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      Two contemporaneous morphs of fossil Chanos Lacepède, 1803 (Gonorynchiformes, Chanidae) from Paleocene (Danian) outcrops near Palenque (Mexico) revealed by geometric morphometrics indicate conservatism in milkfishes after the K/Pg boundary

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          Abstract

          We described chanid material from the Paleocene (Danian) localities of División del Norte and Belisario Domínguez near the archeological site of Palenque, Chiapas State, southeastern Mexico. The parsimony-based morphological phylogeny indicates that the specimens are closely related to the extant milkfish, Chanos chanos (Teleostei, Ostariophysi), and the comparative anatomy reveals a remarkable qualitative similarity of almost every visible bone. Among the synapomorphies for the genus Chanos, is a pleurostyle or caudal complex, which is missing in all other chanid fossils. Extant milkfish are highly variable in meristic and morphometric traits, and we found a signal for quantitative variation with geometric morphometrics tools in the fossil sample. We first dealt with post-mortem body torsion and missing landmarks. The main analysis shows a pattern of two forms present in both localities. A group of specimens shows a bigger head and deeper body than the slender and smaller head of the rest, implying that two types of milkfish coexisted in time and space. We tested for allometry and explored scenarios that can explain the patterns, such as sexual dimorphism or two sympatric and closely related species for the morphotypes, and differential resource utilization for the jaw, head, and depth variations. Furthermore, we argue that, alongside the morphological stasis, Chanos has conserved the life history trait of fry migration towards near-shore nurseries through protracted time (~63mybp). We infer the fish were juvenile, and the paleontological assemblage and taphonomy suggest that the localities exhibit the influence of both marine and transitional environments.

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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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            Extensions of the Procrustes Method for the Optimal Superimposition of Landmarks

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              geomorph: anrpackage for the collection and analysis of geometric morphometric shape data

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                Author and article information

                Contributors
                Role: ConceptualizationRole: Data curationRole: InvestigationRole: MethodologyRole: Project administrationRole: VisualizationRole: Writing – original draftRole: Writing – review & editing
                Role: ConceptualizationRole: Data curationRole: Funding acquisitionRole: Project administrationRole: ResourcesRole: ValidationRole: VisualizationRole: Writing – review & editing
                Role: Editor
                Journal
                PLoS One
                PLoS One
                plos
                PLOS One
                Public Library of Science (San Francisco, CA USA )
                1932-6203
                5 March 2025
                2025
                : 20
                : 3
                : e0313912
                Affiliations
                [1 ] Posgrado en Ciencias Biológicas, Universidad Nacional Autónoma de México, Ciudad de México, México
                [2 ] Departamento de Paleontología, Instituto de Geología, Universidad Nacional Autónoma de México, Ciudad de México, México
                Università degli Studi di Torino, ITALY
                Author notes

                Competing Interests: The authors have declared that no competing interests exist.

                Author information
                https://orcid.org/0009-0005-6413-1098
                Article
                PONE-D-24-33173
                10.1371/journal.pone.0313912
                11882075
                40043070
                695b6f0f-9d84-4643-8b0f-0dab850f62fd
                © 2025 Guadarrama, Cantalice

                This is an open access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.

                History
                : 5 August 2024
                : 18 October 2024
                Page count
                Figures: 15, Tables: 3, Pages: 33
                Funding
                Funded by: funder-id http://dx.doi.org/10.13039/501100003141, Consejo Nacional de Ciencia y Tecnología;
                Award ID: CVU1191092
                Award Recipient :
                Funded by: funder-id http://dx.doi.org/10.13039/501100006087, Dirección General de Asuntos del Personal Académico, Universidad Nacional Autónoma de México;
                Award ID: IA206123
                Award Recipient :
                AG received financial support by the postgraduate scholarship of the Consejo Nacional de Humanidades, Ciencias y Tecnologías (CONAHCYT), with CVU number 1191092. Additional financial support for AG comes from PAEP-UNAM-2023 and DGAPA-PAPIIT project 206123. KMC received the grant from Dirección General de Asuntos del Personal Acádemico-Programa de Apoyo a Proyectos de Investigación e Innovación Tecnológica (DGAPA-PAPIIT) number 206123. The funders had no role in study design, data collection and analysis, decision to publish, or preparation of the manuscript.
                Categories
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                Research and Analysis Methods
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