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Species distribution modelling (SDM), also known as environmental (or ecological) niche modelling (ENM), habitat modelling, predictive habitat distribution modelling, and range mapping uses
computer A computer is a machine that can be programmed to Execution (computing), carry out sequences of arithmetic or logical operations (computation) automatically. Modern digital electronic computers can perform generic sets of operations known as C ...
algorithm In mathematics and computer science, an algorithm () is a finite sequence of rigorous instructions, typically used to solve a class of specific Computational problem, problems or to perform a computation. Algorithms are used as specificat ...
s to predict the
distribution Distribution may refer to: Mathematics *Distribution (mathematics), generalized functions used to formulate solutions of partial differential equations * Probability distribution, the probability of a particular value or value range of a vari ...
of a species across
geographic Geography (from Greek: , ''geographia''. Combination of Greek words ‘Geo’ (The Earth) and ‘Graphien’ (to describe), literally "earth description") is a field of science devoted to the study of the lands, features, inhabitants, and ...
space and time using environmental data. The environmental data are most often climate data (e.g. temperature, precipitation), but can include other variables such as soil type, water depth, and land cover. SDMs are used in several research areas in
conservation biology Conservation biology is the study of the conservation of nature and of Earth's biodiversity with the aim of protecting species, their habitats, and ecosystems from excessive rates of extinction and the erosion of biotic interactions. It is an int ...
,
ecology Ecology () is the study of the relationships between living organisms, including humans, and their physical environment. Ecology considers organisms at the individual, population, community, ecosystem, and biosphere level. Ecology overlaps wi ...
and
evolution Evolution is change in the heritable characteristics of biological populations over successive generations. These characteristics are the expressions of genes, which are passed on from parent to offspring during reproduction. Variation ...
. These models can be used to understand how environmental conditions influence the occurrence or abundance of a species, and for predictive purposes (
ecological forecasting Ecological forecasting uses knowledge of physics, ecology and physiology to predict how ecological populations, communities, or ecosystems will change in the future in response to environmental factors such as climate change. The goal of the approac ...
). Predictions from an SDM may be of a species’ future distribution under climate change, a species’ past distribution in order to assess evolutionary relationships, or the potential future distribution of an invasive species. Predictions of current and/or future habitat suitability can be useful for management applications (e.g. reintroduction or translocation of vulnerable species, reserve placement in anticipation of climate change). There are two main types of SDMs. Correlative SDMs, also known as climate envelope models, bioclimatic models, or resource selection function models, model the observed distribution of a species as a function of environmental conditions. Mechanistic SDMs, also known as process-based models or biophysical models, use independently derived information about a species' physiology to develop a model of the environmental conditions under which the species can exist. The extent to which such modelled data reflect real-world species distributions will depend on a number of factors, including the nature, complexity, and accuracy of the models used and the quality of the available environmental data layers; the availability of sufficient and reliable species distribution data as model input; and the influence of various factors such as barriers to dispersal, geologic history, or
biotic interaction In ecology, a biological interaction is the effect that a pair of organisms living together in a community have on each other. They can be either of the same species (intraspecific interactions), or of different species (interspecific interaction ...
s, that increase the difference between the
realized niche In ecology, a niche is the match of a species to a specific environmental condition. Three variants of ecological niche are described by It describes how an organism or population responds to the distribution of resources and competitors (for ...
and the fundamental niche. Environmental niche modelling may be considered a part of the discipline of
biodiversity informatics Biodiversity informatics is the application of informatics techniques to biodiversity information, such as taxonomy, biogeography or ecology. Modern computer techniques can yield new ways to view and analyze existing information, as well as predict ...
.


