Local adaptation of immunity against a trematode parasite in marine amphipod populations

Resources allocated to defence against parasites are not available for investment in other functions such as growth or reproduction, resulting in trade-offs between different components of an organism’s fitness. In balancing the cost of infection and the cost of immunity, selection should only favour individuals that allocate more energy to resistance and immune responses in populations regularly exposed to debilitating parasites. Here, we compare the ability of amphipods, Paracalliope novizealandiae, to (1) avoid becoming infected and (2) to respond to infection by encapsulating and melanizing parasites, between two natural populations exposed to different risk of parasitism. One population faces high levels of infection by the debilitating trematode parasite Maritrema novaezealandensis, whereas the other population is not parasitised by this trematode nor by any other parasite. Under controlled experimental conditions, with exposure to a standardized dose of parasites, amphipods from the parasite-free population acquired significantly more parasites than those from the population regularly experiencing infection. Furthermore, a lower frequency of amphipods from the parasite-free population succeeded at melanizing (and thus killing) parasites, and they melanized a lower percentage of parasites on average, than amphipods from the parasitised population. These differences persist when individual factors, such as amphipod sex or body length, are taken into account as potential confounding variables. These results support the existence of local adaptation against parasites: an amphipod population that never experiences trematode infections is less capable of resisting infection, both in terms of its first line of defence (avoiding infection) and a later line of defence (fighting parasites following infection), than a population regularly exposed to infection.     

Controls of spatial variation in the prevalence of trematode parasites infecting a marine snail

Geographic variability in abundance can be driven by multiple physical and biological factors operating at multiple scales. To understand the determinants of larval trematode prevalence within populations of the marine snail host Littorina littorea, we quantified many physical and biological variables at 28 New England intertidal sites. A hierarchical, mixed-effects model identified the abundance of gulls (the final hosts and dispersive agents of infective trematode stages) and snail size (a proxy for time of exposure) as the primary factors associated with trematode prevalence. The predominant influence of these variables coupled with routinely low infection rates (21 of the 28 populations exhibited prevalence <12%) suggest broad-scale recruitment limitation of trematodes. Although infection rates were spatially variable, formal analyses detected no regional spatial gradients in either trematode prevalence or independent environmental variables. Trematode prevalence appears to be predominantly determined by local site characteristics favoring high gull abundance.     

Predators selectively graze reproductive structures in a clonal marine organism

Although the fitness consequences of herbivory on terrestrial plants have been extensively studied, considerably less is known about how partial predation impacts the fitness of clonal marine organisms. The trophic role of Caribbean parrotfish on coral reefs is complex: while these fish are important herbivores, as corallivores (consumers of live coral tissue), they selectively graze specific species and colonies of reef-building corals. Though the benefits of parrotfish herbivory for reef resilience and conservation are well documented, the negative consequences of parrotfish grazing for coral reproductive fitness have not been previously determined. We examined recently grazed colonies of Montastraea annularis corals to determine whether grazing was positively associated with coral reproductive effort. We measured gonad number, egg number and size, and proportional reproductive allocation for grazed and intact coral colonies 2–5 days prior to their annual spawning time. We found that parrotfish selectively grazed coral polyps with high total reproductive effort (number of gonads), providing the first evidence that parrotfish selectively target specific tissue areas within a single coral colony. The removal of polyps with high reproductive effort has direct adverse affects on coral fitness, with additional indirect implications for colony growth and survival. We conclude that chronic grazing by parrotfishes has negative fitness consequences for reef-building corals, and by extension, reef ecosystems.     

