Developmental plasticity of limb bone microstructural organization in Apateon: histological evidence of paedomorphic conditions in branchiosaurs

SUMMARY Apateon, a key genus among Branchiosauridae from the Carboniferous—Permian of Europe, is often considered closely related to salamanders on the basis of developmental similarities, anatomical features, and life history. The current work deals with recognition of heterochronies among three “time‐averaged populations” of Apateon based on inference from histological features already studied in extant urodeles. Our study is performed on the long bones of 22 specimens of Apateon pedestris and Apateon caducus. Histological observations show that diaphyseal and epiphyseal ossification patterns of Apateon are similar to those of urodeles. From skeletochronological analysis, the identification of the age of sexual maturity allows us to discriminate juveniles from adults and to confirm the previous hypothesis of a paedomorphic condition based on anatomical data among these species. The current study also suggests a paedomorphic condition of each “population” at the histological level. This heterochrony may have been linked to peculiar ecological conditions such as hypoxic and fresh water environment. Functional reasons may also be invoked to explain differences of ossification between fore‐ and hindlimbs of the “populations” from Odernheim and Niederkirchen because paleoecological conditions are very different from one locality to another. This study illustrates the role that the acquisition of heterochronic features plays at the microevolutionary scale.     

The effect of habitat on modern shark diversification

Sharks occupy marine habitats ranging from shallow, inshore environments to pelagic, and deepwaters, and thus provide a model system for testing how gross habitat differences have shaped vertebrate macroevolution. Palaeontological studies have shown that onshore lineages diversify more quickly than offshore taxa. Among onshore habitats, coral reef‐association has been shown to increase speciation rates in several groups of fishes and invertebrates. In this study, we investigated whether speciation rates are habitat dependent by generating the first comprehensive molecular timescale for shark divergence. Using phylogenetic comparative methods, we rejected the hypothesis that shelf (i.e. onshore) lineages have higher speciation rates compared to those occupying deepwater and oceanic (i.e. offshore) habitats. Our results, however, support the hypothesis of increased speciation rates in coral reef‐associated lineages within the Carcharhinidae. Our new timetree suggests that the two major shark lineages leading to the extant shark diversity began diversifying mostly after the end‐Permian mass extinction: the squalimorphs into deepwater and the galeomorphs into shelf habitats. We suggest that the breakdown of the onshore–offshore speciation rate pattern in sharks is mediated by success in deepwater environments through ecological partitioning, and in some cases, the evolution of morphological novelty.     

Allometric shape change and heterochrony in the freeliving coral Trachyphyllia bilobata (Duncan)

Regression analysis has been used to study the relationship between age, size, shape, and surface area in two ancestral-descendant populations of the Neogene Caribbean coral Trachyphyllia bilobata. Analyses of the relationship between size and age show that the relationship is isometric and that little difference occurs between populations in mean corallite length or height and in their rates of growth. Onset of columella growth is significantly earlier, however, in the descendant population. Studies of the relationship between size and shape show that growth is allometric, with shape change occurring in both corallum elongation and pinching of the corallite wall during ontogeny. In the descendant population, pinching and elongation initiate earlier in the ontogeny of the coral. These results suggest that the evolutionary development of the meandroid form in freeliving corals has been accomplished by heterochrony, involving a complex set of disassociated peramorphic changes in ontogeny accompanied by paedomorphic changes in astogeny. Further analyses show that the observed heterochronic changes serve to decrease corallum surface area which may in turn enhance sediment removal and nutrition in unstable habitats.     

Genetic structure of <Emphasis Type="Italic">Saxifraga tridactylites</Emphasis> (Saxifragaceae) from natural and man-made habitats

In contrast to many declining plant species Saxifraga tridactylites extended its distribution range in the man-made landscape of central Europe. The species naturally colonizes rocks and calcareous grasslands, but has also spread enormously in anthropogenic habitats such as railway constructions during the last decades. To analyze the genetic structure of the species 216 individuals from 8 populations in natural and 5 populations in man-made habitats were studied using AFLPs. The molecular analysis resulted in 250 scorable fragments. Population variability, measured as Nei’s gene diversity, Shannon’s Information Index and percentage of polymorphic bands, was slightly but not significantly higher in populations from natural habitats and was not correlated with population size. Mantel test indicated no significant correlation between pairwise genetic (Φ_PT) and geographic distances. An analysis of molecular variance revealed significant differentiation between the two habitat types. About 9% variability were observed between natural and man-made habitats, 21% among populations within these two habitats and 70% within populations. In a neighbour joining cluster analysis populations from natural and man-made habitats were clearly separated from each other. Populations of S. tridactylites from man-made habitats do, therefore, not suffer from reduced genetic diversity. The observed genetic differentiation between populations from man-made and natural habitats could be due to reduced gene flow and/or habitat specific selection. However, the results of the study clearly demonstrate human impact on the genetic structure of plant populations in man-made landscapes.     

