Evolution Within a Bizarre Phylum: Homologies of the First Echinoderms

SYNOPSIS. The Extraxial/Axial Theory (EAT) of echinoderm skeletalhomologies describes two major body wall types: axial and extraxial.The latter is subdivided into perforate and imperforate regions.Each of the regions has a distinctly different source in earlylarval development. Axial skeleton originates in the rudiment,and develops in association with the pentaradially arrangedhydrocoel according to specific ontogenetic principles. Perforateand imperforate extraxial regions are associated with the leftand right somatocoels respectively, are not governed by ontogeneticprinciples of plate addition, and are products of the non-rudimentpart of the larval body. The morphology of even the most bizarreof the earliest echinoderms can be explored using the EAT. Amongthese, edrioasteroid-like taxa best fit the idea that formsexpressing archimery in the sequential arrangement of axial,perforate extraxial, and imperforate extraxial regions are thefirst echinoderms. Metamorphosis is especially marked in cladesthat have a high axial to extraxial skeleton ratio because structuresdeveloping from the non-rudiment part are suppressed in favorof the developing axial elements during this process. However,inearly echinoderms, extraxial skeleton makes up a far largerproportion of the body wall than axial, implying that metamorphosiswas not as significant a part of the developmental trajectoryas it is in more recently evolved taxa. Echinoderm radiationconsists of a succession of apomorphies that reduced the expressionof extraxial components but increased the influence of axialones, with a concomitant increase in the prominence of metamorphosis.     

Microgeographical variation in shell size of the land snail Chondrina clienta

Microgeographical variation in shell morphology of the rock-dwelling land snail Chondrina clienta , collected from 30 sites within an area of 0.5 km² on the island of Öland, Sweden, was examined in relation to its own population density and that of a potential competitor (the land snail Balea perversa ) and to environmental factors. Dispersal of marked individuals averaged 83 cm per year within a stone pile and 291 cm in an area of exposed bedrock. Local population density of active C. clienta ranged from 5 to 794 individuals per m². Shell characters were highly intercorrelated, both within and between populations. Principal component analysis revealed that most of the interpopulational variation could be expressed by the single character of shell height, which ranged from 5.54 to 6.94 mm. In all populations, snails of a given size had the same whorl number. Shell size was not influenced by habitat type (exposed rock surface, stone pile or stone wall) or proportion of calcareous stones within habitat. It was, however, negatively correlated with local population density, indicating intraspecific competition, and positively correlated with the degree of plant cover within the habitat. Analysis of variance revealed additionally a density effect of B. perversa on shell size in C. clienta , probably as a result of interspecific competition. Breeding experiments using C. clienta from different sites and carried out under unifrom conditions caused most of the phenotypic variation to disappear, demonstrating the high phenotypic plasticity of the species.     

Environmental features influencing Carabid beetle (Coleoptera) assemblages along a recently deglaciated area in the Alpine region

Abstract.  1. The spatio-temporal approach was used to evaluate the environmental features influencing carabid beetle assemblages along a chronosequence of an Italian Alpine glacier foreland. The influence of environmental variables on species richness, morphology (wing and body length), and distribution along the chronosequence was tested.
2. Species richness was found to be a poor indicator of habitat due to weak influences by environmental variables. It seems that the neighbouring habitats of a glacier foreland are not able to determine significant changes in carabid species richness.
3. Instead it appears that history (age since deglaciation) and habitat architecture of a glacier foreland are strongly correlated to species adaptive morphological traits, such as wing morphology and body length. Assemblages characterised by species with reduced wing size are linked to the older stages of the chronosequence, where habitat is more structured. Assemblages characterised by the largest species are linked to the younger sites near the glacier. These morphological differentiations are explained in detail.
4. Habitat age can therefore be considered the main force determining assemblage composition. On the basis of the relationship between morphological traits and environmental variables, it seems likely that age since deglaciation is the main variable influencing habitat structure (primary effect) on the Forni foreland. The strong relationship between carabid assemblages and habitat type indicates that site age has but a secondary effect on carabid assemblages. This may be utilised to interpret potential changes in assemblages linked to future glacier retreat.     

Effects of small‐scale grassland fragmentation and frequent mowing on population density and species diversity of orthopterans: a long‐term study

Abstract 1. Habitat fragmentation is considered one of the major threats to invertebrate diversity in semi‐natural grassland. However, the effects of habitat fragmentation through mowing on the rich insect fauna of these grasslands have not been sufficiently investigated and experiments are especially rare. 2. We studied the impact of small‐scale grassland fragmentation on orthopterans over 7 years in an experiment which allowed us to additionally investigate the effect of frequent mowing on Orthoptera communities. 3. Overall, Orthoptera density and species richness increased over time. This was likely a result of increased small‐scale habitat heterogeneity and the provision of a short‐turf habitat suitable for xerophilous species. The fragmentation affected orthopteran density and species composition but not species richness whose response lagged behind the changes in abundance. 4. Responses differed between suborders. Ensifera density was higher in fragment than in control plots. Caelifera density did not differ between fragment and control plots. The mown matrix was an unsuitable habitat for most of the species, particularly within the Ensifera. 5. Our experiment shows that even small‐scale fragmentation can affect Orthopteran communities and that the effects became more pronounced over time. As the mown matrix was unsuitable for many Ensifera species, they may go locally extinct when large areas are mown simultaneously.     

Are myrmecophytes always better protected against herbivores than other plants?

The present field study compared the degree of defoliation of three Guianian melastome, two myrmecophytes (i.e. plants sheltering ants in hollow structures) and Clidemia sp., a nonmyrmecophytic plant serving as a control. Maieta guianensis Aubl. hosted mostly Pheidole minutula Mayr whatever the area, whereas Tococa guianensis Aubl. hosted mostly Azteca bequaerti Wheeler along streams and Crematogaster laevis Mayr or Azteca sp. 1 in the understory where it never blossomed. Only Tococa , when sheltering A. bequaerti in what can be considered as a truly mutualistic relationship, showed significantly less defoliation than control plants. In the other associations, the difference was not significant, but P. minutula is mutualistic with Maieta because it furnishes some protection (exclusion experiments) plus nutrients (previous studies). When devoid of ants, Tococa showed significantly greater defoliation than control plants; therefore, it was deduced that Tococa probably lacks certain antidefoliator metabolites that control plants possess (both Tococa and control plants are protected by ground-nesting, plant-foraging ants, which is termed 'general myrmecological protection'). Consequently, plant-ants other than A. bequaerti probably also protect Tococa slightly, thus compensating for this deficiency and permitting it to live in the understory until treefall gaps provide the conditions necessary for seed production.  © 2006 The Linnean Society of London, Biological Journal of the Linnean Society , 2006, 89 , 91–98.