Seasonal variation in the diversity and abundance of pelagic larvae of Antarctic marine invertebrates

Most marine benthic macroinvertebrate species reproduce via a larval phase but attempts to explain the occurrence of different larval strategies (feeding or non-feeding, pelagic or benthic) in different habitats have been largely inconclusive. There have been very few year-round surveys of meroplankton at any latitude and in consequence fundamental data on the diversity, abundance, and timings of larval life history phases are lacking. There has been considerable debate regarding the viability of pelagic larvae in cold waters with highly seasonal primary production but there has been only one year-round study of meroplankton in the Southern Ocean, and that was outside of the Antarctic Circle. We present data from the first year-round survey of meroplankton assemblages at a location within the Antarctic Circle. We surveyed abundances of meroplanktonic larvae over 1.5 year at Rothera Point, West Antarctic Peninsula (67°34′S, 68°07′W). Larvae were collected in monthly diver-towed net samples close to the seabed at 20 and 6 m total water depths at each of three locations and were identified and counted live immediately after sampling. A total of 99 operationally defined taxonomic types representing 11 phyla were recorded but this is likely to be an underestimate of true diversity because of inherent difficulties of identification. Larvae were present in all months of the year and although planktotrophic larvae were more abundant in summer, both feeding and non-feeding types were present in all months. Comparisons of seasonal larval abundances with data from a settlement study at the same sites and from the literature show that larvae of mobile adults settle in summer regardless of developmental type, whereas sessile taxa settle in all seasons. We suggest that this is a consequence of differences in the food requirements of mobile and sessile fauna and that the availability of food for post-larval juveniles is more critical for survival than factors affecting the larval stage itself.     

Latitudinal variation in local interactions and regional enrichment shape patterns of marine community diversity

While communities are shaped by both local interactions and enrichment from the regional species pool, we propose a hypothesis that the balance of these forces shifts with latitude, with regional enrichment dominating at high latitudes and local interactions dominating at low latitudes. To test this hypothesis, we conducted a latitudinal-scale experiment with marine epifaunal communities. In four regions of the North Atlantic Ocean and Caribbean Sea, we used mimics of ecosystem engineers to manipulate biogenic structural complexity. We iteratively evaluated diversity patterns of experimental communities up to one year after deployment. Additional data were also collected from one of our tropical sites 2.5 years after initial deployment. As hypothesized, we found a reciprocal latitudinal gradient in the effects of the structurally complex mimics and regional enrichment. In the tropics, local diversity was always higher in association with the mimics than in exposed areas that were more open to predation. This effect was consistent across two spatial scales and beyond the one-year timescale of the experiment. In temperate communities, no consistent effects of the mimics on diversity were observed. However, the proportion of species from the regional species pool that were present at the local scale increased from the tropics to the temperate zone, consistent with the hypothesis that higher-latitude communities may experience greater influence from the regional species pool than communities at low latitudes. This study represents the first large-scale experimental demonstration that suggests that the relative impact of local interactions and regional enrichment on community diversity may depend on latitude.     

Spatial autocorrelation analysis of small-scale genetic structure in a clonal soft coral with limited larval dispersal

The philopatric larval dispesal and small effective population sizes characteristic of many clonal species should promote the development of significant small-scale genetic structure within populations as a result of isolation-by-distance. We used spatial autocorrelation statistics to detect genetic structure, arising from both clonal reproduction and philopatric dispersal of sexual propagules, for five allozyme loci within populations of the soft coral Alcyonium sp. In a population on Tatoosh Island, Washington, USA, sampled in 1991/1992, we found significant positive spatial autocorrelation at all loci among individuals separated by <40 cm, reflecting the presence of significant smallscale genetic structure due to associations among clonemates. For 4 of 5 loci, however, we detected no significant spatial autocorrelation among the different clones within this population over distances of 1 to 40 m. Analysis of soft-coral populations from six additional, topographically diverse sites in the north-east Pacific also did not reveal significant spatial autocorrelation among clones at any loci. This general lack of spatial autocorrelation of genotypes among clones suggests that significant small-scale genetic structure has not arisen in populations of Alcyonium sp. as a consequence of isolation-by-distance.     

