Temporal stability in patterns of genetic diversity and structure of a marine foundation species (Zostera marina).

Genetic diversity and population structure reflect complex interactions among a diverse set of processes that may vary temporally, limiting their potential to predict ecological and evolutionary outcomes. Yet, the stability of these patterns is rarely tested. We resampled eelgrass (Zostera marina) meadows from published studies to determine variability in genetic diversity and structure within and between meadows over 5-12 years. The meadows sampled (San Francisco, Tomales and Bodega Bays in California and the Virginia coastal bays) represent a range of life histories (annual vs perennial), age (well-established vs restored) and environments (rural vs urbanized). In all of these systems, neither diversity nor differentiation (FST) changed over time. Differences among tidal heights within Bodega Bay were also remarkably consistent, with the high intertidal being more diverse than the subtidal, and tidal height differentiation being modest but significant at both time points. Historical studies used only a few microsatellite loci; therefore, our temporal comparisons were based on 4-5 loci. However, analysis of the current data using a set of 12 loci show that 4-5 loci are sufficient to describe diversity and differentiation patterns in this system. This temporal consistency was not because of the resampling of large clones, underscoring the feasibility and relevance of understanding drivers of the differences. Because seagrasses are declining at rapid rates, restoration and conservation are increasingly a coastal management priority. Our results argue that surveys of eelgrass genetic structure and diversity at decadal scales can provide accurate depictions of populations, increasing the utility of published genetic data for restoration and designing networks of reserves.

Color polymorphism and genetic structure in the sea star Pisaster ochraceus.

The sea star Pisaster ochraceus is one of the more striking species on the rocky shores of the Northeast Pacific, in part due to the dramatic color polymorphism of the adults. Along the open Pacific coast, Pisaster populations are 6%-28% orange, with a small percentage of brilliant purple stars and a large percentage of reddish-brown to dull purple ones. However, populations in the San Juan Island Archipelago (Washington, USA) and the southern Strait of Georgia (British Columbia, Canada) are almost entirely brilliant purple. The factors that maintain the color polymorphism, and those that contribute to among-site variation in color frequencies, remain unknown. We examined the relationships between color frequencies and several ecological and morphological variables, and conducted a large-scale phylogeographic survey of Pisaster populations. We found very low population genetic structure, suggesting that gene flow is high and geographic variation in color frequencies is not a vestige of Pleistocene glacial refugia. Color frequencies are also unrelated to adult size and to the frequency of injury within a population. However, there are suggestive relationships between color frequency and diet, and with areas of potentially low salinity. We propose that, although the color polymorphism may have an underlying genetic component, the regional-scale variation in color frequency is ecologically controlled.

Mate selection and the evolution of highly polymorphic self/nonself recognition genes.

Multicellular organisms use the products of highly polymorphic genes to distinguish self from conspecific nonself cells or tissues. These allorecognition polymorphisms may regulate somatic interactions between hosts and pathogens or between competitors (to avoid various forms of parasitism), as well as reproductive interactions between mates or between gametes (to avoid inbreeding). In both cases, rare alleles may be advantageous, but it remains unclear which mechanism maintains the genetic polymorphism for specificity in self/nonself recognition. Contrary to earlier reports, we show that mate selection cannot be a strong force maintaining allorecognition polymorphism in two colonial marine invertebrates. Instead, the regulation of intraspecific competitive interactions appears to promote the evolution of polymorphisms in these species.

Intrapopulation sex ratio variation in the salt grass Distichlis spicata.

ABSTRACT In many dioecious plant populations, males and females appear to be spatially segregated, a pattern that is difficult to explain given its potentially high costs. However, in asexually propagating species, spatial segregation of the sexes may be indistinguishable from superficially similar patterns generated by random establishment of a few genets followed by extensive clonal spread and by gender-specific differences in rates of clonal spread. In populations where a significant fraction of individuals are not flowering and gender cannot be assigned to this fraction, apparent spatial segregation of the sexes may be due to differential flowering between the sexes. We confirm reports that flowering ramets of the clonal, perennial grass Distichlis spicata are spatially segregated by sex. We extend these studies in two fundamental ways and demonstrate that this species exhibits true spatial segregation of the sexes. First, using RAPD markers, we estimated that at least 50% of ramets in patches with biased sex ratios represent distinct genotypes. Second, we identified a RAPD marker linked to female phenotype (eliminating the possibility that gender is environmentally determined) and used it to show that the majority of patches exhibit significantly biased sex ratios for both ramets and genets, regardless of flowering status.

One cell, two cell, red cell, blue cell: The persistence of a unicellular stage in multicellular life histories.

As developmental biologists come closer to understanding at the molecular and genetic levels how a zygote becomes an adult, it is easy to forget that the very phenomenon that gives them an occupation remains a vexing problem to evolutionary biologists: why do unicellular stages persist in life histories of multicellular organisms? There are two explanatory hypotheses. One is that a unicellular stage purges multicellular organisms of deleterious mutations by exposing offspring that are each uniformly of one genotype to selection. Another is that a one-cell stage reduces conflicts of interest among genetically different replicators within an organism.

The analysis of paternity and maternity in the marine hydrozoan Hydractinia symbiolongicarpus using randomly amplified polymorphic DNA (RAPD) markers.

For organisms in which direct observation of mating and subsequent dispersal of offspring and relatives is impossible, patterns of reproductive success and genealogical relationship can only be established using genetic markers. The ideal genetic assay would (1) employ highly polymorphic genetic markers for distinguishing among individuals; (2) use little tissue for analysing early life-history stages; and (3) require minimal investment in time and money for population level studies. From this perspective, DNA polymorphisms revealed by PCR amplification using random ten-base primers [Randomly Amplified Polymorphic DNA (PCR-RAPD) or Arbitrarily Primed DNA (AP-PCR)] have great potential. However, the evidence that RAPD/AP markers are both heritable and can be repeatably amplified remains controversial. This study characterizes patterns of inheritance and polymorphism of RAPD markers in the free-spawning, colonial marine hydrozoan Hydractinia symbiolongicarpus. In all cases, the amplification products were identical among extractions from the same clone. Of 56 primers screened, 13 had sufficient polymorphism and scoreability for an analysis of parentage and higher-order genetic relationships in three matings. These primers generated 156 unique amplification products (putative loci), of which 133 were polymorphic. All but four of these loci were inherited as dominant mendelian markers. Our study suggests that the presence of a marker represents a single allele at a locus; however, what appear to be single null alleles may actually comprise several segregating alleles. When the identity of neither parent was known a priori, inclusion (unique markers present in offspring and only one of the potential parents) proved to be more efficient than exclusion for assigning offspring to parents. The most powerful approach, however, was cluster analysis of all presence/absence information for the marker bands. Clustering avoided the pitfalls caused by the appearance of occasional nonparental bands, and constructed a hierarchical framework that correctly reflected all genealogical relationships.

For adults only? Supply-side ecology and the history of larval biology.

When ecologists study organisms with multiphasic life cycles, they must often confront the problem of which phase to scrutinize. In principle, the dynamics and interactions of all stages could play a major role in the regulation of adult populations and species assemblages. In practice, however, the roles of larger and more sedentary phases - being easier to count and manipulate than motile propagules - have been emphasized. Nonetheless, several recent studies on the small, dispersing larval phase of marine invertebrate life cycles reach the conclusion that the spatial distribution and supply of propagules can control the distribution and abundance of populations of benthic adults. To some, the present emphasis on planktonic propagules amounts to a resurrection of ideas developed during a long and rich history of larval biology. To others, studies of demographic and ecological connections between larval and adult populations represent a substantial revision of ecological paradigms.