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Staged embryonic series are important as reference for different kinds of biological studies. I summarise problems that occur when using 'staging tables' of 'model organisms'. Investigations of developmental processes in a broad scope of taxa are becoming commonplace. Beginning in the 1990s, methods were developed to quantify and analyse developmental events in a phylogenetic framework. The algorithms associated with these methods are still under development, mainly due to difficulties of using non-independent characters. Nevertheless, the principle of comparing clearly defined newly occurring morphological features in development (events) in quantifying analyses was a key innovation for comparative embryonic research. Up to date no standard was set for how to define such events in a comparative approach. As a case study I compared the external development of 23 land vertebrate species with a focus on turtles, mainly based on reference staging tables. I excluded all the characters that are only identical for a particular species or general features that were only analysed in a few species. Based on these comparisons I defined 104 developmental characters that are common either for all vertebrates (61 characters), gnathostomes (26), tetrapods (3), amniotes (7), or only for sauropsids (7). Characters concern the neural tube, somite, ear, eye, limb, maxillary and mandibular process, pharyngeal arch, eyelid or carapace development. I present an illustrated guide listing all the defined events. This guide can be used for describing developmental series of any vertebrate species or for documenting specimen variability of a particular species. The guide incorporates drawings and photographs as well as consideration of species identifying developmental features such as colouration. The simple character-code of the guide is extendable to further characters pertaining to external and internal morphological, physiological, genetic or molecular development, and also for other vertebrate groups not examined here, such as Chondrichthyes or Actinopterygii. An online database to type in developmental events for different stages and species could be a basis for further studies in comparative embryology. By documenting developmental events with the standard code, sequence heterochrony studies (i.e. Parsimov) and studies on variability can use this broad comparative data set.

The anterior end of the mammalian face is characteristically composed of a semimotile nose, not the upper jaw as in other tetrapods. Thus, the therian nose is covered ventrolaterally by the “premaxilla,” and the osteocranium possesses only a single nasal aperture because of the absence of medial bony elements. This stands in contrast to those in other tetrapods in whom the premaxilla covers the rostral terminus of the snout, providing a key to understanding the evolution of the mammalian face. Here, we show that the premaxilla in therian mammals (placentals and marsupials) is not entirely homologous to those in other amniotes; the therian premaxilla is a composite of the septomaxilla and the palatine remnant of the premaxilla of nontherian amniotes (including monotremes). By comparing topographical relationships of craniofacial primordia and nerve supplies in various tetrapod embryos, we found that the therian premaxilla is predominantly of the maxillary prominence origin and associated with mandibular arch. The rostral-most part of the upper jaw in nonmammalian tetrapods corresponds to the motile nose in therian mammals. During development, experimental inhibition of primordial growth demonstrated that the entire mammalian upper jaw mostly originates from the maxillary prominence, unlike other amniotes. Consistently, cell lineage tracing in transgenic mice revealed a mammalian-specific rostral growth of the maxillary prominence. We conclude that the mammalian-specific face, the muzzle, is an evolutionary novelty obtained by overriding ancestral developmental constraints to establish a novel topographical framework in craniofacial mesenchyme.

The origin of turtles is one of the most long-lasting debates in evolutionary research. During their evolution, a series of modifications changed their relatively kinetic and anapsid skull into an elongated akinetic structure with a unique pulley system redirecting jaw adductor musculature. These modifications were thought to be strongly correlated to functional adaptations, especially to bite performance. We conducted a series of Finite Element Analyses (FEAs) of several species, including that of the oldest fully shelled, Triassic stem-turtle Proganochelys , to evaluate the role of force distribution and to test existing hypotheses on the evolution of turtle skull architecture. We found no support for a relation between the akinetic nature of the skull or the trochlear mechanisms with increased bite forces. Yet, the FEAs show that those modifications changed the skull architecture into an optimized structure, more resistant to higher loads while allowing material reduction on specific regions. We propose that the skull of modern turtles is the result of a complex process of progressive correlation between their heads and highly flexible necks, initiated by the origin of the shell.

