases, such as Ubuntu. The electronic
user manual, which covers all of the functions implemented
in NicheA, installation and user support, can be found at
< http://nichea.sourceforge.net/ >.
Because biotic interactions introduce enormous complex-
ity, and relevant data are generally lacking (Soberón 2007,
Colwell and Rangel 2009, Holt 2009), NicheA focuses on
non-interactive (scenopoetic) variables in Grinnellian niche
environments (Soberón 2007), thus effectively ignoring
biotic interactions. Even when ecological niches are related
to the environmental space (E), the current literature focuses
on management, evaluation, and interpretation of ecologi-
cal niches in geographic space (G). To mitigate biases in
G, NicheA allows generating and displaying species’ niches
in E, and visualization of species’ distributions in linked E
and G spaces. Functions and action flows in NicheA are
illustrated in Fig. 1.
NicheA assumes that species’ fundamental ecological
niches are convex in shape, as available evidence suggests
(Birch 1953, Maguire 1967, Hooper et al. 2008, Angilletta
2009, Soberón and Nakamura 2009). Niches can thus be
operationalized as minimum-volume ellipsoids (Van Aelst
and Rousseeuw 2009) or convex polyhedrons (Rissler and
Apodaca 2007, Soberón and Nakamura, 2009, Lloyd 2012,
Monahan and Tingley 2012). NicheA can generate such
forms, calculate their volume, density, shape, position, and

Figure 1. Feasible action flows in NicheA, showing four function modules. 1) The left block is the workflow for creating and displaying a
background cloud with environmental layers supplied by the user. 2) The second block shows generation of occurrence data sets, displaying
them in E, and computing their attributes. 3) The third block shows a simple function used to design barriers in G. 4) The final block
contains processes related to interacting with other modeling algorithms. N ⫽ virtual niche; BC ⫽ background; ENMs ⫽ ecological niche
models.

