Spatial and temporal patterns of recruitment of the tunicate Ciona intestinalis on a mussel farm in Nova Scotia, Canada

Although possibly indigenous to Nova Scotia, the ascidian Ciona intestinalis has become a problem for local mussel aquaculturists. The local population ecology of the ascidian was studied by recording the depth, distribution, and timing of recruitment of C. intestinalis at four different locations (high wave exposure, low wave exposure, recently fallowed, and historically heavily fouled with tunicates) on a mussel farm at Indian Point, Nova Scotia. Animals were collected weekly from 11 collectors deployed at a depth of 4.5 m over two seasons (June-December 2003 and 2004). Recruitment occurred in two peaks during both seasons: the first peak lasted from late June to late July while the second peak lasted from early September to mid-November. Recruitment was highest in the sheltered and historically fouled location, intermediate in the fallowed site, and lowest in the site with highest wave exposure. Collectors were also deployed at different depths. Recruitment was highest at 4.5 m and to a lesser extent at 8.5 m. Little recruitment was seen at the surface (0.5 m) or in deeper water (12.5 and 16.5 m), except during the second peak in 2004. Overall, these results seem to indicate an abbreviated larval dispersal phase with young recruits settling very close to the pool of adults located on the mussel lines.     

Process of invasiveness among exotic tunicates in Prince Edward Island, Canada

Over the past decade, four exotic tunicates (Styela clava, Ciona intestinalis, Botrylloides violaceus and Botryllus schlosseri) have been reported in the Brudenell estuary in Prince Edward Island (PEI), Canada. Styela clava was the first exotic tunicate to arrive in 1997, rapidly establishing, spreading, invading, and eventually becoming a nuisance in several estuaries of PEI. In the Brudenell estuary, S. clava remained the only exotic nuisance tunicate until 2003. In the fall of 2004, the vase tunicate C. intestinalis, was reported in low abundance, followed by the two colonial species, B. schlosseri and B. violaceus, reported in the spring of 2005. The abundance of C. intestinalis rapidly increased post-introduction, eventually replacing S. clava as the foremost nuisance species on mussel farms in the estuary. To date, C. intestinalis continues to colonize this estuary at epidemic proportions, resulting in the continuing drop of S. clava abundance. The current abundance of C. intestinalis is estimated at 5 cm⁻², which is similar to S. clava abundance at its height in 2003. The 2006 abundance of S. clava is estimated to have fallen to near 0 cm⁻². The dominance of C. intestinalis as a fouling organism on mussel farms is considered a serious threat to this aquaculture industry, mainly due to its unmanageable weight. The process of the detection, establishment, invasiveness, and eventual rise to nuisance level of exotic tunicates in the Brudenell River is presented.     

Variation in the ability of Didemnum sp. to invade established communities

Over the past 30 years southern New England, USA has been invaded by several species of ascidians, including Botrylloides violaceus, Diplosoma listerianum, Styela clava, and Ascidiella aspersa. These species have become dominate in coastal embayments and marinas but are usually absent from more open water coastal areas. A colonial ascidian, Didemnum sp. has invaded southern New England during the past 10 years and we first observed this species in eastern Long Island Sound in 2000. It has become the dominant at several field sites while remaining in low abundance at others. We conducted an experiment at two places, a protected marina and an open coast site, to examine its ability to compete with the established fouling community. Small colonies of Didemnum were transplanted onto panels with communities that varied in age from one to four weeks old and these treatments along with controls with only Didemnum were exposed at both sites. In most treatments Didemnum became a dominant species in the communities at both sites but it reached higher abundances at the open coast site. Potential causes of the observed differences are predation on other species of ascidians at the open coast site reducing recruitment of these species and competition, lower tolerance for elevated temperatures at the marina site, or other environmental parameters that might affect growth rates.     

The alien ascidian Styela clava now invading the Sea of Marmara (Tunicata: Ascidiacea)

During the implementation of a large project aimed to investigate the benthic community structures of the Sea of Marmara, specimens of the invasive ascidian species Styela clava were collected on natural substrata (rocks) at 10 m depth at one locality (Karamürsel) in İzmit Bay. The specimens were mature, containing gametes, indicating that the species had become established in the area. The Sea of Marmara seems to provide suitable conditions for this species to survive and form proliferating populations.     