Like other diapsids, Tyrannosaurus rex has two openings in the temporal skull region. In addition, like in other dinosaurs, its snout and lower jaw show large cranial fenestrae. In T . rex , they are thought to decrease skull weight, because, unlike most other amniotes, the skull proportion is immense compared to the body. Understanding morphofunctional complexity of this impressive skull architecture requires a broad scale phylogenetic comparison with skull types different to that of dinosaurs with fundamentally diverging cranial regionalization. Extant fully terrestrial vertebrates (amniotes) provide the best opportunities in that regard, as their skull performance is known from life. We apply for the first time anatomical network analysis to study skull bone integration and modular constructions in tyrannosaur and compare it with five representatives of the major amniote groups in order to get an understanding of the general patterns of amniote skull modularity. Our results reveal that the tyrannosaur has the most modular skull organization among the amniotes included in our study, with an unexpected separation of the snout in upper and lower sub-modules and the presence of a lower adductor chamber module. Independent pathways of bone reduction in opossum and chicken resulted in different degrees of cranial complexity with chicken having a typical sauropsidian pattern. The akinetic skull of opossum, alligator, and leatherback turtle evolved in independent ways mirrored in different patterns of skull modularity. Kinetic forms also show great diversity in modularity. The complex tyrannosaur skull modularity likely represents a refined mosaic of phylogenetic and ecological factors with food processing being probably most important for shaping its skull architecture. Mode of food processing primarily shaped skull integration among amniotes, however, phylogenetic patterns of skull integration are low in our sampling. Our general conclusions on amniote skull integrity are obviously preliminary and should be tested in subsequent studies. As such, this study provides a framework for future research focusing on the evolution of modularity on lower taxonomic levels.

The unparalleled ability of turtle neck retraction is possible in three different modes, which characterize stem turtles, living side-necked (Pleurodira), and hidden-necked (Cryptodira) turtles, respectively. Despite the conservatism in vertebral count among turtles, there is significant functional and morphological regionalization in the cervical vertebral column. Since Hox genes play a fundamental role in determining the differentiation in vertebra morphology and based on our reconstruction of evolutionary genetics in deep time, we hypothesize genetic differences among the turtle groups and between turtles and other land vertebrates. We correlated anterior Hox gene expression and the quantifiable shape of the vertebrae to investigate the morphological modularity in the neck across living and extinct turtles. This permitted the reconstruction of the hypothetical ancestral Hox code pattern of the whole turtle clade. The scenario of the evolution of axial patterning in turtles indicates shifts in the spatial expression of HoxA-5 in relation to the reduction of cervical ribs in modern turtles and of HoxB-5 linked with a lower morphological differentiation between the anterior cervical vertebrae observed in cryptodirans. By comparison with the mammalian pattern, we illustrate how the fixed count of eight cervical vertebrae in turtles resulted from the emergence of the unique turtle shell.

The labyrinth of the vertebrate inner ear is a sensory system that governs the perception of head rotations. Central hypotheses predict that labyrinth shape and size are related to ecological adaptations, but this is under debate and has rarely been tested outside of mammals. We analyze the evolution of labyrinth morphology and its ecological drivers in living and fossil turtles, an understudied group that underwent multiple locomotory transitions during 230 million years of evolution. We show that turtles have unexpectedly large labyrinths that evolved during the origin of aquatic habits. Turtle labyrinths are relatively larger than those of mammals, and comparable to many birds, undermining the hypothesis that labyrinth size correlates directly with agility across vertebrates. We also find that labyrinth shape variation does not correlate with ecology in turtles, undermining the widespread expectation that reptilian labyrinth shapes convey behavioral signal, and demonstrating the importance of understudied groups, like turtles. The size and shape of the inner ear, or bony labyrinth, is thought to be related to ecological adaptations in vertebrates. Here, the authors examine this relationship in turtles across 230 million years of evolution, unexpectedly finding large labyrinth size and no association with ecology.