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other attributes, and quantify similarity among multiple
niches in terms of overlap in n-dimensional environmental
spaces. NicheA generates ecological niche models for virtual
species in E in the form of ellipsoids or polyhedrons, and
projects results to G. However, in view of the heterogeneous
distribution of populations within the niche with high spe-
cies abundance in core areas of the niche and low species
abundance at the edges (Martínez-Meyer et al. 2012, Lira-
Noriega and Manthey 2014), NicheA can relax polyhedron
assumptions, allowing projection of species distributions
based on continuous or logistic functions (Lira-Noriega and
Manthey 2014, Leroy et al. 2015).
Niches of virtual species can be generated using three dif-
ferent options. 1) Virtual niches can be created based on an
ellipsoid drawn manually selecting the shape, position, and
size of ellipsoids using the widget in the software interface,
which allows ellipsoid manipulation from the computer’s
mouse or via writing the ellipsoid’s dimension for detailed
estimations; then, this niche can be exported to the geog-
raphy using the ‘Create a virtual N’ tool in the ‘Toolbox’ of
NicheA. 2) Ecological niches can also be generated based
on environmental parameters (e.g. temperature range) under
diverse suitability distributions established by the user using
the ‘Virtual species – parameter’ tool. 3) Users interested in
generating virtual niches incorporating specific geographic
areas can create a virtual species based on geographic coordi-
nates resembling the site of interest using the ‘Generate N(s)
from occurrences’ tool.
Furthermore, using NicheA, it is possible to increase
complexity of virtual species by, for example, generating dis-
persal barriers in G and exploring associated configurations
in E, to simulate geographic configurations of species distri-
butions and biogeography. Thus, users are able to generate
virtual species of known, user-selected fundamental niche
characteristics; NicheA functions are summarized in Table 1.
These tools are available in NicheA ver. 3.0 9.14.2015, and
steps to develop virtual species are described in Table 2 and
Supplementary material Appendix 2.
Environmental variables
Two input data formats are used in NicheA to character-
ize environmental landscapes: GeoTIFF and ESRI ASCII
raster grid layers. NicheA uses data held in these inputs to
draw a background cloud that characterizes the universe
of environments available to a virtual species. According
to the scientific question and user preferences, environ-
mental variables can be from original (e.g. temperature),
summarized (i.e. principal components; Supplementary
material Appendix 3a), or virtual environments. NicheA
displays the distribution of the virtual niche in E based
on one, two, or three dimensions (Supplementary mate-
rial Appendix 3b). One dimension displays are the density
and range occupied by the virtual species with respect to
each variable, whereas two and three dimensions are visu-
alized in the virtual scenario. Users can calibrate models
and generate multivariate analyses in NicheA using the
n dimensions of E; however, the cost of overfitting and
redundant data layers in analyses should be considered
(Peterson 2007).
Outputs
Different functions in NicheA have different outputs. The
main output, which is defined as the virtual niche, N, in
NicheA, is a folder including the following files: geographic
coordinates of occurrences (ll.txt), plane coordinates (xy.txt),
environmental values of the occurrences (values.txt), and the
species’ geographic distribution in both GeoTIFF (present.
tiff) and PNG (present.png) formats. Models can be gener-
ated in binary and continuous formats. Continuous models
based distribution of suitability using linear and logistic func-
tions as in Leroy et al. (2015), but incorporating the ability
of applying such functions to all the environmental condi-
tions available or restricted to environments falling inside
the virtual niche based on a hyperdimensional ellipsoid.
Niches from virtual species in the form of ellipsoids can be
exported in editable (.elp) and non-editable formats (.mve),
that contain the elements of the ellipsoid (the centroid and
the covariance matrix) and are readable in NicheA.
Occurrence points drawn randomly from within a vir-
tual niche can be exported as a table with geographic coor-
dinates mirroring a continuous model with areas of high or
low suitability, arranging column labels in formats required
by popular ecological niche modeling software packages
(e.g. Maxent and OpenModeller; Phillips and Dudík 2008,
Muñoz et al. 2011; Supplementary material Appendix 3c).
Models of the virtual species in continuous format from
other ecological niche modeling software platforms can be
evaluated in NicheA using the Partial-ROC AUC and AIC
metrics (Peterson et al. 2008 and Warren and Seifert 2011;
Supplementary material Appendix 3d and e). Continuous
models in raster format can also be converted to binary
based on nine threshold methods (Supplementary material
Appendix 3f ). As complementary outputs, NicheA offers
tables and figures associated with several analytical tools,
including niche overlap measurements in E, and descriptive
statistics of environmental variables, occurrence points, and
model evaluation (Supplementary material Appendix 3g).
Scientific workflow management system
Functions in NicheA are connected by the outputs. In other
words, an output from one function can be an input of
another function. NicheA provides a novel method to man-
age connections among functions via a scientific workflow
management system that composes and executes series of
computational or data manipulation steps (Fig. 2). Using
this tool, users can carry out a complex analysis process, and
these workflows can be shared, reused, and adapted. Users
can download shared workflows via the workflows reposi-
tory (Supplementary material Appendix 3h). Users can also
design their own workflows, and share them with other
scientists to develop replicable ecological niche analyses.
Virtual species example
The process to generate virtual species within NicheA is
simple, as we will illustrate by means of some worked exam-
ple analyses. A basic workflow of this example is available
as a default in NicheA for new users. First, we created an
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Table 1. Analytical tools available in NicheA.

