Modelling evolutionary tempo and mode using formal morphological spaces and Markov chain principles.

Evolutionary tempo and mode summarize ancient and controversial subjects of theoretical biology such as gradualism, convergence, contingence, trends, and entrenchment. We employed an integrative methodological approach to explore the evolutionary tempo and mode of Lepidosaurian phalangeal formulae (PFs). This approach involves quantifying the frequencies of morphological changes along an evolutionary trajectory. The five meristic characters encoded by PFs are particularly valuable in revealing evolutionary patterns, owing to their discrete nature and extensive documentation in the literature. Based on a pre-existing dataset of PFs from 649 taxa (35 Lepidosauria families, including fossils), from which there exists a unique repertoire of 53 formulations, our approach simultaneously considers phenetic and phylogenetic data. This culminates in a diagram accounting for the phylogenetic dynamic of evolution traversing across different regions of morphospace. The method involves enumerating phenotypical options, reconstructing phenotypes across the phylogeny, projecting phenotypes onto a morphospace, and constructing a flow network from the frequency of evolutionary transitions between unique phenotypic conditions. This approach links Markovian chains and evolutionary trajectories to formally define parameters that describe the underlying transitions of morphological change. Among other results, we found that (a) PF evolution exhibits a clear trend towards reduction in the phalangeal count and that (b) evolutionary change tends to occur significantly between morphologically similar PFs. Notwithstanding, although minor but not trivial, transitions between distant formulas -jumps- occur. Our results support a pluralistic view including stasis, gradualism, and saltationism discriminating their prevalence in a target character evolution.

Insights on the evolution of Coronavirinae in general, and SARS-CoV-2 in particular, through innovative biocomputational resources.

The structural proteins of coronaviruses portray critical information to address issues of classification, assembly constraints, and evolutionary pathways involving host shifts. We compiled 173 complete protein sequences from isolates belonging to the four genera of the subfamily Coronavirinae. We calculate a single matrix of viral distance as a linear combination of protein distances. The minimum spanning tree (MST) connecting the individuals captures the structure of their similarities. The MST re-capitulates the known phylogeny of Coronovirinae. Hosts were mapped onto the MST and we found a non-trivial concordance between host phylogeny and viral proteomic distance. We also study the chimerism in our dataset through computational simulations. We found evidence that structural units coming from loosely related hosts hardly give rise to feasible chimeras in nature. This work offers a fresh way to analyze features of SARS-CoV-2 and related viruses.

Morphospace analysis leads to an evo-devo model of digit patterning.

Biological forms occupy a constrained portion of theoretical morphospaces. Developmental models accounting for empirical morphospaces are necessary to achieve a better understanding of this phenomenon. We analyzed the phalangeal formulas (PFs) in lizards and relatives' hands by comparing them with a set of simulated PFs that compose a theoretical morphospace. We detected that: (1) the empirical morphospace is severely limited in size, (2) the PFs comply with two properties of phalangeal count per digit, namely the ordering rule (DI ≤ DII ≤ DIII ≤ DIV ≥ DV), and the contiguity relationship (neighbor digits differ on average in one phalanx), (3) the totality of the PFs can be categorized into four categories of hands aligned along a feasibility gradient. We also reconstructed the evolution of PFs and found a stepwise trajectory from the plesiomorphic PF towards reduced conditions. Finally, we propose a developmental model as the generative mechanism behind the PFs. It is consistent with the bulk of evidence managed and involves an ordered digit primordia initialization timed with periodic signals of joint formation coming from digit tips. Our approach is also useful to address the study of other meristic sequences in nature such as dental, floral, and branchial formulas.

Delimiting the boundaries of sesamoid identities under the network theory framework.

Sesamoid identity has long been the focus of debate, and how they are linked to other elements of the skeleton has often been considered relevant to their definition. A driving hypothesis of our work was that sesamoids' nature relies deeply on their connections, and thus we propose an explicit network framework to investigate this subject in Leptodactylus latinasus (Anura: Leptodactylidae). Through the dissection of L. latinasus' skeleton, we modeled its anatomical network where skeletal elements were considered nodes while joints, muscles, tendons, and aponeurosis were considered links. The skeletal elements were categorized into canonical skeletal pieces, embedded sesamoids, and glide sesamoids. We inquired about the general network characterization and we have explored further into sesamoid connectivity behavior. We found that the network is structured in a modular hierarchical organization, with five modules on the first level and two modules on the second one. The modules reflect a functional, rather than a topological proximity clustering of the skeleton. The 25 sesamoid pieces are members of four of the first-level modules. Node parameters (centrality indicators) showed that: (i) sesamoids are, in general terms, peripheral elements of the skeleton, loosely connected to the canonical bone structures; (ii) embedded sesamoids are not significantly distinguishable from canonical skeletal elements; and (iii) glide sesamoids exhibit the lowest centrality values and strongly differ from both canonical skeletal elements and embedded sesamoids. The loose connectivity pattern of sesamoids, especially glides, could be related to their evolvability, which in turn seems to be reflected in their morphological variation and facultative expression. Based on the connectivity differences among skeletal categories found in our study, an open question remains: can embedded and glide sesamoids be defined under the same criteria? This study presents a new approach to the study of sesamoid identity and to the knowledge of their morphological evolution.

