The evolution of the development

The evolution of development required few new features not already present in the eukaryotic cell, as exemplified by the cell cycle. Moreover, the protozoa possess many features of spatial organization and regulation present in metazoan embryos.
The earliest multicellular organism could have been reproduced by a stem cell mechanism or by fission, the latter requiring cell-to-cell interactions that may have favoured cell-interactions and regulation. Regeneration can be considered as a meta-phenomenon related to asexual reproduction and retention of embryonic characters. The origin of embryonic structures like the gastrula may be accounted for in terms of Haeckel's 'Gastrea' theory. Mechanisms based on selection at the level of cell lineage are rejected.
It is not clear what selective forces act on development itself, as distinct from the requirement for reliably producing a functional orgainsm. There is, for example, a major problem why gastrulation should be so variable in related animals. Selection for rate of development in relation to energy utilization may play a role. If many variants are neutral this may facilitate the evolution of novelty.
In general terms there is a requirement for a continuity principle for the evolution of each form in development. Most groups pass through a phylotypic stage with considerable diversity before and after.     

Shaping intraspecific variation: Development,ecology and the evolution of morphology and life history variation in tiger salamanders

The tiger salamander,Ambystoma tigrinum, is a geographically widespread, morphologically variable, polytipic species. It is among the most variable species of salamanders in morphology and life history with two larval morphs (typical and cannibal) and three adult morphs (metamorphosed, typical branchiate, cannibal branchiate) that vary in frequency between subspecies and between populations within subspecies. We report morphometric evidence suggesting that branchiate cannibals arose through intraspecific change in the onset or timing of development resulting in the wider head and hypertrophied tooth-bearing skull bones characteristic of this phenotype. We also quantified bilateral symmetry of gill raker counts and abnormalities, then evaluated fluctuating asymmetry as a measure of the developmental stability of each morph. There was a significant interaction between fluctuating asymmetry of developmental abnormalities in cannibals and typicals and the locality where they were collected, suggesting that relative stability of each phenotype could vary among populations. While altered timing of developmental events appears to have a role in the evolution and maintenance of morphs, novel phenotypes persist only under favorable ecological conditions. Predictability of the aquatic habitat, genetic variation, kinship, body size, intraspecific competition and predation all affect expression and survival of the morphs inA. tigrinum. This taxon provides an excellent model for understanding the diversity and complexity of developmental and ecological variables controlling the evolution and maintenance of novel phenotypes.     

The root of the problem: palaeoecology of distinctive crinoid attachment structures from the Silurian (Wenlock) of Gotland

The holdfast (attachment structure) is the most understudied aspect of the palaeoecology of the endoskeleton of fossil crinoids. A new collection of well-preserved holdfasts from a recently reopened quarry at Hunninge, Gotland, in Homerian (upper Wenlock) strata includes several morphologies. The most common are terminal dendritic radicular holdfasts (TDRHs) that may be derived from the cladid Ennallocrinus d'Orbigny. These have a consistent morphology of five, equally spaced, long radices that spread across the sea floor. These crinoids were gregarious, and TDRHs in a group commonly show the same radice orientations. The radices have a large axial canal compared with those of modern crinoids; each included, at least, nervous tissues. Taken together, these features suggest that, apart from attachment, these distinctive TDRHs may have served a sensory function. Other holdfasts in this assemblage also show monospecific aggregations, perhaps suggesting biochemical attraction such as that shown by certain other sessile invertebrates such as barnacles.     

Ecology and evolution of Jurassic trigoniid bivalves in Europe

 A modern palaeobiological approach to the taxonomy, making full allowance for a wide range of variation, has allowed the distinction of 22 Jurassic trigoniid species in Europe, grouped into 7 genera. Most species were confined to shallow marine habitats ranging up to a few tens of metres in depth but were adapted to low energy environments as well as environments of moderate to high energy. The presence of external ornament on the shell flanks was probably an adaptation to facilitate burrowing, but in the absence of useful external ornament other adaptations to increase burrowing efficiency may have been realized by the development of an elongate shape. Because their occurrence in Jurassic strata is only intermittent, confident inferences on evolutionary patterns are limited to the two commonest genera, Trigonia and Myophorella. Phyletic size increase has been recognized in the latter but not the former genus. The calculated species longevities correspond fairly closely with those established for Jurassic bivalve species in general. The mode of speciation is dominantly punctuational, but only one likely example of species splitting is recognized.     