Function Tool Description

Background data Principal component analysis Generate a principal component analysis (PCA) from selected raster
layers; result is a ‘background’ folder to generate a 2D or 3D
environmental space. This function also generates figures and tables
summarizing the analysis and correlation among variables and the R
script employed during the analysis.
Draw background – folder Select a background cloud from folder created during the PCA to display
a 3D environmental space.
Draw background – files Select a background cloud from raster layers selected by the user to
display a 2D or 3D environmental space.
Niche simulation Open (N)s Open one or several ecological niche (N) models from NicheA (folders)
or other software output (GeoTIFF/ASCII rasters).
Design barriers Open a geographic map that allows the user to create geographic
barriers. Areas selected will result in a portion of the virtual niche.
Clear scenario Remove all items from the current environmental space.
Create a virtual N Export the virtual niche to the geography based in the current editable
ellipsoid (i.e. white ellipsoid with colorful vertexes).
Generate virtual N(s) from Draw a virtual species from a coma delimited file (.csv) including the
occurrences species’ name, longitude, and latitude.
Generate virtual N(s) from ellipsoid Draw a virtual species from a previous saved minimum-volume ellipsoid
file (.mve) not editable.
Virtual species – parameter Create a virtual species based on selected physiological parameters.
User can determine the physiological limits of the virtual species and
the distribution of the frequency (i.e. uniform, normal, binomial,
poisson).
Save current selection Save current ellipsoid selection (.elp) for posterior edition in NicheA.
Open a selection Open an editable virtual niche (.elp).
Open selections Open more than one virtual niche (.elp).
Niche analysis tools Quantify niche overlap Measure the hypervolume overlap of two or more ellipsoids or
convex-polyhedrons.
N attributes Features of the virtual niche.
Import/export Export to ENMs Generate occurrence points from the virtual species to be used in
different ecological niche model software. Occurrences can be drawn
from areas with high or low probability based on a logistic function.
All occurrences can be exported with different suitability values and
then converted from point to raster using the ‘Convert points to raster’
tool.
Close N/ Close N: remove virtual niches from the background.
export N as continuous raster/ Export N as continuous raster: generate a species distribution model in
export MVE configurations GeoTIFF raster format with continuous values of the distance to the
niche centroid.
Export MVE configurations: export virtual niche as ellipsoid file (.elp) to
be open and edit later in an environmental space in NicheA.
Post-ENM analysis Threshold calculator Generate binary maps from continuous models based on 10 different
functions. Binary model and occurrences are required. Environmental
rasters can also be thresholded to show suitable areas according to
each environmental variable.
Occurrence statistics Describe the density distribution in environmental terms of the virtual
niche, according to geographic coordinates, a binary niche model,
and the environmental values available in the study area.
Change the range of the ENM’s Normalization technique to convert continuous model from values
result ranging between 0 and 100 or between 0 and 1.
Calculate AUC value of partial-area Useful for model comparison. Determines if the continuous models
ROC approaches predict independent occurrences better than by chance, considering
the area predicted in a binary format. Diverse binary thresholds can
be tested. Continuous models and occurrences are required (Peterson
2012).
Calculate AIC/BIC values Useful for model comparison. Determine Akaike information criterion
values of Maxent models (Phillips and Dudík 2008) for model
comparisons (Warren and Seifert 2011).
Utility functions Raster conversion Convert raster from GeoTIFF to ASCII files and vice versa.
Variable statistics Maximum, mean, minimum, and range values of the environmental
variables selected.
Read data from multiple raster Extract values from different environmental variables based on one
layers geographic coordinate.
Variable normalization/ Two methods available for normalization of environmental variables for
standardization better visualization of the background to create the virtual niche.
Set no data value Allows users to change the no data value of raster files to fit with
different software requirements.
Convert points to raster Convert occurrence points to raster file.

