Conodont animals were early jawless vertebrates equipped with a feeding apparatus composed of several tooth-like elements. The P1 elements, at the rear of the apparatus, were characterized by a robust shape and rapid morphological evolution. Occlusion occurred between paired right and left P1 elements, occasioning some bilateral asymmetry, which, together with allometric growth, may partially obliterate the temporal differences. The present study aims to disentangle these different components of morphological variation in Late Devonian Polygnathus P1 conodont elements. An extensive 2D geometric morphometric analysis of the platform shape was performed through the Famennian record of two outcrops. This analysis was completed by a 3D study on a subset of conodont elements. The 2D and 3D morphometric quantifications provided highly congruent results, showing that the 2D shape constitutes a good approximation of the element geometry. The 3D analysis delivered further insights into the relationship between the geometry of the elements and the constraints related to occlusion. The 2D analysis allowed a quantitative assessment of the variation among species and through time. Allometry and bilateral asymmetry were differently expressed depending on the species considered, suggesting that constraints imposed on pairing by the morphology of the elements varied even among related species. The within-species variation was so important that it largely obliterated temporal trends; a relationship of Polygnathus shape and conodont biofacies variations through the Famennian nevertheless suggested an evolution driven by ecological interactions between conodont genera.

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Patterns of bilateral asymmetry and allometry in Late Devonian Polygnathus conodonts

This contribution contains the 3D models of the set of Famennian conodont elements belonging to the species Polygnathus glaber and Polygnathus communis analyzed in the following publication: Renaud et al. 2021: Patterns of bilateral asymmetry and allometry in Late Devonian Polygnathus. Palaeontology,

/pdf/3d-models-related-to-the-publication-patterns-of-bilateral-xpjfnq3dj5.pdf

3D models related to the publication: Patterns of bilateral asymmetry and allometry in Late Devonian Polygnathus conodonts

Phenotypic variation is the raw material of evolution. Standing variation can facilitate response to selection along "lines of least evolutionary resistance", but selection itself might alter the structure of the variance. Shape was quantified using 2D geometric morphometrics in Palmatolepis conodonts through the Late Devonian period. Patterns of variance were characterized along the record by the variance-covariance matrix (P-matrix) and its first axis (Pmax). The Late Frasnian was marked by environmental oscillations culminating with the Frasnian/Famennian mass extinction. A shape response was associated with these fluctuations, together with a deflection of the Pmax and the P-matrix. Thereafter, along the Famennian, Palmatolepis mean shape shifted from broad elements with a large platform to slender elements devoid of platform. This shift in shape was associated with a reorientation of Pmax and the P-matrix, due to profound changes in the functioning of the elements selecting for new types of variants. Both cases provide empirical evidences that moving adaptive optimum can reorient phenotypic variation, boosting response to environmental changes. On such time scales, the question seems thus not to be whether the P-matrix is stable, but how it is varying in response to changes in selection regimes and shifts in adaptive optimum.

Morphometric variance, evolutionary constraints and their change through time in Late Devonian Palmatolepis conodonts

Can we predict the evolutionary response of organisms to climate changes? The direction of greatest intraspecific phenotypic variance is thought to correspond to an ‘evolutionary line of least resistance’, i.e. a taxon's phenotype is expected to evolve along that general direction, if not constrained otherwise. In particular, heterochrony, whereby the timing or rate of developmental processes are modified, has often been invoked to describe evolutionary trajectories and it may be advantageous to organisms when rapid adaptation is critical. Yet, to date, little is known empirically as to which covariation patterns, whether static allometry, as measured in adult forms only, or ontogenetic allometry, the basis for heterochrony, may be prevalent in what circumstances. Here, we quantify the morphology of segminiplanate conodont elements during two distinct time intervals separated by more than 130 Myr: the Devonian-Carboniferous boundary and the Carnian-Norian boundary (Late Triassic). We evidence that the corresponding species share similar patterns of intraspecific static allometry. Yet, during both crises, conodont evolution was decoupled from this common evolutionary line of least resistance. Instead, it followed heterochrony-like trajectories that furthermore appear as driven by ocean temperature. This may have implications for our interpretation of conodonts' and past marine ecosystems’ response to environmental perturbations.

/pdf/temperature-driven-heterochrony-as-a-main-evolutionary-3a6o3l49.pdf

Temperature-driven heterochrony as a main evolutionary response to climate changes in conodonts

Identification of relevant taxonomic and evolutionary units is a recurrent issue in the fossil record, and all the more for ancient fossils devoid of modern equivalents such as conodonts. Extensive morphological variation has often led to the description of numerous species, subspecies or morphotypes, which may correspond to end-member morphologies reached through ontogeny. The platform elements of the Late Devonian conodont species Icriodus alternatus were characterized by rows of denticles coming into occlusion between opposite elements; each element grew by the incremental addition of lamellae and by the addition of successive triads during ontogeny. During the late Frasnian and the early Famennian, the important morphological variation within this species led to the description of three subspecies. An extensive sample of early Famennian Icriodus alternatus was quantified using 2D biometric measurements and denticle counts on 2D pictures, showing that the subspecies mainly differed in their size range but not in their general morphology. A 3D morphometric analysis was further performed on a subsample to characterize the shape of the ontogenetically older part of the elements. During ontogeny, early valleys between denticles tended to be filled, and the asymmetry between the inner and outer side of the element increased. These ontogenetic trends are responsible for the morphologies formerly described as the subspecies Ic. alt. mawsonae and Ic. alt. helmsi. Slight discrepancies between temporal ranges of the subspecies may be achieved through variations in range of size reached by the elements as a response to environmental changes. Disparity along ontogeny seems to follow an “hourglass model” suggesting a shift from relatively loose developmental constraints to a pattern of growth modulated by functional constraints during occlusion.

https://fr.pensoft.net/article/80211/download/pdf/

Ronan Ledevin

Papers

Patterns of bilateral asymmetry and allometry in Late Devonian Polygnathus conodonts

3D models related to the publication: Patterns of bilateral asymmetry and allometry in Late Devonian Polygnathus conodonts