Évolution et radiations adaptatives chez les échinides

Evolution and adaptative radiations in echinoids. Echinoids (sea urchins) originated in the Early Paleozoic and, after a first period of diversification during that era, participated in an intense evolutionary radiation during the Mesozoic and Cenozoic. In the context of a reconsideration of homologies for echinoderm body regions, we present several major aspects of echinoid evolution: the origins of irregularity and bilateral symmetry; a model of radiation exemplified by the spatangoids; the diversification of modes of echinoid reproduction. These examples allow us to understand the extent of and capacity for innovation demonstrated by sea urchins during their post-Paleozoic radiation.     

How do changes in parental investment influence development in echinoid echinoderms?

SUMMARY Understanding the relationship between egg size, development time, and juvenile size is critical to explaining patterns of life-history evolution in marine invertebrates. Currently there is conflicting information about the effects of changes in egg size on the life histories of echinoid echinoderms. We sought to resolve this conflict by manipulating egg size and food level during the development of two planktotrophic echinoid echinoderms: the green sea urchin, Strongylocentrotus droebachiensis and the sand dollar, Echinarachnius parma . Based on comparative datasets, we predicted that decreasing food availability and egg size would increase development time and reduce juvenile size. To test our prediction, blastomere separations were performed in both species at the two-cell stage to reduce egg volume by 50%, producing whole- and half-size larvae that were reared to metamorphosis under high or low food levels. Upon settlement, age at metamorphosis, juvenile size, spine number, and spine length were measured. As predicted, reducing egg size and food availability significantly increased age at metamorphosis and reduced juvenile quality. Along with previous egg size manipulations in other echinoids, this study suggests that the relationship between egg size, development time, and juvenile size is strongly dependent upon the initial size of the egg.     

Patterns of species diversity and endemism in comparable temperate brown algal floras

Brown algal species diversity is compared in 100 km sections of the coastlines of four warm temperate regions: southern Australia, California, southwestern Africa, north-central Chile. The highest diversity (over 140 species per section) is found in southern Australia. California has a reasonable diversity (around 70 species per section), and both southern Australia and California have high regional endemism. Sections of north-central Chile and southwestern Africa have similar patterns, with low diversity (< 30 species per 100 km section), low endemism, few or no fucoids, and up to 25% of the brown algal flora are environmentally tolerant species of Scytosiphonales. Species turnover between contiguous sections of coast is generally related to relative change in temperature regime. Thus the high diversity of southern Australia is due to high species diversity within the 100 km sections, with little turnover, except for a rapid reduction in eastern Victoria likely to be related to lack of rocky substatum. It is hypothesized that low diversity and endemism in Chile and southwestern Africa can be explained by the occurrence of major environmental perturbations (upwelling and El Nino effects) in these regions, producing variable inter-annual temperature conditions that select out tolerant species from the local floras.     

The origin of echinoid shell beds in siliciclastic shelf environments: three examples from the Miocene of Sardinia,Italy

Three mass accumulations of sea urchins from the Miocene of Sardinia show a number of taphonomic features which set them apart from previously described echinoid assemblages from the Cenozoic in which they represent: (1) monotypic assemblages; (2) include very well‐preserved remains of either regular or spatangoid echinoids; and (3) originate in deeper water environments. These accumulations are compared using a detailed sedimentological and taphonomic analysis including preservational fabrics, taphonomic signatures, size frequency distributions, density of occurrences and preferred orientations. The possible role of gregarious behaviour contributing to mass occurrences and the specific sedimentary events leading to the excellent preservation are discussed. The interpreted depositional environment of all three deposits is that of a storm‐dominated, siliciclastic shelf environment. A phymosomatid assemblage represents rapid burial through obrution of a highly dense, freshly dead community. A Brissopsis‐dominated spatangoid assemblage represents a mixed accumulation of parautochthonous and transported skeletons. The third assemblage consisting of regular echinoid spines and rare tests represents a composite tempestite. Differences in the depositional environments are related to their position along onshore–offshore gradient with the first two beds originated in a deeper setting than that of the spine accumulation. This study shows that the preservation of assemblages containing complete regular echinoids and spatangoids is higher in deeper water settings than in shallow water environments.     

Evolutionary dynamics at high latitudes: speciation and extinction in polar marine faunas

Ecologists have long been fascinated by the flora and fauna of extreme environments. Physiological studies have revealed the extent to which lifestyle is constrained by low temperature but there is as yet no consensus on why the diversity of polar assemblages is so much lower than many tropical assemblages. The evolution of marine faunas at high latitudes has been influenced strongly by oceanic cooling during the Cenozoic and the associated onset of continental glaciations. Glaciation eradicated many shallow-water habitats, especially in the Southern Hemisphere, and the cooling has led to widespread extinction in some groups. While environmental conditions at glacial maxima would have been very different from those existing today, fossil evidence indicates that some lineages extend back well into the Cenozoic. Oscillations of the ice-sheet on Milankovitch frequencies will have periodically eradicated and exposed continental shelf habitat, and a full understanding of evolutionary dynamics at high latitude requires better knowledge of the links between the faunas of the shelf, slope and deep-sea. Molecular techniques to produce phylogenies, coupled with further palaeontological work to root these phylogenies in time, will be essential to further progress.     