Larger foraminifera and sedimentation around Fongafale Island,Funafuti Atoll,Tuvalu

Larger foraminifera are an important component of coastal sediments around Fongafale Island, Funafuti Atoll, Tuvalu, and at least 10 species are present. In the shallow lagoon, foraminifera (mainly Amphistegina lessonii, A. lobifera, Baculogypsina sphaerulata, Calcarina spengleri, Marginopora vertebralis, and Sorites marginalis) are the dominant component of sand and gravel, followed in decreasing order of abundance by calcareous red and green algae, coral, and molluscs. In deeper water, Halimeda replaces the foraminifera. Close inshore, abrasion removes Halimeda and may reduce the number of foraminiferal tests. There is some sediment movement in both onshore and offshore directions although offshore transport appears minor. On land, dissolution that preferentially removes aragonite may increase the proportion of foraminiferal tests to as much as 83% of the subsurface sediment. Sediments on the ocean side are dominated by coral and coralline red algal debris thrown up in 1972 by cyclone Bebe and later moved inshore and lagoonward.Communicated by P.K. Swart     

Onshore–offshore gradient in metacommunity turnover emerges only over macroevolutionary time-scales

Invertebrate lineages tend to originate and become extinct at a higher rate in onshore than in offshore habitats over long temporal durations (more than 10 Myr), but it remains unclear whether this pattern scales down to durations of stages (less than 5 Myr) or even sequences (less than 0.5 Myr). We assess whether onshore–offshore gradients in long-term turnover between the tropical Eocene and the warm-temperate Plio-Pleistocene can be extrapolated from gradients in short-term turnover, using abundances of molluscan species from bulk samples in the northeast Atlantic Province. We find that temporal turnover of metacommunities does not significantly decline with depth over short durations (less than 5 Myr), but significantly declines with depth between the Eocene and Plio-Pleistocene (approx. 50 Myr). This decline is determined by a higher onshore extinction of Eocene genera and families, by a higher onshore variability in abundances of genera and families, and by an onshore expansion of genera and families that were frequent offshore in the Eocene. Onshore–offshore decline in turnover thus emerges only over long temporal durations. We suggest that this emergence is triggered by abrupt and spatially extensive climatic or oceanographic perturbations that occurred between the Eocene and Plio-Pleistocene. Plio-Pleistocene metacommunities show a high proportion of bathymetric generalists, in contrast to Eocene metacommunities. Accordingly, the net cooling and weaker thermal gradients may have allowed offshore specialists to expand into onshore habitats and maintain their presence in offshore habitats.     

Life-history variation in Crepis tectorum (Asteraceae)

Summary Populations of the monocarpic plant Crepis tectorum were grown in a series of uniform environments to test the hypothesis that weedy populations are more r-selected than populations from a more natural habitat. Weedy populations exhibited a combination of r- and K-selected traits. The relatively rapid growth, the potential for a summer annual habit, and the relatively high fecundity that characterized at least one of the two weed populations studied were considered as r-selected traits favored in habitats of unpredictable duration. However, high levels of competition from other weedy species or from the crop in arable fields may explain at least some presumably K-selected traits observed in the weedy populations, e.g. relatively large seeds and late flowering in the summer. Results indicated that stress due to abiotic factors (strong winds, desiccation and nutrient deficiency) has been a more important selective factor than r- or K-selection, in non-weedy populations from calcareous grasslands (alvars) on the Baltic islands.     