Genetic diversity increases population productivity in a sessile marine invertebrate

Reductions in genetic diversity can have widespread ecological consequences: populations with higher genetic diversity are more stable, productive and resistant to disturbance or disease than populations with lower genetic diversity. These ecological effects of genetic diversity differ from the more familiar evolutionary consequences of depleting genetic diversity, because ecological effects manifest within a single generation. If common, genetic diversity effects have the potential to change the way we view and manage populations, but our understanding of these effects is far from complete, and the role of genetic diversity in sexually reproducing animals remains unclear. Here, we examined the effects of genetic diversity in a sexually reproducing marine invertebrate in the field. We manipulated the genetic diversity of experimental populations and then measured individual survival, growth, and fecundity, as well as the size of offspring produced by individuals in high and low genetic diversity populations. Overall, we found greater genetic diversity increased performance across all metrics, and that complementarity effects drove the increased productivity of our high-diversity populations. Our results show that differences in genetic diversity among populations can have pervasive effects on population productivity within remarkably short periods of time.     

Mother really knows best: host choice of adult phytophagous insect females reflects a within-host variation in suitability as larval food

Non-random distribution patterns of specialized phytophagous insects on their hosts may depend on intraspecific differences in plant tissue quality, including nutrients and secondary compounds. Secondary compounds are involved in plant resistance, but are also important for the recognition and acceptability of plants as resources by specialized insects. If individuals within a plant species vary in their content of such secondary substances, there may also be qualitative differences between them. In such cases, natural selection will favor insects with the ability to distinguish and prefer the more suitable plants. In Sweden, the leaf beetle Gonioctena linnaeana Schrank (Coleoptera, Chrysomelidae) is highly specialized on one host, the native willow Salix triandra L (Salicaceae). Field observations reveal that some host plants in a population harbor many feeding larvae, causing severe defoliation, whereas neighboring plants may have few or no feeding larvae. Our hypothesis is that the distribution pattern of G. linnaeana larvae in this population results from qualitative differences between individual host plants in combination with the ability of G. linnaeana females to distinguish between plants that are suitable and not suitable for offspring performance. We examine whether larval survival differs depending on diet and whether the content of secondary chemical compounds explains female preference. Based on the higher survival rate of larvae reared on leaves from preferred hosts, we conclude that G. linnaeana females have evolved a behavior that maximizes offspring performance and thus positively affects female fitness. A chemical survey of the plants indicates that luteolin-7-glucoside and an unidentified flavonoid are important for separating the preferred from the non-preferred plants.     

Hidden genetic diversity in a key model species of coral

A total of 43 colonies of the scleractinian coral Pocillopora damicornis from lagoonal and reef slope sites in the western Indian Ocean (WIO) region were genetically characterised at one nuclear and two mitochondrial sequence markers and six microsatellite loci. Both mitochondrial and microsatellite data support the existence of two reciprocally monophyletic clusters (F- and NF-types) and provide evidence of the existence of two cryptic species of P. damicornis on reefs in WIO region and put current morphological delineation and geographical boundaries of P. damicornis and Pocillopora molokensis into question. The results add to ongoing studies on the phylogeny and phylogeography within the genus Pocillopora, which all point towards a range of unresolved morphological and molecular species boundaries. Nuclear phylogenies derived from the present and previously published sequences show evidence for incomplete lineage sorting and/or introgressive hybridisation between Pocillopora morphospecies. However, the two WIO types largely remain in separate clusters, further supporting the theory that these represent two different species.     

