The control of the development of a marine benthic community by predation on recruits

Recruitment is an important process in regulating many marine benthic communities and many studies have examined factors controlling the dispersal and distribution of larval immigrants. However, benthic species also have early post-settlement life-stages that are dramatically different from adult and larval stages. Predation on these stages potentially impacts measured recruitment and the benthic populations and communities that ultimately develop.We examined the consequences of post-settlement predation on 1-day-old to 1-month-old recruits of sessile invertebrates at two field sites in southern New England. One site (Breakwater) was in a protected area with few predators and the other (Pine Island) was <1 km away in an open coast area with three different predator guilds: small and large invertebrates and fish. The Breakwater site had been dominated for >10 years by colonial and solitary ascidians. These species were absent from the Pine Island site which was dominated by bryozoans. Our goal was to examine whether post-settlement predation influenced the development and subsequent structure of the epifaunal community.Here we examine long-term changes in community development resulting from post-settlement predation, and contrast these results to those of earlier experiments examining the reductions in observed recruitment by post-settlement predation. Our first long-term experiment examined natural community development at the two sites and whether transplanted communities changed when exposed to the different levels of predation at these sites. The communities that developed at both sites were consistently different from each other and similar to resident communities at their respective sites. On panels transplanted from the Breakwater to Pine Island, solitary ascidians and the colonial ascidian, Botryllus schlosseri, suffered high mortalities on both caged and uncaged treatments, indicative of predation by small predators that could enter cages. Some solitary ascidians did survive inside cages and the colonial ascidian, Botrylloides violaceus, became dominant on all transplanted treatments. On panels transplanted from Pine Island to the Breakwater, ascidians invaded and dominated all treatments except those that were originally caged at Pine Island.In the second long-term experiment, natural communities were allowed to develop on panels exposed at the Breakwater for 1, 2, 3, and 4 weeks. Each set was transplanted to three treatments at Pine Island: open uncaged pilings, caged pilings to exclude fish and large invertebrates, and racks suspended above the bottom to exclude all predators. When 1-week-old communities were transplanted, after 2-3 weeks only bryozoans were found on the open and caged pilings, while colonial ascidians dominated the suspended rack treatment. When older 2-week-old communities were transplanted, colonial ascidians also became dominant in the caged piling treatment and when 3- and 4-week-old communities were transplanted colonial ascidians dominated all three treatments. Solitary ascidians were never abundant on open pilings exposed to fish and large benthic invertebrate predators.Post-settlement predator-prey interactions involved newly settled and juvenile life-stages of a variety of prey species and many invertebrate and vertebrate predator species. The effects of these interactions on recruitment did result in differences in the development and eventual species composition of the communities, even though predators had little if any effect on the adults of the prey species.     

Persistence and space preemption explain species‐specific founder effects on the organization of marine sessile communities

Community assembly may not follow predictable successional stages, with a large fraction of the species pool constituted by potential pioneering species and successful founders defined through lottery. In such systems, priority effects may be relevant in the determination of trajectories of developing communities and hence diversity and assemblage structure at later advanced states. In order to assess how different founder species may trigger variable community trajectories and structures, we conducted an experimental study using subtidal sessile assemblages as model. We manipulated the identity of functionally different founders and initial colony size (a proxy of the time lag before the arrival of later species), and followed trajectories. We did not observe any effects of colony size on response variables, suggesting that priority effects take place even when the time lag between the establishment of pioneering species and late colonizers is very short. Late community structure at experimental panels that started either with the colonial ascidian Botrylloides nigrum, or the arborescent bryozoan Bugula neritina, was similar to control panels allowed natural assembling. In spite of high potential for fast space domination, and hence negative priority effects, B. nigrum suffered high mortality and did not persist throughout succession. Bugula neritina provided complex physical microhabitats through conspecific clustering that have enhanced larval settlement of late species arrivals, but no apparent facilitation was observed. Differently, panels founded by the encrusting bryozoan Schizoporella errata led to different and less diverse communities compared to naturally assembled panels, evidencing strong negative priority effects through higher persistence and space preemption. Schizoporella errata founder colonies inhibited further conspecific settlement, which may greatly relax intraspecific competition, allowing resource allocation to colony growth and space domination, thus reducing the chances for the establishment of other species.     

