Year-round reproduction in a seasonal sea: biological cycle of the introduced ascidian Styela plicata in the Western Mediterranean

The widely introduced ascidian Styela plicata is very common in the Western Mediterranean, an area that can act as a source for secondary introductions due to its high shipping activity. In order to understand the potential of this species to colonize new habitats, its reproductive features were assessed in the Western Mediterranean by means of monthly monitoring of two populations (Vilanova i la Geltrú 41°12′53″N, 1°44′11″E; Blanes 41°40′29″N, 2°47′56″E) from January 2009 to December 2010. The reproductive activity of this species was assessed through gonad histology and a gonad index. Population size-structure was measured monthly in order to study recruitment dynamics. No clear seasonal pattern was observed, and mature gametes and recruits were present all year long. Spawning was potentially continuous, although it seemed punctuated with pulses of gamete release, particularly in spring. A prolonged reproductive period is likely to confer a competitive advantage on S. plicata in temperate seas, where most species reproduce seasonally, and may promote recurrent introductions as larvae are available for settlement on transport vectors over much of the year.     

Metabolic maintenance costs of the suspension feeder Styela plicata

The bioenergetic basis of the biannual reproductive cycle of the solitary tunicate Styela plicata was investigated in order to evaluate hypotheses concerning the lack of larval settlement in summer. The rate of ingestion and absorption efficiency were measured in order to provide an estimate of the rate at which material was made available for maintenance, growth, and reproduction. At a given temperature the rate of ingestion was proportional to the 0.7 power of wet mass. the ingestion rate increased rapidly with increasing temperature between 12° and 18°C (Q₁₀3), but was independent of temperature between 18° and 28°C. Absorption efficiency was independent of temperature and body size and averaged approximately one-third for both carbon and nitrogen. Metabolic maintenance costs were estimated from measurements of oxygen consumption and excretion of ammonia and urea reported for s. plicata. These require only 18±11% of the carbon and 37±22% of the nitrogen absorbed from the gut of S. plicata over the temperature range 12° to 28°C. Metabolic maintenance makes no excessive demands on the material absorbed in the gut at a particular time of year, and a surplus of carbon and nitrogen substrate is available throughout the year for growth and reproduction. Predation on larvae and young adults may be responsible for the low rate of settlement observed in summer months.     

Mixed but not admixed: a spatial analysis of genetic variation of an invasive ascidian on natural and artificial substrates

Following the introduction to a new area (pre-border dispersal), post-border processes determine the success in the establishment of non-indigenous species (NIS). However, little is known on how these post-border processes shape the genetic composition of NIS at regional scales. Here, we analyse genetic variation in introduced populations along impacted coastlines to infer demographic and kinship dynamics at the post-border stage. We used as a model system the ascidian species Microcosmus squamiger that has been introduced worldwide. This species can colonize and grow fast on man-made artificial structures, impacting activities such as mariculture. However, it can also establish itself on natural substrates, thus altering natural communities and becoming an ecological problem. We genotyped 302 individuals from eight populations established on natural and artificial substrates in the north-western Mediterranean Sea, using six microsatellite loci. We then compared the resulting genotypes with those found within the native range of the species. We found high levels of genetic diversity and allelic richness in all populations, with an overall deficit of heterozygotes. Autocorrelation analyses showed that there was no within-population genetic structure (at a scale of tens of metres); likewise, no significant differentiation in pairwise comparisons between populations (tens of kilometres apart) and no isolation-by-distance pattern was found. The results suggest that M. squamiger has a natural capacity for high dispersal from one patch of hard substrate to another and no differences whatsoever could be substantiated between natural and artificial substrates. Interestingly, two groups of genetically differentiated individuals were detected that were associated with the two ancestral source areas of the worldwide expansion of the species. Individual assignment tests showed the coexistence of individuals of these two clusters in all populations but with little interbreeding among them as the frequency of admixed individuals was only 15 %. The mechanism responsible for maintaining these genetic pools unmixed is unknown, but it does not appear to compromise post-border colonization of introduced populations.     