Patterns of correlated character evolution in flightless birds: a phylogenetic approach

Given a robust phylogeny for a particular higher taxon, it is possible to map the evolution of various character changes onto the phylogeny and study the extent to which they co-occur. Of particular interest are the questions of (a) whether particular morphological changes tend to accompany changes in ecology or behaviour to which they bear a functional relationship and (b) whether changes in those ‘primary’ morphological characters tend to be associated with correlated changes in other aspects of morphology, as would be expected given the high level of morphological integration that characterizes most organisms. Here we report a study of this kind, looking at morphological correlates of the evolution of flightlessness in birds, and using the concentrated changes test to determine whether associations are significant. We find that pectoral reduction, pelvic enlargement and changes in skull morphology significantly co-occur, and that these are usually achieved through heterochrony rather than other kinds of developmental reprogramming. This revised version was published online in November 2006 with corrections to the Cover Date.     

Low ecological disparity in Early Cretaceous birds

Ecological divergence is thought to be coupled with evolutionary radiations, yet the strength of this coupling is unclear. When birds diversified ecologically has received much less attention than their hotly debated crown divergence time. Here, we quantify how accurately skeletal morphology can predict ecology in living and extinct birds, and show that the earliest known assemblage of birds (= pygostylians) from the Jehol Biota (≈ 125 Ma) was substantially impoverished ecologically. The Jehol avifauna has few representatives of highly preservable ecomorphs (e.g. aquatic forms) and a notable lack of ecomorphological overlap with the pterosaur assemblage (e.g. no large or aerially foraging pygostylians). Comparisons of the Jehol functional diversity with modern and subfossil avian assemblages show that taphonomic bias alone cannot explain the ecomorphological impoverishment. However, evolutionary simulations suggest that the constrained ecological diversity of the Early Cretaceous pygostylians is consistent with what is expected from a relatively young radiation. Regardless of the proximate biological explanation, the anomalously low functional diversity of the Jehol birds is evidence both for ecological vacancies in Cretaceous ecosystems, which were subsequently filled by the radiation of crown Aves, and for discordance between taxonomic richness and ecological diversity in the best-known Mesozoic ecosystem.     

Morphological and structural adaptations to soft substrates in the Early Jurassic monomyarians Lithiotis and Cochlearites

Lithiotis problematica and Cochlearites loppianus are sessile monomyarian bivalves known from the Early Jurassic of the Tethyan region, where they are found standing vertically in the calcareous mud of lagoonal fades. Their shells are characterized by a long cardinal area, ventrally displaced body space, thick attached valve, and a thin, flat free valve of equal length. A functional ligament is present only in juvenile stages of Cochlearites. To grow straight, the ventral end of the shell had to gape, so in both species it was probably the elasticity of the thin free valve that caused the shell to open. In the adult stage the shell grew only towards the venter with an apparently constant growth rate, while the soft body size remained unchanged. The outer shell is composed of a compact aragonitic layer, while the inner part is filled with loose chalky deposits which are thought to have functioned as supporting the soft body and lifting it upwards. These and other morphologic and structural features, as well as the growth pattern, can be explained as an adaptation of a sessile animal to soft muddy bottoms and rapid sedimentation. Some elongated Crassostrea living in similar environments show remarkable morphologic and structural convergence with Lithiotis and Cochlearites.     

Patterns,trends, and rates of evolution within the Actinistia

Synopsis The interrelationships of 31 actinistian species (including Latimeria chalumnae) are analyzed based on a cladistic analysis of 75 osteological characters. Inference of evolutionary trends (e.g., modification of body shape and skull morphology) from the phylogenetic patterns demonstrates that the morphology of actinistians is less conservative than has been proposed previously. This empirical cladistic approach supports two distinct tempos of evolution during an evolutionary history of 380 million years. Along a phylogenetic pathway originating with a Devonian stem-species and ending with the living Latimeria chalumnae (including 101 morphological changes and 18 cladogenetic events), the first tempo occurred during the Devonian — Permian periods as a decreasing rate of morphological changes, which was followed by a stabilizing tempo during the Permian — Recent periods. The decreasing tempo is characterized by a sequence of gradual versus quantum temporal changes and low versus faster rates, whereas the stabilizing tempo primarily is gradual and low. In contrast to a common assumption, no significant correlation was found between the rates of morphological evolution and the temporal diversity of species.     