Biofouling ascidians on aquaculture gear as potential vectors of harmful algal introductions

Biofouling ascidians are ubiquitous in coastal ecosystems and are among the main colonizers of aquaculture gear. Our study tested the hypothesis that the transport, removal, and transfer of fouling ascidian species by aquaculturists provide a mechanism for concentration and distribution of harmful-algal cells to new areas. Wild-caught specimens of common, biofouling ascidian species (Styela clava, Ciona intestinalis, Molgula manhattensis, Botrylloides violaceus, Didemnum vexillum, and Botryllus schlosseri) were exposed individually to cultured strains of co-occurring harmful algae (Prorocentrum minimum, Alexandrium fundyense, Alexandrium monilatum, Karenia brevis, Aureococcus anophagefferens, or Heterosigma akashiwo) at simulated bloom cell densities of each HAB species. After feeding, ascidians were transferred to ultrafiltered seawater. Immediately after exposure, and after 24 and 48 h in ultrafiltered seawater, biodeposits were collected and observed microscopically for the presence of intact, potentially viable cells. Subsamples of biodeposits were transferred into culture tubes with ultrafiltered seawater and monitored for algal growth during 8 weeks. Cells of all HAB species were found to pass intact through the ascidian digestive system, remained viable, and in many cases were capable of re-establishing populations at least 48 h post-ingestion. The results of our study will inform industry and managers of the potential threat and ecological impact of spreading biofouling ascidians, and practices to mitigate adverse impacts. Additionally, these management practices have been formally incorporated into a new cost-share program developed to help shellfish producers prevent the further spread of ascidians and associated HAB species.     

Distribution and long-term temporal patterns of four invasive colonial ascidians in the Gulf of Maine

Invasive ascidians are a growing concern for ecologists and natural resource managers, yet few studies have documented their short- and long-term temporal patterns of abundance. This study focuses on the invasion of the Gulf of Maine by the colonial ascidians Botryllus schlosseri, Botrylloides violaceus, Diplosoma listerianum and Didemnum sp. A. We examined the time of arrival and potential vectors for these four invasive ascidians using survey data (collected from 1969 onwards) and literature documentation. We also compared the dominance and seasonal patterns of abundance of these species using data from two identical panel studies; one conducted from 1979 to 1980, the other from 2003 to 2005. Didemnum and Botrylloides were most likely first introduced into the Damariscotta River, Maine in the early 1970's through oyster aquaculture while Botryllus and Diplosoma were probably transported by commercial and recreational vessels. The overall abundance of colonial ascidians has increased since 1979 and 1980. Botryllus was the only invasive colonial ascidian present during the 1979 to 1980 study and accounted for an average of 6.16% cover on panels. From 2003 to 2005, the more recently arrived colonial ascidians Botrylloides and Didemnum accounted for 7.38% and 2.08% cover respectively, while Botryllus covered only 1.16%. Our results reveal a shift in seasonal abundance between 1979 to 1980 and 2003 to 2004. In 1979 and 1980, colonial ascidians had the highest percent cover in fall and winter while in 2003 and 2005 they had highest percent cover in summer and fall. Seasonal patterns of space occupation by colonial ascidians were correlated with seasonal changes in seawater temperature.     

The influence of pre‐settlement and early post‐settlement processes on the adult distribution and relative dominance of two invasive mussel species

相似文献   

Isolation and characterization of twelve polymorphic microsatellite markers for the invasive ascidian Styela clava (Tunicata)

The invasive solitary ascidian Styela clava has spread extensively along European coasts since its first occurrence in the early 1950s. In order to characterize spatial and temporal patterns of genetic change during its establishment and subsequent spread, we developed 12 species‐specific loci from an enriched microsatellite library. Polymorphism was explored in one native and two introduced populations (N = 31 ± 1). Number of alleles per locus varied from two to 13 (mean = 7). The average expected heterozygosity within populations ranged from 0.539 to 0.580.     