The reproductive ecology of the invasive ascidian, <Emphasis Type="Italic">Styela clava,</Emphasis> in Auckland Harbour,New Zealand

The ascidian Styela clava, native to the north-west Pacific, is an invasive species affecting New Zealand’s marine ecosystems, biodiversity and aquaculture operations. To provide detailed information on the reproductive biology of S. clava in New Zealand for post-border biosecurity management, long-term seasonal patterns of gametogenesis were determined from May 2006 to May 2008 in Auckland’s Waitemata Harbour (36°49′20″S, 174°45′85″E). Of particular interest was whether the critical 15°C threshold spawning temperature for reproduction observed in the Northern Hemisphere applied here to the first Southern Hemisphere study. S. clava gametogenesis followed a regular seasonal cycle with ripe gametes appearing as early as September and persisting to June; this time frame corresponds to the period when sea surface temperatures in the region first reach 15°C and with spawning occurring mainly during late summer to early autumn. From photoperiod manipulation, it was determined that spawning occurred at approximately 18:20. The extended reproductive period and a short generation time in the Waitemata Harbour provides a lengthy opportunity for S. clava to spread. Findings are discussed in relation to S. clava’s post-border management.     

Growth and reproductive cycles of the marine fouling ascidians Ciona intestinalis,Styela plicata,Botrylloides violaceus,and Leptoclinum mitsukurii at Aburatsubo-Moroiso Inlet (central Japan)

Experimental cultivations of post-metamorphosis juveniles were repeated in different seasons for Ciona intestinalis (L.) and Styela plicata (Lesueur) (simple ascidians) as well as for Leptolinum mitsukurii (Oka) and Botrylloides violaceus Oka (compound ascidians) at Aburatsubo-Moroiso Inlet near Misaki Biological Station (Japan). Growth in body length of simple ascidians was exponential during juvenile development up to near sexual maturity. C. intestinalis grew to sexual maturity in 1 month in summer and in 2 months in winter; S. plicata, in 2 months in summer and in nearly 5 months in winter. The doubling time decreased approximately twofold in the two simple ascidians with a 10C° increase in environmental temperature. Growth of colonies in compound ascidians was exponential after maturation of the first functional zooids. Sexual reproduction of L. mitsukurii and B. violaceus was continuous throughout the year, forming many generations in a year. Doubling time of colony growth in the two compound ascidians decreased approximately threefold with a 10C° increase in temperature.     

Oceanic variability and coastal topography shape genetic structure in a long-dispersing sea urchin

Understanding the scale of marine population connectivity is critical for the conservation and sustainable management of marine resources. For many marine species adults are benthic and relatively immobile, so patterns of larval dispersal and recruitment provide the key to understanding marine population connectivity. Contrary to previous expectations, recent studies have often detected unexpectedly low dispersal and fine-scale population structure in the sea, leading to a paradigm shift in how marine systems are viewed. Nonetheless, the link between fine-scale marine population structure and the underlying physical and biological processes has not been made. Here we show that patterns of genetic structure and population connectivity in the broadcast-spawning and long-distance dispersing sea urchin Centrostephanus rodgersii are influenced by physical oceanographic and geographic variables. Despite weak genetic differentiation and no isolation-by-distance over thousands of kilometers among samples from eastern Australia and northern New Zealand, fine-scale genetic structure was associated with sea surface temperature (SST) variability and geography along the southeastern Australian coast. The zone of high SST variability is characterized by periodic shedding of eddies from the East Australian Current, and we suggest that ocean current circulation may, through its influence on larval transport and recruitment, interact with the genetic consequences of large variance in individual reproductive success to generate patterns of fine-scale patchy genetic structure. If proven consistent across species, our findings suggest that the optimal scale for fisheries management and reserve design should vary among localities in relation to regional oceanographic variability and coastal geography.     

Microgeographic genetic differentiation in a colonial ascidian (Botryllus schlosseri) population

Limited gene flow via the restricted dispersal of larvae and gametes is expected to result in the genetic differentiation of populations of clonal invertebrates on small spatial scales. However, occasional dispersal events over greater distances may generate sufficient gene flow to maintain genetic homogeneity. We applied a spatial autocorrelation approach that does not require a priori definitions of subdivision boundaries to examine genetic differentiation within a continuous population of the colonial ascidian Botryllus schlosseri (Pallas) at two allozyme and five polychromatism loci. Colonies were sampled in July 1992, on a 12 by 18 m grid superimposed on a shallow subtidal (1 to 3 m) population in the Damariscotta River estuary in Maine, USA. Low but significant levels of positive autocorrelation were detected over very small spatial scales (<5 m), with negative autocorrelation occurring on larger scales (>8 m). This pattern indicates significant genetic differentiation over distances of 8 to 21 m, and is consistent with genetic drift and inbreeding creating small scale genetic structure. Received: 18 October 1999 / Accepted: 11 July 2000     

Low genetic differentiation between isolated populations of the colonial ascidian Symplegma rubra Monniot, C. 1972