Maternal energy investment in eggs and jelly coats surrounding eggs of the echinoid Arbacia punctulata

In free-spawning marine invertebrates, the amount of maternal energy that is invested in each egg has profound implications for all life-history stages of the offspring. The eggs of echinoids are freely spawned into the water and are surrounded by several structurally complex extracellular layers. These extracellular layers, or jelly coats, do not contribute energy to embryonic development but must impose an energy cost on the production of each egg. The investment of maternal energy reserves in the jelly coats of echinoid eggs may have important implications for the number of eggs that can be produced (i.e., fecundity) and the amount of energy that can be invested in each egg. We estimated the degree to which maternal energy is invested in the jelly coats surrounding eggs of the echinoid Arbacia punctulata. Estimates were derived from measurements of the amount of energy contained in the combined eggs and jelly coats, and in the eggs alone. The amount of energy contained in A. punctulata eggs ranged from 2.70 to 5.53 x 10(-4) J egg(-1). The amount of energy contained in the jelly coats ranged from 0.13 to 0.48 x 10(-4) J jelly coat(-1). The mean concentration of energy in the eggs was 2.15 mm(-3) and 0.29 J mm(-3) in the jelly coats. These results indicate that between 3% and 11% (mean = 7%) of the total energy invested in each A. punctulata egg is partitioned to the jelly coat alone. A significant positive relationship was found between the volumes of the jelly coats and the amount of energy they contained. Based on this relationship and an analysis of differences in the size of jelly coats between echinoid species, we suggest that the degree to which energy is invested in jelly coats may vary among echinoid species and is therefore likely to be an important life-history characteristic of these organisms.     

Echinoids,epizootics and ecological stability in the rocky subtidal off Nova Scotia,Canada

Mass mortalities ofStrongylocentrotus droebachiensis, attributed to disease, have occurred along the Atlantic coast of Nova Scotia from 1980 to 1982. An amoeboid protist has been tentatively identified as the pathogenic agent. Temperature appears to play an important role in triggering epizootics and determining the extent of mortality. Epizootics have occurred in the autumn of each year during months of record high temperatures. The virtual elimination of echinoids over a large area of coast has enabled regeneration of kelp beds. Disease may play a key role in determining the structure and stability of benthic communities off Nova Scotia and elsewhere by controlling echinoid abundance.     

The role of echinoids in shaping environments

The role of echinoids in shaping the environment of marine and terrestrial biota is considered. The participation of echinoids in bioerosion, bioturbation of sediments, and their relationships with symbionts, parasites and predators, as well as the role of echinoid skeletons in rock formation, are discussed.     

Shallow Traces (Pits) in the Test of the Irregular Echinoid Echinocorys scutata Leske from the Chalk (Upper Cretaceous) of the United Kingdom

Specimens of epifaunal irregular echinoids in the Upper Cretaceous of northern Europe have been reported with patterns of circular, nonpenetrative parabolic pits, Oichnus paraboloides (Bromley), in the apical region. Specimens of Echinocorys scutata Leske from the Chalk at two sites in southeast England were commonly penetrated by this trace, generated by an indeterminate pit-forming organism. Pits commonly surround the apical system and, less commonly, occur within it; they occur preferentially anteriorly. Pits occur within plates, not along margins or sutures. Crosscutting of pits indicates that multiple spatfalls probably occurred. The host echinoid added new test plates adjacent to the apical system; thus, plates bearing O. paraboloides were moved abapically. The reduction in number of pits away from the apex, including those with echinoid tubercles reestablished on the base, indicates that, following death of an infester, the echinoid “reclaimed” and infilled them with calcite. The pit-former was most probably an unmineralized invertebrate that used E. scutata as a domicile which provided good access to food-rich currents for suspension feeding. Although the systematic position of the pit-former is unknown, similar infestations are known from other Upper Cretaceous echinoids and Mississippian crinoids.     

Antarctic, Sub-Antarctic and cold temperate echinoid database

This database includes spatial data of Antarctic, Sub-Antarctic and cold temperate echinoid distribution (Echinodermata: Echinoidea) collected during many oceanographic campaigns led in the Southern Hemisphere from 1872 to 2010. The dataset lists occurrence data of echinoid distribution south of 35°S latitude, together with information on taxonomy (from species to genus level), sampling sources (cruise ID, sampling dates, ship names) and sampling sites (geographic coordinates and depth). Echinoid occurrence data were compiled from the Antarctic Echinoid Database (David et al. 2005a), which integrates records from oceanographic cruises led in the Southern Ocean until 2003. This database has been upgraded to take into account data from oceanographic cruises led after 2003. The dataset now reaches a total of 6160 occurrence data that have been checked for systematics reliability and consistency. It constitutes today the most complete database on Antarctic and Sub-Antarctic echinoids.     