Patterns of biotic change in Middle Jurassic to Early Cretaceous Tethyan radiolaria

The rate of taxic turnover of nearly 400 radiolarian species/subspecies is analyzed in order to document long term biotic change of plankton during the Middle Jurassic to Early Cretaceous (Aalenian to Aptian). The pattern and dynamic of diversity change is described using four indices: rate of species first and last occurrence, rate of diversification and rate of turnover. Plots of cumulative sampling effort suggest that the analyzed data represent an adequate sample of total standing diversity for most examined stages. Rates of species first occurrence exceed rates of last occurrence for most of the Middle Jurassic, except for the middle Bajocian. In contrast, the Late Jurassic was a time of decreasing radiolarian diversity and the Kimmeridgian records the lowest rate of diversification. It is followed by a dramatic increase in first occurrences near the Jurassic–Cretaceous boundary with as a result the highest rate of diversification recorded in the late Tithonian. Regional radiolarian diversity was stable throughout most of the Early Cretaceous. A stratigraphic permutation test was performed to assess the influence of uneven sampling on the observed pattern of taxic turnover and identified the intervals for which randomly obtained patterns are significantly different from the observed pattern. The Kimmeridgian and late Tithonian events coincide with substantial climate-derived perturbations in water cycling, nutrient supply and oceanic productivity. They point to a negative relationship between radiolarian macroevolution and changes in the state of nutrient availability, although further work is needed to refine the temporal resolution of this relationship and to explore ecological aspects of its causal link with respect to radiolarian evolution.     

Patterns and processes of heterochrony in Mesozoic goniasterid sea-stars

Heterochronic processes are obvious in the most common family of Mesozoic asteroids, the Goniasteridae, whether studied from entire fossils or, as is more frequently the case, from isolated ossicles. When phylogeny is poorly known, comparison between species leads to such unsatisfactory interpretations as 'A is more paedomorphic than B' or 'C, compared with other species, has many peramorphic characters'. When the phylogeny is known, heterochronic trends appear amongst the lineages. The peramorphocline Metopaster parkinsoni - M. loirensis-M. trichilae-M. chilipora-M. hypertelicus is studied here from the Cenomanian to the Upper Campanian of France. Different morphological characters evolved at different speeds. Among each polymorphous population, some morphotypes foreshadowed the next step. Juveniles of one species look like the adults of one of the ascendants. The cline developed irrespective of the substrate, and proceeded together with a migration from the Anglo-Paris Basin to the Aquitanian Basin. Metopaster hunteri , during the Coniacian, and Metopaster meudonensis , during the Upper Campanian, both evolved from Metopaster parkinsoni by neoteny, and provide a good example of reiterated and canalized morphological evolution. Some non-heterochronic canalizations are described.     

Morphogenetic evolution of hydroid colony pattern

A scheme of evolution of hydrozoan colony pattern is proposed based upon the consideration of macro-morphogenesis. Four main processes play decisive roles(1) hard skeleton formation by soft tissues, (2) changes in duration of the growth phase relative to the transition to differentiation in interdependent zones of growth, (3) ratio in growth rates between adjacent zones of growth within the rudiment, the shoot, or the whole colony, and (4) spatial relationships among growth zones. The main tendency in morphological evolution of the hydroids is an increasing integration of the colony as revealed by increasing complexity of its structure. That is from a temporary colony towards the permanent one with highly organised shoots, as hydranths and branches are localised in a strictly arranged manner. An analysis of diverse data allows one to state that the main morphogenetic mechanism of increasing complexity in the hydroid colony is convergence, then fusion, of adjacent growth zones, a variant of heterochrony.     

Functional evolutionary developmental biology (evo-devo) of morphological novelties in plants

The origin of morphological and ecological novelties is a long-standing problem in evolutionary biology.Understanding these processes requires investigation from both the development and evolution standpoints,which promotes a new research field called evolutionary developmental biology (evo-devo).The fundamental mechanism for the origin of a novel structure may involve heterotopy,heterochrony,ectopic expression,or loss of an existing regulatory factor.Accordingly,the morphological and ecological traits cont...     

The functional morphology and palaeoecology of the Dinantian brachiopod Levitusia

The functional morphology of Levitusia , based principally upon the type-species L. humerosa (J. Sowerby), is described and compared with living brachiopods. It is concluded that Levitusia species lived in tropical Dinantian seas with agitated waters and at shallow depths, perhaps no more than 50 m. The lithologies in which specimens are found support this view and indicate the need for some revision in our interpretations of some Dinantian sedimentary environments. Further evidence for the synonymy of L. christiani (de Koninck) in L. humerosa is presented.