RESUMEN
Ontogeny is rarely included in phylogenetic analyses of morphological data. When used, the ontogenetic information is reduced to one character for two or three different ontogenetic stages. Several examples show that current methods miss a major part of the information. We here propose a new method for including the ontogenetic dimension in coding schemes of phylogenetic analyses. Our goal was to maximize the phylogenetic information extracted from ontogenetic trajectories. For discrete features, we recommend including precise timings of transformation(s) from one state to another in the ontogenetic trajectories. For continuously varying features, growth laws are modeled on raw data using least-square regressions. Then, parameters of models are included in the coding scheme as continuous characters. This method is employed to reconstruct phylogenetic relationships using the ammonite family Amaltheidae as a test subject. Based on the same data set, a second analysis has been performed only for characters of the adult stage. Comparisons of retention index, bootstrap support, and stratigraphic congruence between the two analyses show that the inclusion of ontogeny yields better phylogenetic reconstruction. Morphological traits in ammonites which are usually the most homoplastic show a better fit to most parsimonious trees by including the ontogenetic dimension. In several cases, growth rates and patterns of growth have better fit to phylogeny than adult shapes, implying that paths of ontogeny can be more relevant than its products.
Asunto(s)
Clasificación/métodos , Filogenia , Animales , Artrópodos/anatomía & histología , Artrópodos/clasificación , Artrópodos/crecimiento & desarrolloRESUMEN
Ammonite phylogeny has mainly been established based on a stratigraphic approach, with cladistics underconsidered. The main arguments against the use of cladistics are the supposed large amount of homoplasy and the small number of characters. Resolving the phylogeny of the Hildoceratidae (Early Jurassic) is especially challenging because of its large diversity and disparity. Many forms that have not been determined as closely related in previous studies exhibit very similar shapes. Moreover, some groups are morphologically very different, adding difficulties to building a unique coding scheme at a low taxonomic resolution (i.e. species). Here we propose an integrated coding scheme of the peristome shape and the ornamentation, allowing an increased level of comparison. The shape of the peristome is used as a new reference to locate ornamental features and propose new homology hypotheses. In total, 105 taxa have been analysed for 47 characters. We code continuous characters by their means and ranges ± one standard deviation. We test two weighting schemes: equal weights standardized by unit range and implied weighting with several concavity constants. This work has led to redefinition of the phylogenetic inclusivenesses of all the hildoceratid subfamilies. The new coding scheme based on peristome shapes provides the fewest homoplastic characters. The schemes appear promising to improve phylogenetic analyses in ammonoids as well as molluscs as a whole by creating a general coding framework.
RESUMEN
Since the introduction of the cladistic method in systematics, continuous characters have been integrated into analyses but no methods for their treatment have received unanimous support. Some methods require a large number of character states to discretise continuous characters in order to keep the maximum level of information about taxa differences within the coding scheme. Our objective was to assess the impact of increasing the character state number on the outcomes of phylogenetic analyses. Analysis of a variety of simulated datasets shows that these methods for coding continuous characters can lead to the generation of well-resolved trees that do not reflect a phylogenetic signal. We call this phenomenon the flattening of the tree-length distribution; it is influenced by both the relative quantity of continuous characters in relation to discrete characters, and the number of characters in relation to the number of taxa. Bootstrap tests provide a method to avoid this potential bias.
Asunto(s)
Clasificación/métodos , Filogenia , Animales , Biología Computacional , Modelos BiológicosRESUMEN
Ammonites are prominent in macroevolutionary studies because of their abundance and diversity in the fossil record, but their paleobiology and position in the marine food web are not well understood due to the lack of preserved soft tissue. We present three-dimensional reconstructions of the buccal apparatus in the Mesozoic ammonite Baculites with the use of synchrotron x-ray microtomography. Buccal mass morphology, combined with the coexistence of food remains found in the buccal mass, suggests that these ammonites fed on plankton. This diet may have extended to all aptychophoran ammonites, which share the same buccal mass morphology. Understanding the role of these ammonites in the Mesozoic food web provides insights into their radiation in the Early Jurassic, as well as their extinction at the end of the Cretaceous/early Paleogene.