Microbial community phylogenetic and trait diversity declines with depth in a marine oxygen minimum zone

Oxygen minimum zones (OMZs) are natural physical features of the world's oceans. They create steep physiochemical gradients in the water column, which most notably include a dramatic draw down in oxygen concentrations over small vertical distances (<100 m). Microbial communities within OMZs play central roles in ocean and global biogeochemical cycles, yet we still lack a fundamental understanding of how microbial biodiversity is distributed across OMZs. Here, we used metagenomic sequencing to investigate microbial diversity across a vertical gradient in the water column during three seasons in the Eastern Tropical South Pacific (ETSP) OMZ. Based on analysis of small subunit ribosomal RNA (SSU rRNA) gene fragments, we found that both taxonomic and phylogenetic diversity declined steeply along the transition from oxygen-rich surface water to the permanent OMZ. We observed similar declines in the diversity of protein-coding gene categories, suggesting a decrease in functional (trait) diversity with depth. Metrics of functional and trait dispersion indicated that microbial communities are phylogenetically and functionally more overdispersed in oxic waters, but clustered within the OMZ. These dispersion patterns suggest that community assembly drivers (e.g., competition, environmental filtering) vary strikingly across the oxygen gradient. To understand the generality of our findings, we compared OMZ results to two marine depth gradients in subtropical oligotrophic sites and found that the oligotrophic sites did not display similar patterns, likely reflecting unique features found in the OMZ. Finally, we discuss how our results may relate to niche theory, diversity-energy relationships and stress gradients.     

Geographic variation in a predator-induced defense and its genetic basis

Predator-induced morphological defenses are a well-known form of phenotypic plasticity, but we continue to have a limited understanding of geographic variation in these responses and its genetic basis. Here we examine genetic variation and geographic differentiation in the inducible defenses of tadpoles (Rana pirica) in response to predatory salamander larvae (Hynobius retardatus). To do so, we crossed male and female frogs from a "mainland" Japanese island having predaceous salamanders and a more isolated island not having predaceous salamanders and raised resulting offspring in the presence and absence of H. retardatus. Mainland tadpoles exhibited a higher capacity to express the inducible morphology (a more bulgy body) than those from the predator-free island, and expression of the bulgy morph in mainland-island hybrids produced phenotypes that were intermediate to those produced by pure crosses. In addition, parental sex had no effect on expression of the bulgy morph. Our results support the hypothesis that geographic variation in inducible defenses is linked to the additive effects of autosomal alleles that are shaped by differences in historical exposure to the inducing predator.     

The consequences of genetic diversity in competitive communities

Several lines of evidence suggest that the species diversity and composition of communities should depend on genetic diversity within component species, but there has been very little effort to directly assess this possibility. Here I use models of competition among genotypes and species to demonstrate a strong positive effect of the number of genotypes per species on species diversity across a range of conditions. Genetic diversity allows species to respond to selection imposed by competition, resulting in both functional convergence and divergence among species depending on their initial niche positions. This ability to respond to selection promotes species coexistence and contributes to a reduction in variation in species composition among communities. These models suggest that whenever individual fitness depends on the degree of functional similarity between a focal individual and its competitors, genetic diversity should promote species coexistence; this prediction is consistent with the few relevant empirical data collected to date. The results point to the importance of considering the genetic origin and diversity of material used in ecological experiments and in restoration efforts, in addition to highlighting potentially important community consequences of the loss of genetic diversity in natural populations.     

Local and regional genetic connectivity in a Caribbean coral reef fish

Coupled bio-physical models of larval dispersal predict that the Costa Rica–Panama (CR–PAN) reefs should constitute a demographically isolated region in the western Caribbean. We tested the hypothesis that CR–PAN coral reef fish populations would be isolated from Mesoamerican Barrier Reef System (MBRS) populations. To test that, we assessed population genetic structure in bicolor damselfish (Stegastes partitus) from both regions. Adult fish were genotyped from five reefs in CR–PAN and from four reefs along the MBRS at 12 microsatellite loci. Between-region F _ST (F _ST = 0.0030, P < 0.005) and exact test (x ² = 74.34, df = 18, P < 0.0001) results indicated that there is weak but significant genetic differentiation between regions, suggesting some restriction in connectivity along the Central American coastline, as predicted by bio-oceanographic models. Additionally, there is among-site genetic structure in the CR–PAN region, relative to the MBRS and between regions, suggesting higher self-recruitment within CR–PAN. This finding may be explained by differences in habitat characteristics.     

