Peramorphosis

Abstract: The bird skull arose from the nonavian dinosaur skull by several episodes of paedomorphosis, in which descendants resemble the juveniles of their ancestors, according to a study of shape change during dinosaur ontogeny and phylogeny. Much of what makes birds so different from other animals relates to the skull, specialized in birds to accommodate the visual and neuromuscular systems needed to coordinate flight and vision, and an ultra-adaptable beak. This comparison of skull morphology in birds with that of extinct theropod dinosaurs suggests that the characteristic features of the bird skull evolved in large part through paedomorphosis — the retention of juvenile features in the adult. With prominent eyes, larger brains and a short face, primitive stem-group birds resemble the embryos and juveniles of archosaurs, the non-avian branch of the theropod dinosaur line that includes the extant crocodylians. The interplay of evolution and development has been at the heart of evolutionary theory for more than a century1. Heterochrony—change in the timing or rate of developmental events—has been implicated in the evolution of major vertebrate lineages such as mammals2, including humans1. Birds are the most speciose land vertebrates, with more than 10,000 living species3 representing a bewildering array of ecologies. Their anatomy is radically different from that of other vertebrates. The unique bird skull houses two highly specialized systems: the sophisticated visual and neuromuscular coordination system4,5 allows flight coordination and exploitation of diverse visual landscapes, and the astonishing variations of the beak enable a wide range of avian lifestyles. Here we use a geometric morphometric approach integrating developmental, neontological and palaeontological data to show that the heterochronic process of paedomorphosis, by which descendants resemble the juveniles of their ancestors, is responsible for several major evolutionary transitions in the origin of birds. We analysed the variability of a series of landmarks on all known theropod dinosaur skull ontogenies as well as outgroups and birds. The first dimension of variability captured ontogeny, indicating a conserved ontogenetic trajectory. The second dimension accounted for phylogenetic change towards more bird-like dinosaurs. Basally branching eumaniraptorans and avialans clustered with embryos of other archosaurs, indicating paedomorphosis. Our results reveal at least four paedomorphic episodes in the history of birds combined with localized peramorphosis (development beyond the adult state of ancestors) in the beak. Paedomorphic enlargement of the eyes and associated brain regions parallels the enlargement of the nasal cavity and olfactory brain in mammals6. This study can be a model for investigations of heterochrony in evolutionary transitions, illuminating the origin of adaptive features and inspiring studies of developmental mechanisms.

Abstract: Heterochrony can be defined as change to the timing or rate of development relative to the ancestor. Because organisms generally change in shape as well as increase in size during their development, any variation to the duration of growth or to the rate of growth of different parts of the organism can cause morphological changes in the descendant form. Heterochrony takes the form of both increased and decreased degrees of development, known as “peramorphosis” and “paedomorphosis,” respectively. These are the morphological consequences of the operation of processes that change the duration of the period of an individual’s growth, either starting or stopping it earlier or later than in the ancestor, or by extending or contracting the period of growth. Heterochrony operates both intra- and interspecifically and is the source of much intraspecific variation. It is often also the cause of sexual dimorphism. Selection of a sequence of species with a specific heterochronic trait can produce evolutionary trends in the form of pera- or paedomorphoclines. Many different life history traits arise from the operation of heterochronic processes, and these may sometimes be the targets of selection rather than morphological features themselves. It has been suggested that some significant steps in evolution, such as the evolution of vertebrates, were engendered by heterochrony. Human evolution was fuelled by heterochrony, with some traits, such as a large brain, being peramorphic, whereas others, such as reduced jaw size, are paedomorphic.

Abstract: A model is proposed, based on examples that have been interpreted as phylogenetic trends, to explain how directional morphological evolution at the species level can arise by heterochrony. The examples illustrated are of Tertiary to Recent rhynchonellide brachiopods, Cambrian olenellid trilobites, living spatangoid echinoids, Tertiary to Recent schizasterid echinoids, Cenomanian ammonites and Si- lurian monograptids. Morphological discontinuities between species along morphological gradients (which can be recognised both spatially and/or temporally), and temporal morphological stasis within species, are both consistent with the punctuated equilibria model of macroevolution. It is argued that morpho- logical discontinuities have arisen by selection of morphological novelties produced by heterochronic processes. These novelties are preadaptations which allow ecological and, consequently, genetic isolation from ancestral species. Establishment of a heterochronic morphological gradient is only possible given a suitable environmental gradient. The terms "paedomorphocline" and "peramorphocline" are proposed for these heterochronic morphological gradients. Paedomorphoclines and peramorphoclines each comprise a number of species occupying a series of adaptive peaks, which have evolved sequentially through time by selection along an environmental gradient.

Abstract: Sturgeons (Acipenseridae) are an ancient and unique assemblage of fishes historically important to discussions of actinopterygian evolution. Despite their basal position within Actinopterygii, rigorous comparative morphological studies of acipenserids have never been made, and most ideas about acipenserid evolution hinge on an untested impression that shovelnose sturgeons (Scaphirhynchini) are phylogenetically primitive. This impression promoted ideas that: (1) the earliest acipenserids were highly benthic and evolved secondarily into pelagic predators, and (2) paedomorphosis has dominated mechanisms affecting their morphological change. Using cladistic methods, this study examines generic level interrelationships within Acipenseridae. Representatives of the four acipenserid genera Huso, Acipenser, Pseudoscaphirhynchus, and Scaphirhynchus, as well as their acipenseriform outgroups Polyodontidae, †Peipiaosteidae, and †Chondrosteidae, were surveyed for skeletal characters. Sixty-nine characters are identified and described to support the first generic level cladogram of Acipenseridae. Huso is phylogenetically primitive within Acipenseridae and the sister group to a redefined subfamily Acipenserinae. Acipenser is not supported by any characters identified in this study, but the tribe Scaphirhynchini comprising Scaphirhynchus and Pseudoscaphirhynchus is found to be monophyletic. The cladogram contradicts historical ideas about acipenserid evolution because Huso defines an outgroup morphology and life history founded on pelagic habitats and piscivory. In contrast, acipenserines, and more markedly scaphirhynchines, are benthic predators possessing character complexes for benthic feeding, respiration, locomotion, and protection. Also, the pattern of character acquisition within Acipenseridae suggests that peramorphosis played a central role in acipenserid evolution. Peramorphic addition and enlargement of the skeleton and scalation defines most characters at all nodes within Acipenseridae, and repudiates paedomorphosis as a major trend in evolution within the family Acipenseridae.