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      Morphological Species Delimitation in The Western Pond Turtle ( Actinemys): Can Machine Learning Methods Aid in Cryptic Species Identification?

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          Synopsis

          As the discovery of cryptic species has increased in frequency, there has been an interest in whether geometric morphometric data can detect fine-scale patterns of variation that can be used to morphologically diagnose such species. We used a combination of geometric morphometric data and an ensemble of five supervised machine learning methods (MLMs) to investigate whether plastron shape can differentiate two putative cryptic turtle species, Actinemys marmorata and Actinemys pallida. Actinemys has been the focus of considerable research due to its biogeographic distribution and conservation status. Despite this work, reliable morphological diagnoses for its two species are still lacking. We validated our approach on two datasets, one consisting of eight morphologically disparate emydid species, the other consisting of two subspecies of Trachemys ( T. scripta scripta, T. scripta elegans). The validation tests returned near-perfect classification rates, demonstrating that plastron shape is an effective means for distinguishing taxonomic groups of emydids via MLMs. In contrast, the same methods did not return high classification rates for a set of alternative phylogeographic and morphological binning schemes in Actinemys. All classification hypotheses performed poorly relative to the validation datasets and no single hypothesis was unequivocally supported for Actinemys. Two hypotheses had machine learning performance that was marginally better than our remaining hypotheses. In both cases, those hypotheses favored a two-species split between A. marmorata and A. pallida specimens, lending tentative morphological support to the hypothesis of two Actinemys species. However, the machine learning results also underscore that Actinemys as a whole has lower levels of plastral variation than other turtles within Emydidae, but the reason for this morphological conservatism is unclear.

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          Cryptic species as a window on diversity and conservation.

          The taxonomic challenge posed by cryptic species (two or more distinct species classified as a single species) has been recognized for nearly 300 years, but the advent of relatively inexpensive and rapid DNA sequencing has given biologists a new tool for detecting and differentiating morphologically similar species. Here, we synthesize the literature on cryptic and sibling species and discuss trends in their discovery. However, a lack of systematic studies leaves many questions open, such as whether cryptic species are more common in particular habitats, latitudes or taxonomic groups. The discovery of cryptic species is likely to be non-random with regard to taxon and biome and, hence, could have profound implications for evolutionary theory, biogeography and conservation planning.
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            Species concepts and species delimitation.

            The issue of species delimitation has long been confused with that of species conceptualization, leading to a half century of controversy concerning both the definition of the species category and methods for inferring the boundaries and numbers of species. Alternative species concepts agree in treating existence as a separately evolving metapopulation lineage as the primary defining property of the species category, but they disagree in adopting different properties acquired by lineages during the course of divergence (e.g., intrinsic reproductive isolation, diagnosability, monophyly) as secondary defining properties (secondary species criteria). A unified species concept can be achieved by treating existence as a separately evolving metapopulation lineage as the only necessary property of species and the former secondary species criteria as different lines of evidence (operational criteria) relevant to assessing lineage separation. This unified concept of species has several consequences for species delimitation, including the following: First, the issues of species conceptualization and species delimitation are clearly separated; the former secondary species criteria are no longer considered relevant to species conceptualization but only to species delimitation. Second, all of the properties formerly treated as secondary species criteria are relevant to species delimitation to the extent that they provide evidence of lineage separation. Third, the presence of any one of the properties (if appropriately interpreted) is evidence for the existence of a species, though more properties and thus more lines of evidence are associated with a higher degree of corroboration. Fourth, and perhaps most significantly, a unified species concept shifts emphasis away from the traditional species criteria, encouraging biologists to develop new methods of species delimitation that are not tied to those properties.
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              A guide to machine learning for biologists

              The expanding scale and inherent complexity of biological data have encouraged a growing use of machine learning in biology to build informative and predictive models of the underlying biological processes. All machine learning techniques fit models to data; however, the specific methods are quite varied and can at first glance seem bewildering. In this Review, we aim to provide readers with a gentle introduction to a few key machine learning techniques, including the most recently developed and widely used techniques involving deep neural networks. We describe how different techniques may be suited to specific types of biological data, and also discuss some best practices and points to consider when one is embarking on experiments involving machine learning. Some emerging directions in machine learning methodology are also discussed.
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                Author and article information

                Contributors
                Journal
                Integr Org Biol
                Integr Org Biol
                iob
                Integrative Organismal Biology
                Oxford University Press
                2517-4843
                2024
                02 April 2024
                02 April 2024
                : 6
                : 1
                : obae010
                Affiliations
                Department of Ecology and Evolution, Stony Brook University , Stony Brook, NY 11794, USA
                Center for Inclusive Education, Stony Brook University , Stony Brook, NY 11794, USA
                Department of Geological Sciences, California State University , Fullerton, CA 92834, USA
                Section of Research and Collections , NC Museum of Natural Sciences, Raleigh, NC 27601, USA
                952 NW 60th St., Seattle , Washington, WA 98107, USA
                Negaunee Integrative Research Center, Field Museum of Natural History , Chicago, IL 60605, USA
                Author notes
                Article
                obae010
                10.1093/iob/obae010
                11058871
                38689939
                e3c6ffa7-d0b0-4504-9ac7-0abfb296d7c4
                © The Author(s) 2024. Published by Oxford University Press on behalf of the Society for Integrative and Comparative Biology.

                This is an Open Access article distributed under the terms of the Creative Commons Attribution License ( https://creativecommons.org/licenses/by/4.0/), which permits unrestricted reuse, distribution, and reproduction in any medium, provided the original work is properly cited.

                History
                : 07 December 2023
                : 29 February 2024
                : 30 April 2024
                Page count
                Pages: 21
                Funding
                Funded by: National Science Foundation, DOI 10.13039/100000001;
                Award ID: DBI-0306158
                Award ID: DBI-0353797
                Funded by: National Institute of General Medical Sciences, DOI 10.13039/100000057;
                Funded by: National Institutes of Health, DOI 10.13039/100000002;
                Award ID: K12GM102778
                Categories
                Article
                AcademicSubjects/SCI00960

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