Observations on the composition and larval developmental modes of polydorid pests of farmed oysters (Crassostrea gigas) and abalone (Haliotis midae) in South Africa

This study provides preliminary insights into how the culture method of molluscs and the larval developmental mode of pest polydorids may determine which species become pests. A total of 10 species were identified on oysters and abalone in offshore and oysters in onshore culture on the west coast of South Africa. One-factor PERMANOVA showed that abalone and oysters cultured offshore housed similar communities of worms, but that they differed significantly from the communities housed by oysters cultured onshore. Oysters from onshore were infested mainly by Polydora hoplura, but also by Polydora cf. websteri and Boccardia pseudonatrix. All these species produce adelphophagic larvae or a mixture of these and planktotrophic larvae. Adelphophagic larvae usually only emerge soon before they settle, which would probably enhance their ability to recruit locally and establish local populations. By contrast, worms from molluscs grown offshore were infested by high abundances of P. hoplura and lower abundances of up to five more species per individual host. All hatched broods observed with these female P. hoplura contained planktotrophic larvae which have a long planktonic phase before settling. This would presumably enhance their ability to reach hosts in the offshore.     

Nutritional differentiation among populations of the mediterranean shrub Dittrichia viscosa (Asteraceae) in siliceous and calcareous habitats

Summary Dittrichia (ex Inula) viscosa is a ruderal species that has recently become an invading plant in the northwest Mediterranean basin. A previous study failed to demonstrate the occurrence of morphologically differentiated ecotypes among populations of the species but suggested the existence of nutritional ecotypes. This latter possibility is examined here by comparing the ability of plants from contrasting habitats to control cation accumulation balance. Dittrichia viscosa plants, from eight siliceous habitats and nine calcareous habitats of southern France and neighbouring Spain and Italy, were cloned and grown together hydroponically with a solution simulating an acid soil with an aluminium constraint. Two independent hydroponic units containing solution supplemented with two levels of Al were used (2 Al levels x17 populations x3 genotypes x3 replicates). The growth and cation content (K, Ca, Mg and Na) of plant shoots and the chemical composition of the soil of each habitat were analysed. At the high Al level (1.1mm), populations differed in K, Ca and Mg plant proportions. Two groups could be distinguished: one containing all but one siliceous populations and the other containing all but one calcareous populations. Plants of the siliceous group accumulated proportionally more K and less Ca, and had better growth, than plants of the calcareous group, in the same way as calcifuge and calcicole species when grown on acid soil. At the lowest Al level (0.37mm), differences between siliceous and calcareous populations were less marked. The results suggest that differences in the ability of plants to control K and Ca balance, which appear to be of adaptive significance, could have arisen through selection, and that Dittrichia viscosa has evolved calcifuge and calcicole nutritional ecotypes in siliceous and calcareous habitats respectively. Various degrees of calcifugy, and to a lesser extent of calcicoly, can thus be suggested to occur among the studied populations, some in relation to the intensity of mineral stress in the natural habitats. So far, only functional traits have provided evidence of ecotypic differentiations within Dittrichia viscosa.deceased     

Genome‐scale data reveal that endemic Poecilia populations from small sulphidic springs display no evidence of inbreeding

Populations with limited ranges can be highly vulnerable to changes in their environment and are, thus, of high conservation concern. Populations that experience human‐induced range reductions are often highly inbred and lack genetic diversity, but it is unknown whether this is also the case for populations with naturally small ranges. The fishes Poecilia sulphuraria (listed as critically endangered) and Poecilia thermalis, which are endemic to small hydrogen sulphide‐rich springs in southern Mexico, are examples of such populations with inherently small habitats. We used geometric morphometrics and population genetics to quantify phenotypic and genetic variation within and among two populations of P. sulphuraria and one population of P. thermalis. Principal component analyses revealed phenotypic and genetic differences among the populations. Evidence for inbreeding was low compared to populations that have undergone habitat reduction. The genetic data were also used to infer the demographic history of these populations to obtain estimates for effective population sizes and migration rates. Effective population sizes were large given the small habitats of these populations. Our results imply that these three endemic extremophile populations should each be considered separately for conservation purposes. Additionally, this study suggests that populations in naturally small habitats may have lower rates of inbreeding and higher genetic diversity than expected, and therefore may be better equipped to handle environmental perturbations than anticipated. We caution, however, that the inferred lack of inbreeding and the large effective population sizes could potentially be a result of colonization by genetically diverse ancestors.     