Sesamoids in tetrapods: the origin of new skeletal morphologies.

Along with supernumerary bones, sesamoids, defined as any organized intratendinous/intraligamentous structure, including those composed of fibrocartilage, adjacent to an articulation or joint, have been frequently considered as enigmatic structures associated with the joints of the skeletal system of vertebrates. This review allows us to propose a dynamic model to account for part of skeletal phenotypic diversity: during evolution, sesamoids can become displaced, attaching to and detaching from the long bone epiphyses and diaphysis. Epiphyses, apophyses and detached sesamoids are able to transform into each other, contributing to the phenotypic variability of the tetrapod skeleton. This dynamic model is a new paradigm to delineate the contribution of sesamoids to skeletal diversity. Herein, we first present a historical approach to the study of sesamoids, discussing the genetic versus epigenetic theories of their genesis and growth. Second, we construct a dynamic model. Third, we present a summary of literature on sesamoids of the main groups of tetrapods, including veterinary and human clinical contributions, which are the best-studied aspects of sesamoids in recent decades. Finally, we discuss the identity of certain structures that have been labelled as sesamoids despite insufficient formal testing of homology. We also propose a new definition to help the identification of sesamoids in general. This review is particularly timely, given the recent increasing interest and research activity into the developmental biology and mechanics of sesamoids. With this updated and integrative discussion, we hope to pave the way to improve the understanding of sesamoid biology and evolution.

Bone indicators of grasping hands in lizards.

Grasping is one of a few adaptive mechanisms that, in conjunction with clinging, hooking, arm swinging, adhering, and flying, allowed for incursion into the arboreal eco-space. Little research has been done that addresses grasping as an enhanced manual ability in non-mammalian tetrapods, with the exception of studies comparing the anatomy of muscle and tendon structure. Previous studies showed that grasping abilities allow exploitation for narrow branch habitats and that this adaptation has clear osteological consequences. The objective of this work is to ascertain the existence of morphometric descriptors in the hand skeleton of lizards related to grasping functionality. A morphological matrix was constructed using 51 morphometric variables in 278 specimens, from 24 genera and 13 families of Squamata. To reduce the dimensions of the dataset and to organize the original variables into a simpler system, three PCAs (Principal Component Analyses) were performed using the subsets of (1) carpal variables, (2) metacarpal variables, and (3) phalanges variables. The variables that demonstrated the most significant contributions to the construction of the PCA synthetic variables were then used in subsequent analyses. To explore which morphological variables better explain the variations in the functional setting, we ran Generalized Linear Models for the three different sets. This method allows us to model the morphology that enables a particular functional trait. Grasping was considered the only response variable, taking the value of 0 or 1, while the original variables retained by the PCAs were considered predictor variables. Our analyses yielded six variables associated with grasping abilities: two belong to the carpal bones, two belong to the metacarpals and two belong to the phalanges. Grasping in lizards can be performed with hands exhibiting at least two different independently originated combinations of bones. The first is a combination of a highly elongated centrale bone, reduced palmar sesamoid, divergence angles above 90°, and slender metacarpal V and phalanges, such as exhibited by Anolis sp. and Tropidurus sp. The second includes an elongated centrale bone, lack of a palmar sesamoid, divergence angles above 90°, and narrow metacarpal V and phalanges, as exhibited by geckos. Our data suggest that the morphological distinction between graspers and non-graspers is demonstrating the existence of ranges along the morphological continuum within which a new ability is generated. Our results support the hypothesis of the nested origin of grasping abilities within arboreality. Thus, the manifestation of grasping abilities as a response to locomotive selective pressure in the context of narrow-branch eco-spaces could also enable other grasping-dependent biological roles, such as prey handling.