Development and application of tools for incursion response: Lessons learned from the management of the fouling pest Didemnum vexillum

In October 2001, an unidentified didemnid ascidian, was recorded for the first time in New Zealand, smothering wharf piles and moorings in a northern harbour. A heavily-fouled barge then translocated the ascidian to an international shipping port some 500 km south, near the heart of the New Zealand mussel industry. The species was subsequently identified as Didemnum vexillum, but its status as indigenous or non-indigenous was disputed. Nonetheless, its presence was regarded as a significant threat to the mussel industry because of its demonstrated invasiveness on artificial structures, and its ability to over-grow and smother mussels.From the barge's mooring area, D. vexillum subsequently spread to the seabed beneath, and to nearby vessels and artificial structures (i.e., barges, recreational vessels, moorings, salmon cages and wharf piles). Given the likelihood that infected vectors would spread the ascidian to mussel farms in the region, and in consideration of a benefit-cost analysis, an eradication program for D. vexillum was instigated. This paper provides a chronology of events surrounding the initial detection and spread of the ascidian, and describes the development of incursion response tools for the different substrata that were infected. The treatments included smothering soft-sediment habitats with uncontaminated dredge spoil, wrapping wharf piles with plastic, smothering rip-rap habitats using a geotextile fabric, and various other approaches based on water blasting, air drying or chlorine dosing. While many of the response methods were completely effective at eliminating D. vexillum from different substrata, the program overall failed to eradicate the organism from the region. The reasons for this failure are documented, and the important lessons learned are highlighted, as a contribution to the successful management of invasive species in the future.     

Effects of native and invasive macroalgal canopies on composition and abundance of mobile benthic macrofauna and turf-forming algae

We examined the effects of native kelps, Laminaria longicruris de la Pylaie and L. digitata (Hudson) Lamouroux, and of the invasive alga, Codium fragile ssp. tomentosoides (Van Goor) Silva, on the composition and abundance of mobile benthic macrofauna and of turf algae by measuring the response of these assemblages to experimental removal of the respective macroalgal canopy. From June 2003 to November 2004, we censused macrofauna and measured canopy cover within 4 × 10 m strips of alternating Canopy Intact (control) and Canopy Removed treatments in both a Codium- and a Laminaria-dominated habitat in the rocky subtidal zone of a semi-protected embayment on the Atlantic coast of Nova Scotia, Canada. Macroalgal canopy cover fluctuated seasonally, peaking in September/October (69% cover in 2003; 55% cover in 2004) for Codium and in May 2004 (70% cover) for kelps, and with both canopy types reaching a winter minimum in January 2004 (22 and 28% cover, respectively). In both Codium and Laminaria habitats, significant effects of canopy removal on the overall macrofaunal assemblage were evident only during periods in which canopy cover in the Canopy Intact treatment was ≥ 50%. In the Codium habitat, 4 out of 11 characteristic taxa were more abundant in the Canopy Intact treatment, where taxonomic diversity also was higher. In contrast, 4 out of 11 characteristic taxa in the Laminaria habitat were more abundant in the Canopy Removed treatment and diversity was similar between treatments. Turf algae were sampled in November 2004 and, despite between-treatment differences in the light regime, there was no significant effect of canopy removal. Our results indicate that selection of algal habitats by mobile macrofauna is likely determined by the different shelter and foraging opportunities offered by these morphologically dissimilar ecosystem engineers.     

Identification and population genetic comparison of three ascidian species based on mtDNA sequences

Ascidians are sessile marine chordate invertebrates found along seashores worldwide and are typically regarded as invasive organisms. Knowledge concerning their global genetic structure and subsequent invasive potential is limited. Here, we identified three ascidians—Ciona robusta, Ciona savignyi, and Styela clava from the northeast region of China using morphological characteristics and mitochondrial cytochrome c oxidase subunit I (cox1) as genetic marker. We additionally used phylogenetics to aid in the identification of these three species. The results of a population genetic analysis showed that among the three species, the level of haplotype diversity was particularly high within C. savignyi, and nucleotide diversity varied moderately. We divided the three species separately into native and invasive populations using 170 cox1 sequences from global resources to explore population genetic structure and invasive potential. Although in the network analysis Ciona spp. formed haplogroups of native and invasive populations, some haplotypes were still shared. We found that the haplotypes did not cluster within the network of S. clava. Our AMOVA results also showed that Ciona spp. had a weak genetic structure, and less genetic differentiation was present in S. clava. These data suggest that there are extensive incursions of these three ascidians into different geographical regions. Global comparisons of ascidian populations will help in the understanding of their population genetic structure and invasive potential, hence providing important insights regarding conservation as well as management.     