Diurnal variation of nitrogen cycling in coastal,marine sediments

A closed chamber technique was developed to determine the emission of microbially produced N₂O from an estuarine sediment. A diurnal variation was observed; maximum emissions of 0.4 to 4.0 mol N₂O–N m^-2 h^-1 were recorded at night whereas the rates were low or even negative, -0.4 to 0.4 mol N₂O–N m^-2 h^-1, during the day. The bacterial denitrification located in the uppermost centimeter was apparently the major source of the emitted N₂O. The diurnal emission pattern was thus inversely related to the O₂ availability at the sediment surface; in the dark, the lack of O₂ production by benthic photosynthesis allowed the denitrification to occur closer to the sediment-water interface and was likely to enhance the release of N₂O to the water. The daily averages for the emission were about 40 mol N₂O–N m^-2 d^-1 for three investigation periods in autumn (November), winter (February) and spring (April), whereas no significant emission was recorded in the NO ₃ ^- -depleted sediment in early summer (June). In this estuary, the N₂O emissions from the sediment were significant contributions to the overall release of N₂O to the atmosphere.     

Local genetic structure and relatedness in a solitary mammal, Neotoma fuscipes

We used DNA microsatellites to investigate temporal and spatial patterns of local genetic differentiation and relatedness in a solitary mammal, the dusky-footed woodrat (Neotoma fuscipes). Patterns of genetic variation were measured relative to spatial clusters, or neighborhoods, of woodrats. We detected significant genetic differentiation among woodrat neighborhoods in two populations spanning multiple habitat types and densities. Estimates of θ _ST among neighborhoods ranged 0.034–0.075 and were comparable to levels reported in social mammals. Genetic differentiation at such a local scale is noteworthy because it occurred in the absence of any physical barriers to gene flow, suggesting that the patterns observed are linked to the nonrandom patterns of mating and dispersal that characterize woodrat social structure. Genetic differentiation and relatedness among neighborhoods were even higher when only resident females were analyzed. These results are consistent with a pattern of female philopatry and male-biased dispersal in woodrats. Geographic distance and relatedness were inversely correlated in adult females at intermediate densities, but not at low densities. Nonetheless, matrilineal genetic structure was apparent even at low woodrat densities based on estimates of θ _ST among neighborhoods of resident females that were significantly greater than zero and consistently greater than estimates including all individuals. In summary, this study demonstrates a matrilineal genetic structure in dusky-footed woodrats. In addition, our results support the idea that intermediate densities may be better at facilitating the formation of spatial kin clusters than either extreme. An erratum to this article can be found at     

Bathymetric patterns of genetic variation in a deep-sea protobranch bivalve, Deminucula atacellana

The origin of the deep-sea benthic fauna is poorly understood and represents an enormous gap in our understanding of basic evolutionary phenomena. One obstacle to studying evolutionary patterns in the deep sea has been the technical difficulty of measuring genetic variation in species that are typically minute, rare, and must be recovered from extreme depths. We used molecular genetic techniques to quantify variation in the 16S rRNA mitochondrial gene within and among populations of the common protobranch bivalve Deminucula atacellana (Schenck, 1939). We analyzed 89 individuals from nine samples collected in the 1960s along a depth gradient from 1100 to 3800 m in the western North Atlantic. Genetic variability within populations is much lower than between populations, and peak haplotype numbers occur near the center of its depth distribution. Continental slope (<2500 m) and rise (>2500 m) populations were genetically distinct despite the lack of any obvious topographic or oceanographic features that would impede gene flow. These findings indicate that the deep-sea macrofauna can have strong population structure over small (134 km) spatial scales, similar to that observed in shallow-water and terrestrial organisms. This surprisingly high biodiversity at the genetic level affords the potential for adaptation and evolutionary diversification, the ultimate historical causes of high species diversity in the deep-sea benthos. Received: 24 July 1997 / Accepted: 26 January 1998     