Whereas hundreds of breeds of domestic dogs are known, only several dozen domestic cat breeds are currently recognized, and the ferret is not classified into specific breeds. We studied pre- and postnatal patterns of development and growth in the domesticated forms of these three carnivoran species. We present the most comprehensive staging system for domestic dog embryos to date and define qualitative characters for phylogenetic comparisons. For postnatal development, we present analyses of new and literature measurements of cranial and limb proportions. We analyze changes in the progress of growth among different domestic dog and domestic cat breeds. All three domesticated forms drastically differ in the relative timing of prenatal development. This is correlated with ontogenetic plasticity at birth, which enables artificial selection to act. For postnatal development, we detected a greater shape variance in domestic dog ontogeny when compared to that of the domestic cat. We conclude that ontogenetic preconditions as well as body size constrain the species’ capability for artificial selection in domestic dogs and cats. However, we speculate that the human requirements for functional performance of their domesticates might render some developmental biases substantially. Although ferrets would be preferable for artificial selection given their plastic embryonic development, they have been of less interest for domestication due to their small body size - by which they were already well adapted for hunting in burrows - and due to the fact that other relevant tasks were already assumed by domestic cats and dogs since earlier phases of human cultural evolution.

Following the traditional and holistic concept of Vergleichende Entwicklungsgeschichte , Wolfgang Maier studied different aspects of vertebrate morphology, including dentition, the locomotor apparatus, nasal and ear regions. His work comprises investigations on pre- to postnatal stages of extant species as well as fossils and is mainly based on histological serial sections, but also on µCT data in recent years. This resulted in an integrative research agenda on the evolutionary biology of mammals and other vertebrates. Most of his studies are designed around the interrelationship of ontogenetic and functional adaptations and evolutionary transformations. The present collection in Vertebrate Zoology collates a series of research articles related to and in honor of Wolfgang Maier’s work. Invited colleagues of Maier provide current insights to their own research, in many cases inspired by his scholarship, ranging from mammalian to reptilian and fish comparative anatomy. In total, this volume contains 19 publications. They apply modern and traditional techniques to investigate the diversity of biological form. In so doing, they integrate traditional conceptual frameworks from the earliest days of morphological research more than two centuries ago.

The evolution of skull diversity in Crocodylia is rather well documented, but the developmental foundation of their cranial architecture remains poorly understood. Here, we present the first three-dimensional reconstruction of the embryonic cartilaginous skull of Caiman crocodilus based on histological sections. We provide a comprehensive anatomical description and morphometric analysis of the chondrocranium of this species, integrating linear measurements and comparative anatomical data to assess interspecific variation and evaluate whether closely related taxa exhibit greater similarity in chondrocranial morphology. We identified both qualitative and quantitative differences between major crocodilian clades, which may reflect diverse ecological demands. Within Crocodylidae, orbitotemporal proportions strongly influence chondrocranial morphology, likely reflecting adaptations related to bite force and visual acuity, linked to prey type and diverse aquatic habits. Within Alligatoridae, by contrast, the emphasis on nasal capsule proportions suggests a reliance on olfactory and respiratory functions, consistent with their occupation of densely vegetated environments and more restricted geographic ranges. Additionally, we identified a set of embryonic diagnostic features located in the splanchnocranium of Caiman and in the neurocranium of Alligatoridae. These findings shed new light on the developmental basis of cranial diversity in Crocodylia and emphasize the evolutionary significance of chondrocranial traits in shaping macroevolutionary patterns.

Development in marsupials is specialized towards an extremely short gestation and highly altricial newborns. As a result, marsupial neonates display morphological adaptations at birth related to functional constraints. However, little is known about the variability of marsupial skull development and its relation to morphological diversity. We studied bony skull development in five marsupial species. The relative timing of the onset of ossification was compared to literature data and the ossification sequence of the marsupial ancestor was reconstructed using squared-change parsimony. The high range of variation in the onset of ossification meant that no patterns could be observed that differentiate species. This finding challenges traditional studies concentrating on the onset of ossification as a marker for phylogeny or as a functional proxy. Our study presents observations on the developmental timing of cranial bone-to-bone contacts and their evolutionary implications. Although certain bone contacts display high levels of variation, connections of early and late development are quite conserved and informative. Bones that surround the oral cavity are generally the first to connect and the bones of the occipital region are among the last. We conclude that bone contact is preferable over onset of ossification for studying cranial bone development.