History

A. F. W. Schimper used geographical and environmental factors to explain plant distributions in his 1898 ''Pflanzengeographie auf physiologischer Grundlage'' (''Plant Geography Upon a Physiological Basis''). Andrew Murray used the environment to explain the distribution of mammals in his 1866 ''The Geographical Distribution of Mammals''. Robert Whittaker's work with plants and Robert MacArthur's work with birds strongly established the role the environment plays in species distributions. Elgene O. Box constructed environmental envelope models to predict the range of tree species. His computer simulations were among the earliest uses of species distribution modelling. The adoption of more sophisticated
generalised linear model In statistics, a generalized linear model (GLM) is a flexible generalization of ordinary linear regression. The GLM generalizes linear regression by allowing the linear model to be related to the response variable via a ''link function'' and by ...
s (GLMs) made it possible to create more sophisticated and realistic species distribution models. The expansion of
remote sensing Remote sensing is the acquisition of information about an object or phenomenon without making physical contact with the object, in contrast to in situ or on-site observation. The term is applied especially to acquiring information about Earth ...
and the development of GIS-based environmental modelling increase the amount of environmental information available for model-building and made it easier to use.


Correlative vs mechanistic models


Correlative SDMs

SDMs originated as correlative models. Correlative SDMs model the observed distribution of a species as a function of geographically referenced climatic predictor variables using multiple regression approaches. Given a set of geographically referred observed presences of a species and a set of climate maps, an algorithm finds the most likely environmental ranges within which a species lives. Correlative SDMs assume that species are at equilibrium with their environment and that the relevant environmental variables have been adequately sampled. The models allow for interpolation between a limited number of species occurrences. For these algorithms to be effective, it is required to gather observations not only of species presences, but also of absences, that is, where the species does not live. Records of species absences are typically not as common as records of presences, thus often "random background" or "pseudo-absence" data are used to fit these models. If there are incomplete records of species occurrences, pseudo-absences can introduce bias. Since correlative SDMs are models of a species’ observed distribution, they are models of the
realized niche In ecology, a niche is the match of a species to a specific environmental condition. Three variants of ecological niche are described by It describes how an organism or population responds to the distribution of resources and competitors (for ...
(the environments where a species ''is'' found), as opposed to the
fundamental niche In ecology, a niche is the match of a species to a specific environmental condition. Three variants of ecological niche are described by It describes how an organism or population responds to the distribution of resources and competitors (for ...
(the environments where a species ''can'' be found, or where the abiotic environment is appropriate for the survival). For a given species, the realized and fundamental niches might be the same, but if a species is geographically confined due to dispersal limitation or species interactions, the
realized niche In ecology, a niche is the match of a species to a specific environmental condition. Three variants of ecological niche are described by It describes how an organism or population responds to the distribution of resources and competitors (for ...
will be smaller than the
fundamental niche In ecology, a niche is the match of a species to a specific environmental condition. Three variants of ecological niche are described by It describes how an organism or population responds to the distribution of resources and competitors (for ...
. Correlative SDMs are easier and faster to implement than mechanistic SDMs, and can make ready use of available data. Since they are correlative however, they do not provide much information about causal mechanisms and are not good for extrapolation. They will also be inaccurate if the observed species range is not at equilibrium (e.g. if a species has been recently introduced and is actively expanding its range).