Pelagic larvae are highly important for maintaining the gene flow among populations of sessile marine invertebrates. Colonial ascidians consist, exclusively, of brooding species, with lecithotrophic larvae that have a limited dispersal. As a result, there is a marked differentiation among populations. In this work, we used allozyme electrophoresis to access the genetic variation in four populations of Symplegma rubra, a colonial ascidian frequently found in the intertidal zone of Southeastern Brazilian coast. High variability was found at three of the four sites sampled, the exception being Praia Grande in the State of Rio de Janeiro. At this site, there was a great preponderance of clones, which possibly reflected the enclosed nature of the location and its low water circulation that reduce the dispersal capabilities of these animals. S. rubra did not conform to expectations for random mating (Hardy–Weinberg equilibrium): there was a deficit of heterozygotes that was more related to the small population size than to inbreeding processes, since F _is analysis per locus revealed a deficiency of heterozygotes at only one locus—MDH*. The greatest variation in allele frequency was found for GPI-2*. Analyses of genetic variability revealed moderate differentiation among the populations (F _ST=0.051), which was unexpected for a species with a low dispersal capability. Rafting, a frequently underestimated means of dispersal, may be the main mode of gene flow in this species over large areas, since colonies of S. rubra are frequently seen growing on drift material and there is no evidence that the larva survive for a long time in the plankton.     

土壤空间变异性研究评述

在研究方法上，土壤空间变异性研究经历了由定性描述到传统统计学、再到地统计学的不断改进过程，而研究内容也由对单一特性的研究逐渐转向多特性的研究。土壤空间变异性研究虽取得了明显的进展，但也存在地统计学方法在实际应用中主观性太强，土壤综合特性的空间变异性研究、时空结合的变异性研究以及与数学模型有机结合研究不够深入等问题。文章在分析国内外研究现状的基础上，提出了亟待深入研究的若干问题，对这些问题的研究有助于“数字土壤”、“精准农业”和生态环境建设的实施。     

Grazing regulates the spatial variability of periphyton biomass

The presence of consumers not only alters the mean biomass of the prey assemblage, but also affects the spatial heterogeneity of biomass distribution. Whereas the mean prey biomass is generally reduced by consumer presence, the effect on spatial heterogeneity is less clear-cut. A meta-analysis of almost 600 field experiments manipulating the presence of benthic invertebrate or vertebrate grazers was conducted to analyze the effect of grazers on both the absolute spatial variability of periphyton biomass and the relative variability, which was standardized to the mean. Effects on absolute variability were measured as the log response ratio of the standard deviation of biomass (LR-SD), whereas effects on relative variability were measured as the log response ratio of the coefficient of variation of biomass (LR-CV). The overall magnitude and range of LR-SD and LR-CV indicated that grazers not only reduced periphyton biomass, but also substantially altered their spatial distribution. However, grazer effects differed strongly for absolute and relative variability. On average, grazers reduced the absolute spatial variability in prey biomass by 50% (average LR-SD = -0.68) but increased the relative variability by 24% (average LR-CV = 0.22). The magnitude of LR-SD strongly depended on the efficiency of grazing, with strong biomass removal leading to strong homogenization. Moreover, LR-CV and LR-SD were significantly affected by habitat type (freshwater vs. coastal) and substrata. Given the importance of spatial heterogeneity for resource uptake, competition and the maintenance of diversity, grazer presence has potentially strong indirect effects on the interactions within prey assemblages.     

Multispecies genetic objectives in spatial conservation planning

Growing threats to biodiversity and global alteration of habitats and species distributions make it increasingly necessary to consider evolutionary patterns in conservation decision making. Yet, there is no clear‐cut guidance on how genetic features can be incorporated into conservation‐planning processes, despite multiple molecular markers and several genetic metrics for each marker type to choose from. Genetic patterns differ between species, but the potential tradeoffs among genetic objectives for multiple species in conservation planning are currently understudied. We compared spatial conservation prioritizations derived from 2 metrics of genetic diversity (nucleotide and haplotype diversity) and 2 metrics of genetic isolation (private haplotypes and local genetic differentiation) in mitochondrial DNA of 5 marine species. We compared outcomes of conservation plans based only on habitat representation with plans based on genetic data and habitat representation. Fewer priority areas were selected for conservation plans based solely on habitat representation than on plans that included habitat and genetic data. All 4 genetic metrics selected approximately similar conservation‐priority areas, which is likely a result of prioritizing genetic patterns across a genetically diverse array of species. Largely, our results suggest that multispecies genetic conservation objectives are vital to creating protected‐area networks that appropriately preserve community‐level evolutionary patterns.     