Genetic structure of lake and stream populations in a Pyrenean amphibian (Calotriton asper) reveals evolutionary significant units associated with paedomorphosis

Differences in environmental conditions such as those between lakes and streams can produce phenotypic variation and ultimately promote evolutionary diversification. Some species of newts and salamanders can occupy these habitats and express alternative phenotypes: metamorphs that lose gills at metamorphosis and paedomorphs that retain them at the adult stage. Whereas this process is facultative in some species, it is obligatory in others, thus suggesting that isolation and environmental pressures may have canalized developmental pathways. In this study, we focused our research on the Pyrenean brook newt, Calotriton asper, which is present in both lakes and streams, but whose fully aquatic paedomorphic individuals are only present in lakes. We aimed to determine the genetic structure and differentiation of two paedomorphic populations, including their surrounding stream and lake metamorphic populations, to test whether populations of paedomorphs can constitute evolutionary significant units. Although gene flow was identified between lakes and nearby stream populations, there was a low percentage of dispersers, and the paedomorphic populations were genetically differentiated from the populations of metamorphs. It is likely that the studied lakes have offered peculiar conditions that have allowed the development of a paedomorphic phenotype. These populations and phenotypes therefore constitute good models to understand local adaptations. As each of these populations of paedomorphs can be considered evolutionary significant units that cannot be replaced by other nearby populations in case of a population crash, conservation actions should be focused directly on them.     

The incertae sedis <Emphasis Type="Italic">Carpathoporella</Emphasis> Dragastan, 1995, from the Lower Cretaceous of Albania: skeletal elements (sclerites,internodes/branches,holdfasts) of colonial octocorals

The incertae sedis Carpathoporella Dragastan, 1995, reported from the Lower Cretaceous of the Western Tethyan domain, is usually interpreted as remains of calcareous algae (Dasycladales or Characeae). New thin-section material from the Aptian of Albania sheds light not only on its biogenic nature but also on the morphological variability of this taxon. In fact, Carpathoporella represents the debris of colonial, bushy, most likely gorgonid octocorals with tuberculated spheroids that may be fused at least near the basal root-like holdfast. Colony branching originates from longitudinally grooved calcareous branches or internodes. Possible relationships to other Upper Cretaceous to Palaeogene genera are discussed and a revised critical inventory of Cretaceous octocorals is presented. Due to the evidenced morphological features, Carpathoporella could either represent an ancestral isidid octocoral of the order Alcyonacea such as Moltkia Steenstrup or, due to the likely primary aragonitic skeletal mineralogy, a representative of Epiphaxum Lonsdale of the order Helioporacea. Due to morphological analogies, the new combination Carpathoporella elliotti (Radoičić) is proposed. In any case, the Lower Cretaceous record from Tethyan peri-reefal shallow-water carbonates is highlighted since numerous skeletal findings of fossil gorgonid Octocorallia were so far only known from Upper Cretaceous and younger strata of outer shelf environments of the boreal realm. The origin of deep-water Upper Cretaceous octocorals from Lower Cretaceous shallow-water taxa such as Carpathoporella is proposed as a possible further example of onshore/offshore evolutionary pattern.     

Natural hybridization in eastern-Mediterranean firs: The case of Abies borisii-regis

The genus Abies is represented in southern Balkans by A. alba, A. cephalonica, and A. borisii-regis. To infer the status of southern-Balkans firs, as well as the extent and patterns of introgression within this taxonomical complex, we analyzed genetic variation patterns of 29 indigenous fir populations in Bulgaria, Macedonia, Greece, and Calabria using a combination of maternally and biparentally inherited markers. Three mitochondrial lineages were observed, one comprising Calabrian populations and two distributed in the Balkans, coinciding with A. alba and A. cephalonica. The boundary between lineages is sharp; only two populations containing a mixture of haplotypes were found. Bayesian analysis of population structure based on seven nuclear microsatellite (nSSR) loci revealed the existence of two clusters whose proportions exhibited a latitudinal cline with a width of 2.3° ( ≈ 255 km). Populations in the center of the latitudinal cline exhibit the most symmetrical, the flattest, and the broadest distributions of cluster proportions within individual tree genomes. A neighbor-net network reflects the cline resulting from the Bayesian analysis. The observed variation patterns are not consistent with the hypothesis of A. borisii-regis as a monophyletic taxon or a stabilized hybridogenous species resulted from ancient hybridization; the taxon rather is a product of recent introgression.     