Mechanistic SDMs

Mechanistic SDMs are more recently developed. In contrast to correlative models, mechanistic SDMs use physiological information about a species (taken from controlled field or laboratory studies) to determine the range of environmental conditions within which the species can persist. These models aim to directly characterize the fundamental niche, and to project it onto the landscape. A simple model may simply identify threshold values outside of which a species can't survive. A more complex model may consist of several sub-models, e.g.
micro-climate A microclimate (or micro-climate) is a local set of atmospheric conditions that differ from those in the surrounding areas, often with a slight difference but sometimes with a substantial one. The term may refer to areas as small as a few squ ...
conditions given macro-climate conditions, body temperature given micro-climate conditions, fitness or other biological rates (e.g. survival, fecundity) given body temperature (thermal performance curves), resource or energy requirements, and
population dynamics Population dynamics is the type of mathematics used to model and study the size and age composition of populations as dynamical systems. History Population dynamics has traditionally been the dominant branch of mathematical biology, which has ...
. Geographically referenced environmental data are used as model inputs. Because the species distribution predictions are independent of the species’ known range, these models are especially useful for species whose range is actively shifting and not at equilibrium, such as invasive species. Mechanistic SDMs incorporate causal mechanisms and are better for extrapolation and non-equilibrium situations. However, they are more labor-intensive to create than correlational models and require the collection and validation of a lot of physiological data, which may not be readily available. The models require many assumptions and parameter estimates, and they can become very complicated. Dispersal, biotic interactions, and evolutionary processes present challenges, as they aren’t usually incorporated into either correlative or mechanistic models. Correlational and mechanistic models can be used in combination to gain additional insights. For example, a mechanistic model could be used to identify areas that are clearly outside the species’ fundamental niche, and these areas can be marked as absences or excluded from analysis. See for a comparison between mechanistic and correlative models.


Niche modelling algorithms (correlative)

There are a variety of mathematical methods that can be used for fitting, selecting, and evaluating correlative SDMs. Algorithms include "profile" methods, which are simple statistical techniques that use e.g. environmental distance to known sites of occurrence such a
BIOCLIM
ref name="Nix1986a">
and DOMAIN; "regression" methods (e.g. forms of generalized linear models); and "
machine learning Machine learning (ML) is a field of inquiry devoted to understanding and building methods that 'learn', that is, methods that leverage data to improve performance on some set of tasks. It is seen as a part of artificial intelligence. Machine ...
" methods such as maximum entropy (MAXENT). Ten machine learning algorithms used in SDM can be seen in. An incomplete list of algorithms that have been used for niche modelling includes:


Profile techniques

* BIOCLIM *
DOMAIN Domain may refer to: Mathematics *Domain of a function, the set of input values for which the (total) function is defined **Domain of definition of a partial function **Natural domain of a partial function **Domain of holomorphy of a function * Do ...
*
Ecological niche factor analysis Ecology () is the study of the relationships between living organisms, including humans, and their physical environment. Ecology considers organisms at the individual, population, community, ecosystem, and biosphere level. Ecology overlaps w ...
(ENFA) * Mahalanobis distance * Isodar analysis


Regression-based techniques

*
Generalized linear model In statistics, a generalized linear model (GLM) is a flexible generalization of ordinary linear regression. The GLM generalizes linear regression by allowing the linear model to be related to the response variable via a ''link function'' and b ...
(GLM) *
Generalized additive model In statistics, a generalized additive model (GAM) is a generalized linear model in which the linear response variable depends linearly on unknown smooth functions of some predictor variables, and interest focuses on inference about these smooth func ...
(GAM) *
Multivariate adaptive regression splines In statistics, multivariate adaptive regression splines (MARS) is a form of regression analysis introduced by Jerome H. Friedman in 1991. It is a non-parametric regression technique and can be seen as an extension of linear models that automaticall ...
(MARS) * Maxlike * Favourability Function (FF)


Machine learning techniques

*
MAXENT The principle of maximum entropy states that the probability distribution which best represents the current state of knowledge about a system is the one with largest entropy, in the context of precisely stated prior data (such as a proposition ...
*
Artificial neural networks Artificial neural networks (ANNs), usually simply called neural networks (NNs) or neural nets, are computing systems inspired by the biological neural networks that constitute animal brains. An ANN is based on a collection of connected unit ...
(ANN) *
Genetic Algorithm for Rule Set Production Genetic Algorithm for Rule Set Production (GARP) is a computer program based on genetic algorithm that creates Environmental niche modelling, ecological niche models for species. The generated models describe environmental conditions (precipitation ...
(GARP) * Boosted regression trees (BRT)/gradient boosting machines (GBM) *
Random forest Random forests or random decision forests is an ensemble learning method for classification, regression and other tasks that operates by constructing a multitude of decision trees at training time. For classification tasks, the output of th ...
(RF) *
Support vector machines In machine learning, support vector machines (SVMs, also support vector networks) are supervised learning models with associated learning algorithms that analyze data for classification and regression analysis. Developed at AT&T Bell Laboratorie ...
(SVM) * XGBoost (XGB) Furthermore, ensemble models can be created from several model outputs to create a model that captures components of each. Often the mean or median value across several models is used as an ensemble. Similarly, consensus models are models that fall closest to some measure of central tendency of all models—consensus models can be individual model runs or ensembles of several models.