Long-term variability in the structure of subtidal benthic communities in Puget Sound,Washington, USA

Data on benthic infauna from 4 permanent stations in Puget Sound off Seattle, USA, collected during 1963–1964, 1967, and 1969, revealed considerable stability in numbers of species and specimens and in diversity within stations among sampling dates. The species composition of the faunal assemblages also remained rather constant during the period of investigation, but the relative dominance among the numerically important species varied somewhat. Biomass data did not differ significantly in 1964 and 1969, but the 1967 data were considerably lower at all stations.     

Fine-scale genetic structure in the surface brooding Caribbean octocoral,Antillogorgia elisabethae

Larval dispersal is a critical component of marine species’ life histories because it controls their population dynamics. Dispersal distance can be inferred by the presence and scale of spatial genetic structure (SGS). The larvae of Antillogorgia elisabethae, a surface brooding Caribbean octocoral, have been observed to settle within meters of maternal colonies, although many disperse over greater distances. Using a spatially hierarchical sampling design, A. elisabethae recruits were collected from a site off the coast of Great Abaco, The Bahamas, in June of 2009–2011. Seven microsatellite loci were used to determine whether genetic structure was present among recruits on scales of <1 m to 10s of meters. Multilocus genotypes were autocorrelated within spatial distance classes using Moran’s I coefficient as an estimate of genetic similarity, and Rousset’s genetic distance (â). Genetic structure was present on a scale of <10 m in 2010, but not in the other 2 years, suggesting that larval dispersal was more localized in 2010. Moran’s I coefficients were positive and significant for 0–1, 1–2 and 2–5 m classes in 2010, and the average genetic distance (â) among recruits in the 0–1 m class was significantly lower than the other classes. Autocorrelograms suggested that genetic patch sizes were 7.5 m in 2010, and 20 m in 2009 and 2011. Differences in the SGS of recruits between years might reflect temporal variation in temperature, wind and/or current speeds affecting larval dispersal, or variation in reproduction at the site.     

Mitochondrial control region variability and global population structure in the swordfish,Xiphias gladius

Little is known about what limits genetic exchange in highly vagile, open ocean vertebrate species, such as the swordfish Xiphias gladius L. Reduced abundance of swordfish in some regions, and increased fishing pressure in others, has raised concerns and fueled interest in a more complete evaluation of the resource. In this study, global population structure in swordfish was assessed by sequencing a 300 base pair segment of the 5 end of mitochondrial DNA control region from 159 swordfish collected in three ocean basins: the Mediterranean, Atlantic and Pacific, over the years 1988 to 1994. Among the 159 individuals, 95 polymorphic sites delineated 121 unique haplotypes, indicating a high level of polymorphism on a global scale. A phylogenetic analysis of the unique DNA haplotypes revealed two divergent clades with differing geographic distributions. Phylogeographic concordance of this pattern with that of two other pelagic fish species suggests a biogeographic explanation for this structure. An analysis of molecular variance (AMOVA) revealed significant geographic partitioning of molecular variation among the three ocean basins, indicating that swordfish populations are structured on a global scale. Estimates of genetic exchange among populations within an ocean basin were high, indicating panmixia within ocean basins. Since the haplotypic diversity exhibited by the swordfish control regions is extremely high, much larger sample sizes may be necessary to detect subdivision within ocean basins.     

Structural, morphological and genetic variability in Halophila stipulacea (Hydrocharitaceae) populations in the western Mediterranean

Halophila stipulacea (Forssk.) Ascher. is a dioecious seagrass that colonized the Mediterranean basin probably following the opening of the Suez Canal (1869). Natural meadows have been reported since the end of the last century on the eastern side of the basin and only recently along the northern coast of Sicily. In the present study we examined the morphological and genetic variability of two natural meadows located along the Sicilian coast (Vulcano Island and Oliveri-Tindari coastal lakes). In order to determine morphological and genetic polymorphism, samples were collected at different depths (5, 15 and 25 m depth at the Vulcano site) and positions (edge vs mid-bed) within the meadows. Statistically significant differences in phenotypic features were found between the factors “depth” and “position” within the same meadow and between the two localities. Genetic diversity was assessed using randomly amplified polymorphic DNA and found to be high. Deep and shallow stands of the Vulcano Island meadow clustered in different positions in the UPGMA tree. The shallow Vulcano stand was found to be closer to the shallow Oliveri-Tindari meadow than to the deeper stand from Vulcano. Mantel's test did not allow rejection of the null hypothesis of independence of morphological and molecular distance matrices. We conclude that (i) H. stipulacea shows high morphological and genetic polymorphism, (ii) environmental and/or ecological barriers exist between different depths, and (iii) trends of morphological and genetic variability may be influenced by different environmental and/or ecological factors. Received: 15 November 1998 / Accepted: 21 May 1999