An introduction to microevolution: rate,pattern, process

This special issue of Genetica brings together a diverse collection of contributions that examine evolution within and among populations (i.e., microevolution), and the role that microevolution plays in the formation of new species and morphological forms (i.e., macroevolution). Many of the papers present evidence of microevolution occuring over contemporary time frames, further validating the near ubiquity of ongoing evolution in the world around us. Several synthetic reviews of empirical work help to define the conditions under which microevolution is or is not likely to occur. Some of the studies speak directly to current controversies in evolutionary biology, such as the relative roles of determinism and contigency, and the nature of the relationship between microevolution and macroevolution. In general, microevolution seems driven largely by deterministic mechanisms, particularly natural selection, but contingency plays a role in (1) determining whether or not suitable conditions are present for evolution to proceed, and (2) guiding the precise manner by which evolution proceeds. Several theoretical treatments and empirical reviews confirm previous research in showing that microevolutionary processes are at least capable of generating macroevolutionary trends. Macroevolution may indeed reflect microevolution writ large but the pattern by which it arises is perhaps best charcaterized as microevolution writ in fits and starts.     

Population Structure and Stock Identification of Eulachon (Thaleichthys pacificus), an Anadromous Smelt, in the Pacific Northwest

The genetic structure of eulachon (Thaleichthys pacificus) populations was examined in an analysis of variation of 14 microsatellite loci representing approximately 1900 fish from 9 sites between the Columbia River and Cook Inlet, Alaska. Significant genetic differentiation occurred among the putative populations. The mean F_ST for all loci was 0.0046, and there was a significant correlation between population genetic differentiation (F_ST) and geographic distance. Simulated mixed-stock samples comprising populations from different regions suggested that variation at microsatellite loci provided reasonably accurate estimates of stock composition for potential fishery samples. Marine sampling indicated that immature eulachons from different rivers, during the 2 to 3 years of prespawning life in offshore marine waters, do not mix thoroughly. For eulachons captured incidentally in offshore trawl fisheries, there was a clear geographic cline in relative abundance of eulachons from different geographic areas. The sample from northern British Columbia was dominated by northern and central coastal populations of British Columbia, the sample from central British Columbia was composed of eulachons from all regions, and the sample from southern British Columbia was dominated by Columbia River and Fraser River populations. These results have implications for the management of trawl fisheries and conservation of spawning populations in some rivers where abundance is at historically low levels.     

Gammaglobulin allotypes of Melanesians from Malaita and Bougainville, Solomon Islands

Data are presented on the distribution of the Gm and Inv allotypes of human IgG in samples from Melanesian populations, three from Malaita and three from Bougainville of the Solomon Islands. The Lau from Malaita are polymorphic for the phenogroup, Gm^{1, 2, 5, 13, 14}. This phenogroup is not known to be polymorphic in any other population of the world. The Inv¹ frequencies of the populations from Malaita are lower than the lowest observed in samples from Bougainville, and this may indicate an extension of the north-south cline for Inv¹ previously reported for Bougainville. Samples from Aita in the north of Bougainville and from the Nagovisi in the south confirm the existence of the north-south cline for Inv¹ in Bougainville and suggest the presence of a Gm cline.     

Local adaptation along an environmental cline in a species with an inversion polymorphism

Polymorphic inversions are ubiquitous across the animal kingdom and are frequently associated with clines in inversion frequencies across environmental gradients. Such clines are thought to result from selection favouring local adaptation; however, empirical tests are scarce. The seaweed fly Coelopa frigida has an α/β inversion polymorphism, and previous work demonstrated that the α inversion frequency declines from the North Sea to the Baltic Sea and is correlated with changes in tidal range, salinity, algal composition and wrackbed stability. Here, we explicitly test the hypothesis that populations of C. frigida along this cline are locally adapted by conducting a reciprocal transplant experiment of four populations along this cline to quantify survival. We found that survival varied significantly across treatments and detected a significant Location x Substrate interaction, indicating local adaptation. Survival models showed that flies from locations at both extremes had highest survival on their native substrates, demonstrating that local adaptation is present at the extremes of the cline. Survival at the two intermediate locations was, however, not elevated at the native substrates, suggesting that gene flow in intermediate habitats may override selection. Together, our results support the notion that population extremes of species with polymorphic inversions are often locally adapted, even when spatially close, consistent with the growing view that inversions can have direct and strong effects on the fitness of species.     