Niche modelling software (correlative)

SPACES
is an online Environmental niche modeling platform that allows users to design and run dozens of the most prominent algorithms in a high performance, multi-platform, browser-based environment.
MaxEnt
is the most widely used method/software uses presence only data and performs well when there are few presence records available.
ModEco
implements various algorithms. DIVA-GIS has an easy to use (and good for educational use) implementation of BIOCLIM
The Biodiversity and Climate Change Virtual Laboratory (BCCVL)
is a "one stop modelling shop" that simplifies the process of biodiversity and climate impact modelling. It connects the research community to Australia's national computational infrastructure by integrating a suite of tools in a coherent online environment. Users can access global climate and environmental datasets or upload their own data, perform data analysis across six different experiment types with a suite of 17 different algorithms, and easily visualise, interpret and evaluate the results of the models. Experiments types include: Species Distribution Model, Multispecies Distribution Model, Species Trait Model (currently under development), Climate Change Projection, Biodiverse Analysis and Ensemble Analysis. Example of BCCVL SDM outputs can be foun
here
Another example is Ecocrop, which is used to determine the suitability of a crop to a specific environment. This database system can also project crop yields and evaluate the impact of environmental factors such as
climate change In common usage, climate change describes global warming—the ongoing increase in global average temperature—and its effects on Earth's climate system. Climate change in a broader sense also includes previous long-term changes to E ...
on plant growth and suitability. Most niche modelling algorithms are available in the R package
'dismo'


an
'mopa'
. Software developers may want to build on th
openModeller
project. The Collaboratory for Adaptation to Climate Chang
adapt.nd.edu
has implemented a
online version of openModeller
that allows users to design and run openModeller in a high-performance, browser-based environment to allow for multiple parallel experiments without the limitations of local processor power.


See also

*
Biogeography Biogeography is the study of the distribution of species and ecosystems in geographic space and through geological time. Organisms and biological communities often vary in a regular fashion along geographic gradients of latitude, elevation, ...
*
Ecosystem model An ecosystem model is an abstract, usually mathematical, representation of an ecological system (ranging in scale from an individual population, to an ecological community, or even an entire biome), which is studied to better understand the re ...
*
Quantum evolution Quantum evolution is a component of George Gaylord Simpson's multi-tempoed theory of evolution proposed to explain the rapid emergence of higher taxonomic groups In biology, taxonomy () is the scientific study of naming, defining ( circumscr ...


References


Further reading

* * *


External links


Climate Envelope Modeling Working Group
- Online gathering place for scientists, practitioners, managers, and developers to discuss, support, and develop climate Environmental Niche Modeling tools and platforms
BioVeL Ecological Niche Modeling (ENM)
- online tool with workflows to generate ecological niche models
EUBrazilOpenBio SpeciesLab Virtual Research Environment
- online working environment to support the production of ecological niche modeling by (i) simplifying access to occurrence points and environmental parameters and (ii) offering a powerful version of openModeller benefitting from a distributed computing infrastructure;
openModeller
- open source niche modelling library
lifemapper
- niche modelling project from Kansas University
Lifemapper 2.0
- video of presentation by Aimee Stewart, Kansas University, at O'Reilly Where 2.0 Conference 2008
AquaMaps
- global predictive maps for marine species
Ecological Modelling
- International Journal on Ecological Modelling and Systems Ecology {{modelling ecosystems, expanded=other Biogeography Ecological niche Information science Landscape ecology Environmental modelling