Chaetognaths of the United States South Atlantic Bight: Distribution, abundance and potential interactions with newly spawned larval fish

Chaetognaths are one of the most numerous organisms in the zooplankton community off the coast of North Carolina. During two years of offshore sampling in the late winter to early spring, sixteen chaetognath species were identified, four of which had not previously been reported in the waters of the United States South Atlantic Bight. Offshore samples were dominated by Sagitta enflata Grassi, 1881, one of the larger species, which contributed > 61% of total chaetognath abundance while dominant coastal species were S. tenuis Conant, 1896 and S. hispida Conant, 1895. Abundances, body sizes and spatial distributions were determined for the most abundant chaetognath species along with the overall abundance of three common co-occurring larval fish species (spot, Atlantic croaker and Atlantic menhaden). In addition, laboratory feeding experiments were conducted using S. tenuis and S. hispida to estimate the potential impact of chaetognath predation on representative North Carolina larval fish which spawn offshore and subsequently migrate into local estuaries. Feeding rates (no. prey items day^− 1) varied with prey type and the condition (starved/fed) of the chaetognath. Weight specific daily rations (SDR) were found to vary inversely with chaetognath size, decreasing exponentially with an increase in chaetognath length. The observed abundance and distribution data indicate that wintertime chaetognath populations in offshore waters of Onslow Bay, North Carolina have the potential to interact with recently spawned larval fish and may simultaneously act as competitors, predators, or prey. Furthermore, calculations using published values of chaetognath abundance, zooplankton standing crops, and our SDR estimates indicate that chaetognaths in a representative North Carolina estuary would require a minimum of 5.96 cal m^− 3 day^− 1 to sustain their biomass. Allowing for the reported spatial and temporal variability in zooplankton abundance in these systems, chaetognaths should consume 4.4%-20.9% of the estimated total zooplankton production day^− 1. This further emphasizes the role of chaetognaths not only as predators, but also as competitors with larval fish for zooplankton food stocks in southeastern United Sates estuaries.     

Life cycle of the lilac pyralid Palpita nigropunctalis (Bremer) (Lepidoptera: Crambidae) on five oleaceous tree species

The seasonal abundance of Palpita nigropunctalis larvae was studied on five oleaceous tree species in Ibaraki, central Japan, for two years. The larval population peaked on some tree species in both spring and autumn while it peaked on other tree species only in autumn. In bimodal populations, the spring peaks consisted of larvae infesting leaves, while the autumn peaks consisted of larvae infesting various tree parts (flowers, fruits and leaves). Larval development was longer and pupal weights were lower on Ligustrum lucidum fruits than on Ligustrum japonicum fruits. Thus, L. japonicum fruits were suitable for larval development in autumn. First-instar larvae appeared to occur three times per year (in late April to early May, mid to late September and early to mid October). Adults were observed from late March to early April, mid May to early June, and early September to mid November at the census sites, showing that P. nigropunctalis had three generations a year. The percentage of females having premature ovaries were 64.3 and 12.5% at 15 °C under 14-h and 15-h photophases, respectively, and 28.6 and 25% at 20 °C under the respective photophases. These temperatures and photoperiods are typical of those in May and June in the census sites. The ovaries of females collected in the field between late May and mid July were premature, in agreement with the laboratory experiments. The pupal duration was prolonged under the short photoperiod, especially at reduced temperatures. We discuss a possible life cycle of P. nigropunctalis in Ibaraki.     

Fine-scale spatial and temporal variation in settlement of the intertidal mussel Perna perna indicates differential hydrodynamic delivery of larvae to the shore