Behavioral Plasticity in Ant Queens: Environmental Manipulation Induces Aggression among Normally Peaceful Queens in the Socially Polymorphic Ant Leptothorax acervorum

The behavioral traits that shape the structure of animal societies vary considerably among species but appear to be less flexible within species or at least within populations. Populations of the ant Leptothorax acervorum differ in how queens interact with other queens. Nestmate queens from extended, homogeneous habitats tolerate each other and contribute quite equally to the offspring of the colony (polygyny: low reproductive skew). In contrast, nestmate queens from patchy habitats establish social hierarchies by biting and antennal boxing, and eventually only the top-ranking queen of the colony lays eggs (functional monogyny: high reproductive skew). Here we investigate whether queen-queen behavior is fixed within populations or whether aggression and high skew can be elicited by manipulation of socio-environmental factors in colonies from low skew populations. An increase of queen/worker ratio and to a lesser extent food limitation elicited queen-queen antagonism in polygynous colonies from Nürnberger Reichswald similar to that underlying social and reproductive hierarchies in high-skew populations from Spain, Japan, and Alaska. In manipulated colonies, queens differed more in ovarian status than in control colonies. This indicates that queens are in principle capable of adapting the magnitude of reproductive skew to environmental changes in behavioral rather than evolutionary time.     

Morphology,ontogeny, and heterochrony in lower and middle Cambrian Gogiids (Eocrinoidea,Echinodermata) from Guizhou Province,China

Gogiid eocrinoids from the Lower Cambrian (Balang Formation — Guizhoueocrinus) and basal Middle Cambrian (Kaili Formation — Sinoeocrinus, Globoeocrinus) in Guizhou Province, China are found in great numbers and are extremely well preserved as high fidelity molds in shale/mud sized siliciclastics. Because of their numbers, complete ontogenetic growth sequences have been observed. Significant differences in growth patterns (heterochrony) are present between Lower and Middle Cambrian genera: thecal plates tend to be paedomorphic in development, while the sutural pores developed between them are peramorphic in their development. Because of the large surface area of theca and brachioles relative to the size of the attachment area, considerable drag in ambient currents would necessitate a strong attachment medium. Anchoring by “biogluing”, possibly by collagen, directly to the substrate or to biodetritus is proposed as the method of attachment.     

Ecological and historical determinants of population genetic structure and diversity in the Mediterranean shrub Rosmarinus officinalis (Lamiaceae)

Population genetic studies of widespread Mediterranean shrubs are scarce compared with those of trees and narrow endemics or studies from phylogeographical perspectives, despite the key role these species may play in Mediterranean ecosystems. Knowledge on the effect of ecological factors in shaping their genetic patterns is also limited. In this study we investigate genetic diversity and population structure across 18 populations of Rosmarinus officinalis, a Mediterranean shrubland plant. Populations were sampled along two elevational gradients, one each on calcareous and siliceous soils in a mountain system in the eastern Iberian Peninsula, to decipher the effect of ecological factors on the genetic diversity and structure based on 11 microsatellite loci. We found overall high levels of genetic diversity and weak population structure. Genetic diversity increased with elevation, whereas population differentiation was stronger among populations growing on siliceous soils. The nested analysis of elevational gradients within soil types revealed that these general patterns were mostly driven by siliceous populations, whereas calcareous populations were more homogeneous along elevational belts. Bayesian analysis of population structure revealed genetic membership of lowland and high‐elevation populations to different genetic clusters and a higher admixture of intermediate‐elevation populations to both clusters. High‐elevation populations were less differentiated from a hypothetical ancestral cluster, suggesting the persistence of their gene pool during the Pleistocene glaciations. In contrast, lowland populations resulted from more recent divergence. We propose that life‐history and reproductive traits mostly contribute to explain the high levels of genetic diversity and weak population structure, whereas ecological and historical factors mostly contribute to the stronger differentiation of siliceous populations and a rapid expansion of R. officinalis on calcareous soils possibly mediated by human landscape transformations, © 2015 The Linnean Society of London, Botanical Journal of the Linnean Society, 2016, 180 , 50–63.