Regional versus latitudinal variation in the life-history traits and demographic rates of a reef fish,Centropyge bispinosa,in the Coral Sea and Great Barrier Reef Marine Parks,Australia

Environmental temperature is an important determinant of physiological processes and life histories in ectotherms. Over latitudinal scales, variation in temperature has been linked to changes in life-history traits and demographic rates, with growth and mortality rates generally being greatest at low latitudes, and longevity and maximum length being greater at higher latitudes. Using the two-spined angelfish, Centropyge bispinosa, as our focal species, we compared growth patterns, growth rates, longevity, mortality, asymptotic length and maximum length across 22 reefs that span 13° of latitude within the Great Barrier Reef Marine Park (GBRMP) and the Coral Sea Marine Park (CSMP), Australia. We found no predictable latitudinal variation in mortality rates, growth patterns, growth rates, asymptotic or maximum length of C. bispinosa at regional to biogeographic scales. However, C. bispinosa consistently exhibited reduced longevity at lower, warmer latitudes within the CSMP. The greatest differences in mean maximum length of C. bispinosa were between continental (GBRMP) and oceanic (central CSMP) reefs of similar latitude, with individuals being larger on average on continental versus oceanic reefs. The lack of predictable life-history and demographic variation in C. bispinosa across a 13° latitudinal gradient within the CSMP, coupled with differences in mean maximum length between continental and oceanic reefs at similar latitudes, suggest that local environmental conditions have a greater influence than environmental temperature on the demographic rates and life-history traits of C. bispinosa.     

Loriolella,a key taxon for understanding the early evolution of irregular echinoids

Loriolella Fucini is an enigmatic and poorly understood Early Jurassic sea-urchin that has been interpreted both as a regular and an irregular echinoid. The discovery of new and well-preserved material of the type species L. ludovicii (Meneghini) from the Upper Pliensbachian (Lower Jurassic) of Lombardy, Italy, has clarified the morphology of this genus for the first time. Loriolella resembles regular echinoids in having a large (almost certainly monocyclic) apical disc enclosing the periproct, and large primary interambulacral tubercles and spines. However, its peristome is extremely small with only vestigial buccal notches and auricles, and the ambulacra expand adorally forming cassiduloid-like phyllodes. Cladistic analysis suggests that Loriolella is the earliest known microstomatid irregular echinoid. Its unique mixture of primitive and derived characters is important for interpreting the initial steps by which irregular echinoids arose from diadematoid-like regular ancestors.     

IRREGULAR ECHINOIDS—AN INSUFFICIENTLY KNOWN GROUP

Some aspects of echinoid evolution from an endocyclic towards an exocyclic apical system, in the light of embryological and palaeontological data, are discussed. A primitive feature—imbrication of interamhulacral plates in Jurassic irregular echinoids, recently described—proves once more that this group is insufficiently known. Every revision of taxonomic or phylogenetic concepts concerning this group of echinoids should be preceded by careful morphological analysis of the material.     

Echinoid grazing as a structuring force in coral communities: Whole reef manipulations

To examine the effects of echinoid grazing on shallow water coral community structure, the entire Diadema antillarum Philippi population (> 3000 individuals) was eliminated from a patch reef in Discovery Bay, Jamaica W.I. in January 1974. All remaining regular echinoids, notably including Echinometra viridis A. Agassiz (> 7000 individuals), were removed from one-half of the same reef 6 months later.Where Echinometra was present, its grazing was highly patchy, even at densities of up to 50/m². On the other hand, the effects of grazing by Diadema were more uniformly distributed at equivalent or lower densities.Percent-cover of adult corals was enhanced in the presence of Echinometra viridis, however, it was greatly reduced in the absence of all echinoid grazing as a result of massive overgrowth by algae. Success of coral recruitment (density of coral spat) increased in the absence of Diadema due to the alleviation of biological disturbance imposed by this urchin's feeding activities. The presence of Echinometra viridis, even at the high densities naturally present on the reef, did not deter coral settlement.Coral diversity was highest in the presence of all echinoids, intermediate in the presence of E. viridis (without Diadema), and lowest in the absence of all echinoids. This was due to the disproportionately successful recruitment and resultant high relative abundance of Agaricia at low echinoid densities. Favia, Millepora, Helioseris and Acropora all exhibited universally low recruitment levels. Predation by Diadema and Echinometra is responsible for damaging larger coral colonies, particularly Porites.As echinoid species composition and density was altered, the species composition of benthic algae and associated invertebrates shifted. In this way, the array of species competing for space with coral spat shifted as did their competitive relationships. Major epifaunal competitors included spirorbid polychaetes, various foraminifera (particularly Gypsina), and sponges. Algal competitors included filamentous chlorophytes, corallines (e.g. Jania), and numerous other red algae; the species composition of algal competitors shifted through time. Competitive success in corals varied as echinoid density changed and was species-specific with respect to coral genus. Interspecific competition between coral spat (i.e., interspecific aggression via extracoelenteric digestion) was negligible under all treatments.The presence of Echinometra viridis in high densities (in the absence of Diadema) created conditions where growth and fusion in Agaricia spat were optimized. Under these conditions, fusion allows a young coral colony to attain a large enough size to survive damage incurred from grazing or from competitive overgrowth, decreasing the probability of whole-colony mortality during the early stages of development.     