Loss of genetic diversity in Harpacticoida near offshore platforms

Offshore oil and gas production platforms can be a source of chronic stress that could lead to sublethal impacts on resident benthic organisms. In June 1993 and January 1994, genetic diversity of Harpacticoida (Copepoda) living proximal to operating, offshore platforms in the Gulf of Mexico was estimated to test if platforms are associated with strong selective pressures. Because harpacticoids have short generation times and direct benthic development, they are suitable organisms for examining population responses. Genetic diversity was estimated by comparing restriction fragment length polymorphisms generated from mitochondrial DNA amplified by the polymerase chain reaction on individuals of five species (Cletodes sp., Enhydrosoma pericoense, Normanella sp., Robertsonia sp., and Tachidiella sp.). Populations living at Near regions (stations<50 m from a platform) had significantly less haplotype diversity than populations of the same species living at Far regions (stations>3 km from a platform). The levels of haplotype diversity exhibited by the Far populations were similar at three different platforms located hundreds of kilometers apart. The differences in haplotype diversity between Near and Far regions were the result of a higher proportion of dominant haplotypes, and a loss of less common haplotypes. Haplotypic diversity was inversely correlated with a multivariate measurement of levels of sediment contaminants. The pattern of haplotype diversity on the Gulf of Mexico continental shelf seems to consist of a uniform level of haplotype diversity, punctuated by islands of lower diversity around oil and gas platforms. The selective pressures that lead to a loss of genetic diversity may be the result of contaminants, other differences in the physico-chemical environment, or disturbance in general.     

Regional variation in the hierarchical partitioning of diversity in coral-dwelling fishes

The size of the regional species pool may influence local patterns of diversity. However, it is unclear whether certain spatial scales are less sensitive to regional influences than others. Additive partitioning was used to separate coral-dwelling fish diversity to its alpha and beta components, at multiple scales, in several regions across the Indo-Pacific. We then examined how the relative contribution of these components changes with increased regional diversity. By employing specific random-placement null models, we overcome methodological problems with local-regional regressions. We show that, although alpha and beta diversities within each region are consistently different from random-placement null models, the increase in beta diversities among regions was similar to that predicted once heterogeneity in coral habitat was accounted for. In contrast, alpha diversity within single coral heads was limited and increased less than predicted by the null models. This was correlated with increased intraspecific aggregation in more diverse regions and is consistent with ecological limitations on the number of coexisting species at the local scale. These results suggest that, apart from very small spatial scales, variation in the partitioning of fish diversity along regional species richness gradients is driven overwhelmingly by the corresponding gradients in coral assemblage structure.     

Patterns of genetic subdivision in populations of a clonal cnidarian,Zoanthus coppingeri,from the Great Barrier Reef

Samples of an intertidal zoanthid, Zoanthus coppingeri, Haddon and Shackelton, 1891, were collected from three localities in the Great Barrier Reef region during 1992–1993, and subjected to allozyme electrophoretic analysis at seven polymorphic loci. The reduced ratio of observed to expected genotypic diversity indicated that populations were partly clonal, but they were not dominated by a few clones as occurs in some other cnidarians. Regular disturbance by wave action is postulated to prevent the formation of large stands of particular clones by clearing space and mixing genotypes over small scales. The sexual origin of clonal genotypes was confirmed by conformance to Hardy-Weinberg predictions of genotype frequencies at all but one locus. Values of the standardised genetic variance among populations, F _ST , were highly significant between localities and between replicate sites within localities separated by only 50 m. Strong genetic structure has not previously been described in a Great Barrier Reef invertebrate species, and is considered to be the consequence of stochastic changes in gene frequencies as a result of low levels of gene flow. High clonal longevity and low recruitment rates may maintain genetic differences over long periods. Similar effects may be seen in other Great Barrier Reef invertebrate species with comparable reproductive patterns.