Scales of spatial and temporal variation in the settlement of early and late plantigrades of the mussel Perna perna to artificial substrata were measured on the south coast of South Africa. One study examined daily settlement over one month at three sites of similar tidal height and aspect. The sites were separated by 100′s of m with 5 replicated artificial settlement substrata approximately 1 m apart within each site. Early (< 0.5 mm) and large late plantigrades (1 - 3.5 mm) dominated samples with few individuals of 0.5-1 mm. Settler abundance showed clear peaks, synchronised among sites and apparently following spring tides, though the relationship between tide and settlement was not significant. Two-way ANOVA indicated significant spatial variation, which varied with time, at the 100 m-scale among sites. A second study investigated the local effects of substratum aspect. Seaward facing surfaces had consistently lower settlement (but higher adult cover) than horizontal and landward facing surfaces, this effect was significant for large late plantigrades. This suggests that either juveniles migrate after settlement, or that post-settlement mortality differs markedly among aspects. The high degree of temporal variation at the scale of days and weeks was expected, but the results also suggest a degree of pulsing in settlement, with maximum intensities around spring tide. A previous study of nearshore waters indicated a common pool of larvae at scales of 100 s m. Thus, consistent ranking of sites for all size classes of settlers indicates predictably differential hydrodynamic delivery of larvae to sites separated by 100 s m. The effect of aspect on settlement highlights the importance of smaller scale hydrodynamics, suggesting that the stronger turbulence experienced by seaward facing surfaces may affect larval settlement negatively.     

Recent population expansions of non-native ascidians in The Netherlands

Over the last 50 yrs seven non-native ascidians have settled in The Netherlands, concentrated in the two periods 1974-1977 and 1991-2004 (i.e., Styela clava, Aplidium glabrum, Diplosoma listerianum, Didemnum sp., Botrylloides violaceus, Molgula complanata and Perophora japonica). The year of the introduction of B. violaceus remains a matter of dispute because many of the Botrylloides specimens that are recorded in western Europe, have been identified as the closely resembling species B. leachi. Only Didemnum sp. has become a true invasive species and has become a threat to native ecosystems, especially in the province of Zeeland, by its ability to overgrow virtually all hard substrata present. This includes rocks, stones, sand, algae and almost all sessile marine animals. The sudden population expansion of the didemnid from 1996 onward, coincided with the cold winter of 1995-1996, which caused decreased population sizes of many marine animals. The resulting increase in the availability of suitable substrates for settlement and the strong decrease of grazing sea urchins, may have triggered the population expansion. Studying its population dynamics, the optimal growing temperature for Didemnum sp. appears to be 14-18 °C. Virtually all colonies die when the water temperature gets colder than 5 °C. Colonies growing on live marine animals seem to be more resistant to the cold, than those growing on rocks, stones and plants. Two potential predators of the didemnid have also been recorded in Dutch waters: the gastropods Trivia arctica and Lamellaria sp.     

Patterns of adult abundance in Chthamalus stellatus (Poli) and C. montagui Southward (Crustacea: Cirripedia) emerge during late recruitment

The aim of this study was to investigate when adult distribution patterns are established in the barnacles Chthamalus stellatus and C. montagui. Adult ‘zones’ were identified by analysing field counts of both species at mid and upper shore heights. Monthly collections of cyprids, < 1 month old metamorphs and recruits (all metamorphosed individuals older than approximately 1 month) were made for C. stellatus and C. montagui in natural barnacle beds at six shores in SW Ireland. This was carried out over one year in 1996/1997, using a hierarchical sampling design. Abundance of total recruits (0-3 months old) was compared between adult zones after the main settlement season had ended. In addition, scales of variability in 0-3 month recruitment into adult zones were compared between the species at two scales: shores (1000s of metres) and sites within shores (10s of metres). Older recruits of each species, up to 11 months of age, were also compared between adult zones.The majority of settlement (measured as attached cyprids) occurred between August and October 1996. In October, there was no effect of adult zone on the abundance of total (0-3 month) recruitment up to that point in either species. Despite this homogeneity in recruitment between adult zones, significant spatial variation was found in 0-3 month recruits of both species at both of the scales examined. In C. stellatus the amount of variation associated with the larger scale (shore) was more than twice that of sites or of the residual variation (replicates within sites). 0-3 month recruitment in C. montagui was also most variable at the scale of shores but the residual variability (between replicates within site) was of similar magnitude to that of shores. Variability in 0-3 month C. montagui recruitment was relatively low at the scale of sites.There was a small but consistent input of recruits to adult zones over 9 months of the year, complicating the assessment of when adult patterns were set-up in these species. By June 1997, characteristic patterns of adult dominance had been established at all shores. Settlement had completely ceased by this time and individual barnacles were potentially 11 months old. Neither settlement nor early recruitment are significant in determining adult zonation patterns in these species. Instead, differential mortality patterns in individuals up to the age of 11 months are implicated in determining patterns of distribution of both species.     