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Table 2. Steps to create virtual species using NicheA. Next, we collected 100 occurrences drawn at random
Steps Process Tool from N and generated new virtual niches from this sample.
The niche estimates that resulted were projected to geog-
Step 1 Generate background Principal component analysis,
draw background – folder raphy according to models based on a minimum-volume
Step 2 Draw virtual species Draw a virtual niche manually ellipsoid and a convex polyhedron. A detailed example and
using the widget in the left explanation are in the online manual (Supplementary mate-
panel of NicheA, generate rial Appendix 3j). Figure 3 illustrates the completed outputs
virtual N(s) from occurrences
or virtual species – parameter
of the process above.
Step 3 Import virtual species Save current selection To display the E and G link, which is a function unique
(ellipsoid), export to ENMs to NicheA, we generated geographic barriers (e.g. ocean, des-
(coordinates), export N as ert, river) for the virtual species in the geographic view, and
continuous raster (map), used the ‘Design barriers’ tool to divide G into four subsets
export MVE configurations
(ellipsoid) that could then be displayed in E (Fig. 4, 5). Finally, we mea-
Step 4 Virtual species analyses Design barriers, quantify niches sured the overlap between the resulting minimum-volume
overlap, N attributes, ellipsoids from the geographic footprint, as divided by the
threshold calculator hypothesized geographic barriers. Details of this process can
be found in the online manual (Supplementary material
environmental background using 19 bioclimatic variables Appendix 3k).
from the WorldClim climate data archive (Hijmans et al. Additional analytical tools in multivariate environmen-
2005) at 10’ spatial resolution. In NicheA, we calculated tal spaces, and the use of NicheA to manage other ENM
the principal components of the 19 variables, and drew a outputs are described in detail in NicheA’s user manual
background cloud using the first 3 principal components in (< http://nichea.sourceforge.net/ >). Video tutorials and case
the E viewer. Then, we created a virtual niche using widgets studies are available in both English and Spanish (< http://
from the NicheA interface to draw an ellipsoid representing bit.ly/1O9tmaQ >).
a virtual species’ niche (Supplementary material Appendix
3i; Fig. 2). To save the virtual species, we used the ‘Create a
virtual N’ tool. We then visualized the virtual niche, N, in Discussion
the geographic space based on environmental values inside
the minimum-volume ellipsoid we created, and projected NicheA provides a GUI-based tool by which to generate
the environments corresponding to a convex polyhedron virtual species based on physiological ranges, geographic
from the same ellipsoid to the geography. coordinates generated by the user, or a manual selection

Figure 2. NicheA interface. (A) The set of tool options by which to generate, analyze, and manage ecological niche models in environmen-
tal space. (B) Widget by which to generate virtual species manually using the mouse to move the bars or via entering specific dimensions
of the niche. (C) Environmental space window where ecological niches can be viewed in a three-dimensional environmental scenario as a
background cloud (gray points) with three virtual ecological niches (green, yellow, and red ellipsoids).

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Figure 3. Process by which to generate a virtual species’ niche. (A) Virtual species niche based on an ellipsoid generated manually (white
ellipsoid) using NicheA’s widgets to specify external vertices as environmental semi-axes X, Y, and Z (red points) and the niche centroid
(green point). (B) Final virtual ecological niche in the form of a minimum volume ellipsoid (MVE; yellow ellipsoid). (C) Convex polyhe-
dron from the virtual ecological niche (CP; green lines). (D) Points falling inside the virtual ecological niche (red points).

of hypothetical environmental spaces occupied by the vir- and geographic spaces. NicheA provides users with tools
tual species. NicheA allows users to explore the Hutchinson with which to explore G and E spaces for virtual spe-
duality through visualizations of linked environmental cies, as opposed to the manipulations in G only typical of

Figure 4. Linking environmental and geographic spaces using NicheA. (A) Virtual ecological niche based on a minimum-volume ellipsoid
(yellow). (B) Geographic distribution corresponding to the minimum-volume ellipsoid (yellow). (C) Virtual ecological niche based on a
convex-polyhedron (green). (D) Geographic distribution corresponding to the convex-polyhedron (green). Notice that niche estimation
using minimum-volume ellipsoid generates broader predictions compared to convex-polyhedron.

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Figure 5. Linking environmental and geographic space using barriers. (A) Virtual species’ niche in geographic space showing different por-
tions of the original fundamental niche. The original virtual species’ potential distribution split into distinct populations: North America
(red), South America (green), South Africa (yellow), and Australia (blue). (B) Each ellipsoid represents a sub-portion of the species’ funda-
mental niche. Notice that although populations of different continents are spatially distant their niches overlap broadly in E.

