Morphological characters in light of new molecular phylogenies: the caudal-fin skeleton of Ovalentaria

The Ovalentaria is a taxon of teleosts that has been proposed based on molecular analyses only. Previously widely separated families are assembled in this taxon. For the first time, the Ovalentaria are analysed using a comparative morphological approach. The caudal-fin skeleton of 355 species covering all 48 ovalentarian families are examined in cleared and stained specimens, µCT datasets and X-ray images as well as from the literature. A total of 38 morphological characters are evaluated and used for ancestral character state reconstructions and phylogenetic analyses. Results provide hypotheses for a scenario of the evolution of the caudal-fin skeleton and its ground plan in Ovalentaria. An evolutionary trend towards the reduction of skeletal elements in the caudal fin is observed. Connections between the evolution of the caudal-fin skeleton and modes of locomotion found in ovalentarian taxa are discussed. Phylogenetic analyses based on the caudal-fin morphology provide topologies for intra-ovalentarian relationships that largely agree with molecular hypotheses.

The accessory neural arch: development, morphology, and systematic distribution

The accessory neural arch is an oddly distributed character present in several non-acanthomorph teleostean taxa. Its homology was often implied but never satisfyingly tested. In this study, we attended this pending problem. We analyzed the morphology, development, and systematic distribution of the accessory neural arch in teleosts. Using a comprehensive taxon sampling of cleared and stained specimens, we evaluated if the accessory neural arch fulfils existing homology criteria. We then combined these data with recent genetic phylogenies and ancestral character state estimation to reconstruct the evolutionary history of the accessory neural arch. While its gross morphology and development fit homology criteria, results from ancestral character state estimations suggest multiple independent evolutions within teleosts. Although the accessory neural arch cannot be homologous between several teleostean taxa, the concept of parallelism may explain the presence of such a similar character in a variety of non-acanthomorph teleostean taxa.

A New Cynodont from the Late Triassic Los Colorados Formation (Argentina, South America) Reveals a Novel Paleobiogeographic Context for Mammalian Ancestors

Probainognathia is a derived lineage of cynodonts which encompass Mammalia as their crown-group. The profuse record of probainognathians from the Carnian of Argentina contrasts with their Norian representation, with only one named species. Here we describe a new probainognathian, Tessellatia bonapartei gen. et sp. nov., from the Norian Los Colorados Formation of the Ischigualasto-Villa Unión Basin of Argentina. The new taxon, represented by a partial cranium with articulated lower jaws, was analyzed through neutron and X-rays micro-tomography (µCT). The high-resolution neutron µCT data allowed the identification of a unique character state combination, including features inaccessible through traditional techniques. We constructed the largest phylogenetic data-matrix of non-mammalian cynodonts. The new species and its sister-taxon, the Brazilian Therioherpeton, are recovered as probainognathians, closely related to Mammaliamorpha. We conducted the first quantitative paleobiogeographic analysis of non-mammalian cynodonts, focusing in probainognathians. The results indicate that Probainognathia and Mammaliamorpha originated in Brazil, which was an important center of diversification during the Triassic. Finally, China is identified as the ancestral area of Mammaliaformes. These new findings, besides adding to the knowledge of the poorly represented Norian cynodonts from the Los Colorados Formation, are significant to improve our understanding of probainognathian diversity, evolution, and paleobiogeographic history.

Loss of a larval generic character: an interesting and new description for Isoperla vevcianensis Ikonomov, 1980 (Plecoptera: Perlodidae) with updated adult characters

The larva of the Balkan microendemic Isoperla vevcianensis Ikonomov, 1980 (Plecoptera: Perlodidae: Isoperlinae) is described on the basis of larvae associated with adults by means of cox1 sequences similarities. Eggs and everted penis of the male are described for the first time. The larva possessed blunt paraprocts which were previously sharp among Palaearctic larvae of this huge Holarctic and Oriental genus. The commonly pointed paraproct remains as the only distinguishing generic character for the morphologically diverse larvae of Palaearctic Isoperla, but should not be considered as an exclusive character state for the genus. Morphological characters are illustrated in comparison with the sympatric larvae of the Isoperla tripartita species complex.

New Remains of Scandiavis mikkelseni Inform Avian Phylogenetic Relationships and Brain Evolution

Although an increasing number of studies are combining skeletal and neural morphology data in a phylogenetic context, most studies do not include extinct taxa due to the rarity of preserved endocasts. The early Eocene avifauna of the Fur Formation of Denmark presents an excellent opportunity for further study of extinct osteological and endocranial morphology as fossils are often exceptionally preserved in three dimensions. Here, we use X-ray computed tomography to present additional material of the previously described taxon Scandiavis mikkelseni and reassess its phylogenetic placement using a previously published dataset. The new specimen provides novel insights into the osteological morphology and brain anatomy of Scandiavis. The virtual endocast exhibits a morphology comparable to that of modern avian species. Endocranial evaluation shows that it was remarkably similar to that of certain extant Charadriiformes, yet also possessed a novel combination of traits. This may mean that traits previously proposed to be the result of shifts in ecology later in the evolutionary history of Charadriiformes may instead show a more complex distribution in stem Charadriiformes and/or Gruiformes depending on the interrelationships of these important clades. Evaluation of skeletal and endocranial character state changes within a previously published phylogeny confirms both S. mikkelseni and a putative extinct charadriiform, Nahmavis grandei, as charadriiform. Results bolster the likelihood that both taxa are critical fossils for divergence dating and highlight a biogeographic pattern similar to that of Gruiformes.