Hierarchical metapopulation structure in a highly mobile marine predator: the southern Australian coastal bottlenose dolphin (<Emphasis Type="Italic">Tursiops</Emphasis> cf. <Emphasis Type="Italic">australis</Emphasis>)

Little is known about the population ecology of the recently described bottlenose dolphin species Tursiops australis. The classification of this species is still under debate, but this putative species is thought to be comprised of small and genetically distinct populations (including sub-populations under increasing anthropogenic threats) and is likely endemic to coastal southern Australia. Mitochondrial DNA (mtDNA) control region sequences and microsatellite loci were used to assess genetic variation and hierarchical population structure of coastal T. cf. australis across a range of spatial scales and environmental discontinuities between southern Western Australia (WA) and central South Australia (SA). Overall, genetic diversity was similar to that typically found for bottlenose dolphins, although very low mtDNA diversity was found in Gulf St. Vincent (GSV) dolphins. We found historical genetic subdivision and likely differences in colonisation between GSV and Spencer Gulf, outer- and inner-gulf locations, and SA/WA and previously identified Victorian/Tasmanian populations. A hierarchical metapopulation structure was revealed along southern Australia, with at least six genetic populations occurring between Esperance, WA and southern Tasmania. In addition, fine-scale genetic subdivision was observed within each SA/WA population. In general, contemporary migration was limited throughout southern Australia, but an important gene flow pathway was identified eastward along the Great Australian Bight. Management strategies that promote gene flow among populations should be implemented to assist with the maintenance of the inferred metapopulation structure. Further research into the population ecology of this species is needed to facilitate well-informed management decisions.     

Completeness of a fossil record: the Pleistocene echinoids of the Antilles

Diversity of Pleistocene marine invertebrates with single component, robust, easily identifiable skeletons shows a good correspondence to that of the Recent when compared between similar environments in the same region. Such comparisons are less easily made for multi-element skeletons. In the Antilles there are 24 extant species of echinoids in 0–30  m water depth. Considering all available fossils - complete tests, disarticulated spines, and test fragments and ossicles - the Pleistocene echinoids of the Antillean region have a high similarity with the extant shallow water fauna (c. 63% specific and 72% generic similarity). Many of the extant, shallow water, regular echinoid taxa are known from the Pleistocene, although irregular echinoids are less well represented. These results are comparable with those determined for coeval benthic invertebrates from this and other geographic regions, and indicate a high degree of similarity from a survey which is so far limited to only a few of the islands. Not all Pleistocene units considered herein were deposited in shallow water, but they include autochthonous specimens of taxa whose depth range extends beyond shallow water at the present day or presumed allochthonous specimens derived by downslope transport from shallower water, or both.     

Effects of capability for dispersal on the evolution of diversity in antarctic benthos

The likelihood of marine invertebrates to maintain large geographic ranges is widely dependent on the ability of their early ontogenetic stages to disperse over long distances. Marine benthic invertebrates inhabiting the cold-stenothermal environment of the Southern Ocean are known for their overall reduced number of pelagic larvae, or drifting stages of any kind, when compared with organisms elsewhere in the sea. The diversity of organisms thriving in Antarctic waters is the result of evolution in situ and of the intrusion of species from surrounding seas. The reasons for a high level of endemism and a stunning diversity of benthic invertebrates found today are frequently discussed in the literature, but the mechanisms whereby diversity has been controlled over time remain largely theoretical. Here, I suggest that, indeed, early life-history patterns play a key role in defining the radiation and the speciation potential of Antarctic benthic invertebrates. In arguing this case, I synthesize the growing body of molecular studies on population connectivity in Antarctic benthic invertebrates, and compare this information with knowledge of their life histories and biogeography. I conclude that differences in early life-history patterns are key to the resilience potential of species in response to late Cenozoic glacial periods and propose that there is a direct relationship between rate of speciation and the ability of taxa to disperse.