geographic information systems applications. NicheA gen-
erates virtual species based on five approaches: minimum-
volume ellipsoids, convex polyhedrons, physiological ranges,
and linear and logistic functions (Leroy et al. 2015). Virtual
niches are displayed in one, two, or three environmental
dimensions, but analyses can be developed in any number
of dimensions.
Furthermore, NicheA is a toolkit including several inde-
pendent functions with which to analyze the complexity of
ecological niche models. Such functions can be arranged,
exported, and imported in the form of a workflow. By merg-
ing different functions via this tool, users can analyze diverse
problems creatively, without the limitations of single-func-
tion applications. Thus, NicheA is a workbench at which
users can address a variety of questions related to species’
ecological niches and geographic distributions.
Compared to existing tools for creating virtual species
including SDMvspecies and virtualspecies (Duan et al. 2015
and Leroy et al. 2015), NicheA has a GUI to manage the
virtual species’ niche in a multidimensional environmental
space (Table 3), something we found critical for robust study
design and accurate interpretation of outputs (Escobar et al.
2014). Beyond creating virtual niches in an environmental
space from a variety of possibilities (e.g. manual or via occur-
rence data), NicheA allows the user to analyze the features
of the virtual niche and to visualize and evaluate continuous
model outputs from diverse ecological niche model algo-
rithms (e.g. Maxent). A virtual niche from NicheA can also
Table 3. Environmental scenario manipulation in NicheA using the
computer’s mouse.
Operation Commands
Rotate scenario Left click ⫹ mouse movement
Zoom in/out scenario Alt ⫹ left click ⫹ mouse movement
Move scenario Right click ⫹ mouse movement

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be opened using environmental backgrounds from different
geographic areas or past or future climates. The ability of
NicheA to draw virtual niches from geographic coordinates
created by the user, when specific areas are selected to be
included in the virtual niche, allows importing real coordi-
nates from real species. Owing to the versatility of NicheA,
the software has already been employed in a diversity of stud-
ies using virtual and real species, in fields ranging from epi-
demiology to evolutionary biology. To date, NicheA has been
used to facilitate the study of hantavirus ecology in Brazil (de
Oliveira et al. 2013), the potential for the spread of white-
nose syndrome in bats (Escobar et al. 2014), species limits in
opossums (Tocchio et al. 2014), spatial and environmental
distribution of bats (Escobar et al. 2015a), niche divergence
in Asian gazelles (Hu et al. 2015), sympatric speciation in
butterflies (Rosser et al. 2015), effects of niche breath on
species survival (Saupe et al. 2015), anticipating the global
spread of Vibrio cholerae in marine ecosystems (Escobar et al.
2015b), and evaluating performance of ecological niche
model algorithms under diverse niche scenarios (Qiao et al.
2015).
NicheA is free, open-source software, available under the
GNU Lesser General Public License. All source code can be
retrieved via a subversion repository (< svn://mmweb.animal.
net.cn/nichea/trunk >). Because ecological niche modeling is
a rapidly-developing field that is seeing impressive research
use in addressing various questions in ecology, NicheA is
based in a community platform to meet increasing demand
of assistance in the form of forums, videos, an online man-
ual, and free remote support. Inclusion of additional model-
ling techniques (Blonder et al. 2014) is expected for future
NicheA versions. Finally, although NicheA has the capability
for developing complex analyses under multivariate environ-
mental scenarios; the user-friendly interface makes this soft-
ware an ideal resource for teaching in ecology.
To cite NicheA or acknowledge its use, cite this Software
note as follows, substituting the version of the application
that you used for ‘version 0’:
Qiao, H., Peterson, A. T., Campbell, L. P., Soberón, J., Ji, L. and
Escobar, L. E. 2016. NicheA: creating virtual species and
ecological niches in multivariate environmental scenarios.
– Ecography 39: 000–000 (ver. 0).
Acknowledgements – The authors thank Sergio Estay for his com-
ments on the NicheA installation process. This study was supported
by the National Natural Science Foundation of China (A New
Method to Predict the Species Distributions, 31100390), the Sci-
ence and Technology Supporting Project of Ministry of Science,
Microsoft Research Grant #47780, and NSF grant DEB-
1208472.
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