Multiple trait breeding programs with genotype-by-environment interactions based on reaction norms, with application to genetic improvement of disease resilience

Background Genotype-by-environment interactions for a trait can be modeled using multiple-trait, i.e. character-state, models, that consider the phenotype as a different trait in each environment, or using reaction norm models based on a functional relationship, usually linear, between phenotype and a quantitative measure of the quality of the environment. The equivalence between character-state and reaction norm models has been demonstrated for a single trait. The objectives of this study were to extend the equivalence of the reaction norm and character-state models to a multiple-trait setting and to both genetic and environmental effects, and to illustrate the application of this equivalence to the design and optimization of breeding programs for disease resilience. Methods Equivalencies between reaction norm and character-state models for multiple-trait phenotypes were derived at the genetic and environmental levels, which demonstrates how multiple-trait reaction norm parameters can be derived from multiple-trait character state parameters. Methods were applied to optimize selection for a multiple-trait breeding goal in a target environment based on phenotypes collected in a healthy and disease-challenged environment, and to optimize the environment in which disease-challenge phenotypes should be collected. Results and conclusions The equivalence between multiple-trait reaction norm and multiple-trait character-state parameters allow genetic improvement for a multiple-trait breeding goal in a target environment to be optimized without recording phenotypes and estimating parameters for the target environment.

Evolutionary development of the cephalopod arm armature: a review

The cephalopod arm armature is certainly one of the most important morphological innovations responsible for the evolutionary success of the Cephalopoda. New palaeontological discoveries in the recent past afford to review and reassess origin and homology of suckers, sucker rings, hooks, and cirri. Since a priori character state reconstructions are still ambiguous, we suggest and discuss three different evolutionary scenarios. Each of them is based on the following assumptions: (1) Neocoleoidea uniting extant Decabrachia and Octobrachia is monophyletic (= proostracum-bearing coleoids); (2) extinct Belemnitida and Diplobelida are stem decabrachians; (3) proostracum-less coleoids (Hematitida, Donovaniconida, Aulacoceratida) represent stem-neocoleoids; (4) Ammonoidea and Bactritoidea are stem coleoids. We consider a scenario where belemnoid hooks derived from primitive suckers as well-supported. Regarding belemnoid hooks and suckers as homologues implies that belemnoid, oegopsid, and probably ammonoid arm hooks arose through parallel evolution. Our conclusions challenge the widespread opinion, whereupon belemnoid hooks evolved de novo, and instead support earlier ideas formulated by Sigurd von Boletzky.

Character correlation and its use for identification

A method is presented for correlating phylogenetic characters through cladistic analysis. It extends the use of phylogenetic datasets for diagnostic purposes. It improves matrix-based identification tools by predicting novel character-state combinations that were not observed when the key was constructed. By interpreting homoplasy as analytical error, hypothetical character-state combinations are tested for the homoplasy that they would add to the shortest tree(s). The correlation is equal to the homoplasy summed across all state combinations, divided by a maximum possible value. The results depend on uncertainty about the sequence of state transitions and their overlap among characters. A correlation index r is proposed for sets of non-additive characters; it is a kind of multiple-regression value, and its ensemble value R is a statistic of a whole matrix. This approach can be used to select sets of the best "proxy" characters to substitute for unobservable characters of interest. The concept can be extended to continuous characters. Worked examples are given with datasets of various insect orders.

Hidden-state-only speciation and extinction models provide accurate tip estimates of diversification rates

1. State-dependent speciation and extinction (SSE) models provide a framework for testing potential correlations between the evolution of an observed trait and speciation and extinction rates. Recent expansions of these models allow for the inclusion of "hidden states" that, among other things, allow for rate heterogeneity often observed among lineages sharing a particular character state. However, in reality, multiple circumstances and interacting traits related to a focal character play a role in changing diversification dynamics of a lineage over time, restricting the use of available SSE models that require trait information to be assigned at the tips. 2. Here we introduce MiSSE, an SSE approach that infers diversification rate differences from hidden states only. It can be used similarly to other trait-free methods to estimate varying speciation, extinction, but also different functions of these parameters such as net-diversification, turnover rates, and extinction fraction. Given the size of the model space, we also describe an algorithm designed for efficiently searching through a reasonably large set of models without having to be exhaustive. 3. We compare the accuracy of rates inferred at the tips of the tree by MiSSE against popular character-free methods and demonstrate that the error associated with tip estimates is generally low. Due to certain characteristics of the SSE models, this method avoids some of the recent concerns with parameter identifiability in diversification analyses and can be used alongside regular phylogenetic comparative methods in trait-related diversification hypotheses. 4. Finally, we apply MiSSE, with a renewed focus on classic comparative methods, to understand processes happening near the present, rather than deep in the past, to examine how variation in plant height has impacted turnover rates in eucalypts, a species-rich lineage of flowering plants.

Character-state evaluation when discriminating Thysanoptera taxa (Insecta)

Problems are discussed in evaluating character state variation in the insect order Thysanoptera, particularly in the family Phlaeothripidae, illustrated by two such problems in the genus Haplothrips Amyot & Serville. The synonymy of Jironiella Retana-Salazar & Soto-Rodriguez with Haplothrips is reconfirmed, and Haplothrips aliakbarii Mirab-balou & Miri is recognised as a new synonym of H. globiceps Bagnall.