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      Allometric shell growth in infaunal burrowing bivalves: examples of the archiheterodonts Claibornicardia paleopatagonica (Ihering, 1903) and Crassatella kokeni Ihering, 1899

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          Abstract

          We present two cases of study of ontogenetic allometry in outlines of bivalves using longitudinal data, a rarity among fossils, based on the preserved post-larval record of shells. The examples are two infaunal burrowing bivalves of the southern South America, Claibornicardia paleopatagonica (Archiheterodonta: Carditidae) (early Paleocene) and Crassatella kokeni (Archiheterodonta: Crassatellidae) (late Oligocene–late Miocene). Outline analyses were conducted using a geometric morphometric approach (Elliptic Fourier Analysis), obtaining successive outlines from shells’ growth lines, which were used to reconstruct ontogenetic trajectories. In both taxa, ontogenetic changes are characterized by the presence of positive allometry in the extension of posterior end, resulting in elongated adult shells. This particular allometric growth is known in others infaunal burrowing bivalves ( Claibornicardia alticostata and some Spissatella species) and the resulting adult morphology is present in representatives of several groups (e.g., Carditidae, Crassatellidae, Veneridae, Trigoniidae). Taxonomic, ecological and evolutionary implications of this allometric growth pattern are discussed.

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          Heterochrony and allometry: the analysis of evolutionary change in ontogeny.

          The connection between development and evolution has become the focus of an increasing amount of research in recent years, and heterochrony has long been a key concept in this relation. Heterochrony is defined as evolutionary change in rates and timing of developmental processes; the dimension of time is therefore an essential part in studies of heterochrony. Over the past two decades, evolutionary biologists have used several methodological frameworks to analyse heterochrony, which differ substantially in the way they characterize evolutionary changes in ontogenies and in the resulting classification, although they mostly use the same terms. This review examines how these methods compare ancestral and descendant ontogenies, emphasizing their differences and the potential for contradictory results from analyses using different frameworks. One of the two principal methods uses a clock as a graphical display for comparisons of size, shape and age at a particular ontogenic stage, whereas the other characterizes a developmental process by its time of onset, rate, and time of cessation. The literature on human heterochrony provides particularly clear examples of how these differences produce apparent contradictions when applied to the same problem. Developmental biologists recently have extended the concept of heterochrony to the earliest stages of development and have applied it at the cellular and molecular scale. This extension brought considerations of developmental mechanisms and genetics into the study of heterochrony, which previously was based primarily on phenomenological characterizations of morphological change in ontogeny. Allometry is the pattern of covariation among several morphological traits or between measures of size and shape; unlike heterochrony, allometry does not deal with time explicitly. Two main approaches to the study of allometry are distinguished, which differ in the way they characterize organismal form. One approach defines shape as proportions among measurements, based on considerations of geometric similarity, whereas the other focuses on the covariation among measurements in ontogeny and evolution. Both are related conceptually and through the use of similar algebra. In addition, there are close connections between heterochrony and changes in allometric growth trajectories, although there is no one-to-one correspondence. These relationships and outline links between different analytical frameworks are discussed.
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            A guide to the nomenclature of heterochrony

            Since Haeckel's Biogenetic Law (‘ontogeny recapitulates phylogeny’) fell into disrepute early in the twentieth century, there has been intermittent debate, particularly in recent years (de Beer, 1958; Gould, 1977; Alberch et al., 1979; Alberch, 1980; Bonner, 1982; McNamara, 1982a), on the nature of the relationship between an individual's development and phylogenetic history. Important questions under discussion include the following: If a strong causal relationship does exist, what is its nature? How does it work? What is its importance in evolution? How can it be recognized in the fossil record?
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              SHAPE: a computer program package for quantitative evaluation of biological shapes based on elliptic Fourier descriptors.

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                Author and article information

                Contributors
                Journal
                PeerJ
                PeerJ
                peerj
                peerj
                PeerJ
                PeerJ Inc. (San Francisco, USA )
                2167-8359
                19 June 2018
                2018
                : 6
                : e5051
                Affiliations
                [-1] División Paleoinvertebrados, Museo Argentino de Ciencias Naturales “Bernardino Rivadavia” , Buenos Aires, Argentina
                Article
                5051
                10.7717/peerj.5051
                6014312
                e67ca309-b4d2-45b7-b925-7ae96a4ca748
                ©2018 Perez and Santelli

                This is an open access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, reproduction and adaptation in any medium and for any purpose provided that it is properly attributed. For attribution, the original author(s), title, publication source (PeerJ) and either DOI or URL of the article must be cited.

                History
                : 1 March 2018
                : 2 June 2018
                Funding
                Funded by: CONICET
                The authors were supported by CONICET (Argentina) through doctoral and post-doctoral grants. The funders had no role in study design, data collection and analysis, decision to publish, or preparation of the manuscript.
                Categories
                Evolutionary Studies
                Marine Biology
                Paleontology
                Zoology

                geometric morphometrics,ontogeny,crassatella,claibornicardia,allometric growth,paleoecology,archiheterodonta,elliptic fourier analysis

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