Benthic ctenophore (Order Platyctenida) reproduction,recruitment, and seasonality in south Florida

Reproductive structures, modes, and seasonal patterns of size–class abundances are examined in two benthic platyctene (Family Coeloplanidae) ctenophore species present in dissimilar shallow marine environments in subtropical southeast Florida. Coeloplana waltoni, a minute (1–3 mm body length) epizoic associate of octocorals, occurs in exposed environments often under turbulent conditions, and Vallicula multiformis (2–10 mm) commonly occurs epiphytically on macroalgae in protected, calm‐water environments. Reproductive activity in C. waltoni is most active during the warm‐water summer season (June–October); gonadal development in V. multiformis occurs year‐round, and is most pronounced during sea‐warming periods in late spring (May) and late summer to early autumn (August–October), with release of cydippid larvae. Both species are hermaphroditic brooders, exhibiting paedogenesis (early gonadal development) at body lengths approximately one‐third (Coeloplana) to one‐sixth (Vallicula) of maximum adult size. Juvenile individuals (<0.6 mm) increased in abundance in C. waltoni during the summer reproductive period, and large (≥1 mm) pink‐colored individuals comprised 50% or more of samples from July through September. Seasonal abundance of gravid individuals and the timing of cydippid larval release in V. multiformis did not correspond closely with juvenile or adult population densities. Asexual fragmentation occurred in both ctenophore species, but was observed more frequently in individuals of V. multiformis. This asexual mode of reproduction probably accounted in part for the discordance between ctenophore abundances and larval recruitment events by sexual means. Morphological structures and behaviors associated with reproduction are described in this study. Uncommon images of reproductive products (gametes, embryos, larvae), spawning events, brooding, and asexual fragmentation are included, some for the first time in the published literature.     

Environmental factors associated with invasion: modelling occurrence data from a coordinated sampling programme for Pacific oysters

Documenting establishment and spread of invasive species requires extensive co-ordinated sampling programmes. Identifying the factors promoting or inhibiting local establishment of an invasive species can improve capacity to predict further spread and underpin strategies to limit spread. Here, a structured sampling programme was used to assess the current distribution of feral populations of Pacific oysters, Crassostrea gigas, in Ireland. Sixty-nine sites were sampled using a standardised protocol combining semi-quantitative and quantitative approaches. Sites were chosen to represent variation in proximity to aquaculture and a range of environmental variables. Oyster populations were found at 18 locations, with densities ranging from single individuals to nine individuals per m². The broad size range of oysters found is indicative of more than one recruitment event. Logistic regression indicated that feral oysters were positively associated with the presence of hard substrata or biogenic reef, long residence times of embayments and large intertidal areas. There was also a tendency for oysters to occur disproportionately in bays with aquaculture, but >500 m from it. Small-scale analysis within sites showed that oysters were almost exclusively attached to hard substrata and mussel shell. The approach taken here provides a rigorous repeatable methodology for future monitoring and a detailed basis for the prediction of further spread.     

The presence of GABA in gastropod mucus and its role in inducing larval settlement

Chemical substances that induce larval settlement have been the focus of many gastropod studies due to the importance of wild stock recruitment and production within aquaculture facilities. Gamma-aminobutyric acid (GABA), GABA analogs, and GABA-mimetics associated with certain crustose coralline algae (CCA), are known to induce larval settlement in commercial abalone (Haliotis) species, and other gastropods. Furthermore, mucus secreted from these gastropods has been shown to induce larval settlement, but the stimulatory components of mucus have not been thoroughly investigated. We now present data confirming that GABA is the settlement-inducing effector molecule contained within abalone mucus. To do this, we initially generated anti-GABA for use in immunoenzyme and immunofluorescent microscopy. Using these techniques GABA was identified in the nerves and epithelial cells of the foot, including mucus. Dried mucus samples subject to HPLC analysis revealed a mean concentration of 0.68 mM GABA after sample rehydration. The presence of GABA in these samples was confirmed by time-of-flight mass spectroscopy (TOF-MS). In addition, GABA was detected in the mucus of several abalone species and other gastropods by immunocytochemistry. Subsequent bioassays using both dry and fresh mucus strongly promoted induction of larval settlement.