The influence of resident adults on larval settlement: experiments with four species of ascidians

Residents within any community can affect the larval settlement of both their own and other species. In marine sessile communities resident adults can affect larval settlement by preying on settling larvae, removing or adding space for the larvae to colonize, or stimulating or prohibiting larval settlement on available substratum nearby. To examine those processes by which residents affect settlement, we exposed experimental substrata with three densities of adults of a single species at a site in eastern Long Island Sound, USA for a 24-h period. Four species of common ascidians, Botryllus schlosseri (Pallas), Botrylloides diegensis Ritter and Forsyth, Diplosoma macdonaldi Herdman, and Molgula manhattensis (De Kay), were used in 11 separate experiments. Few individuals of any species settling attached to the surfaces of these species and this resulted in the main effect of these residents being the usurpation of space and the restricting of settlement to unoccupied areas. A model is also presented to explain the apparent aggregated settlement of several species in open areas adjacent to the resident ascidians. From this model we suggest that the aggregated settlement can result from limited larval mobility such that some larvae that contact and reject the resident species as settlement sites may subsequently contact open surfaces of the same substratum and increase settlement densities there over those observed on control substrata. Finally, settlement data for several species indicate that Molgula may influence settlement by preying on larvae.     

Temperature,space availability,and species assemblages impact competition in global fouling communities

Competition drives community composition in many ecosystems and can influence the spread of invasive species. Marine fouling communities are excellent study systems for competition because of space limitation and the abundance of invasive species. While many studies have examined individual or site-specific responses to changes in temperature or presence of invasive species, it is difficult to predict ecological impacts without assessing interspecific interactions over a wide geographic range. This study compared interactions between several globally distributed invasive fouling species over a broad geographic range. Weekly examination of photographs of settlement panels in marinas at 18 sites around the world allowed for the quantification of competitive outcomes. In the north Atlantic, experimental panels became covered with fouling organisms exponentially faster at warmer temperatures, while northeast and south Pacific sites did not. An invasive ascidian (Diplosoma listerianum) and bryozoan (Bugula neritina) were strong competitors, but most species displayed a negative response in high competition settings where there was little available space. Two species (Botryllus schlosseri and Botrylloides violaceus) had better competitive outcomes at cooler temperatures, possibly due to fewer strong competitors at these sites. Thus, warmer sites with little open space and multiple strong competitors are likely most resistant to future invasions, while colder sites with more open space and weaker competitors would be more susceptible to invasive species. These results suggest that the establishment and spread of invasive fouling species is likely to be influence by seawater temperature, available space, and the competitive abilities of community members.     

Effects of small-scale disturbance on invasion success in marine communities

Introductions of non-indigenous species have resulted in many ecological problems including the reduction of biodiversity, decline of commercially important species and alteration of ecosystems. The link between disturbance and invasion potential has rarely been studied in the marine environment where dominance hierarchies, dynamics of larval supply, and resource acquisition may differ greatly from terrestrial systems. In this study, hard substrate marine communities in Long Island Sound, USA were used to assess the effect of disturbance on resident species and recent invaders, ascidian growth form (i.e. colonial and solitary growth form), and the dominant species-specific responses within the community. Community age was an additional factor considered through manipulation of 5-wk old assemblages and 1-yr old assemblages. Disturbance treatments, exposing primary substrate, were characterized by frequency (single, biweekly, monthly) and magnitude (20%, 48%, 80%) of disturbance. In communities of different ages, disturbance frequency had a significant positive effect on space occupation of recent invaders and a significant negative effect on resident species. In the 5-wk community, magnitude of disturbance also had a significant effect. Disturbance also had a significant effect on ascidian growth form; colonial species occupied more primary space than controls in response to increased disturbance frequency and magnitude. In contrast, solitary species occupied significantly less space than controls. Species-specific responses were similar regardless of community age. The non-native colonial ascidian Diplosoma listerianum responded positively to increased disturbance frequency and magnitude, and occupied more primary space in treatments than in controls. The resident solitary ascidian Molgula manhattensis responded negatively to increased disturbance frequency and magnitude, and occupied less primary space in treatments than in controls. Small-scale biological disturbances, by creating space, may facilitate the success of invasive species and colonial organisms in the development of subtidal hard substrate communities.     

Seasonal non-indigenous species succession in a marine macrofouling invertebrate community

We report results from a 2-year study on the succession pattern of macrofouling assemblages in the Taranto Sea, an important alien species hotspot in the Mediterranean Sea. Four sets of PVC panels were used as macrofouling collectors; each unit was installed at a different time (April 2013, July 2013, October 2013 and January 2014) and then surveyed quarterly for 1 year. The macrofouling community consisted of 93 sessile invertebrate species, of which 16 were NIS and five were cryptogenic. In both years non-indigenous species (NIS) recruitment occurred mainly in the quarter July/October in concert with the settlement of pioneer autochthonous species. This recruitment is independent of immersion time, occurring on both bare substrates and on previously colonized panels. This increase in NIS coverage is influenced by the development stage of the community, suggesting that NIS grow better without potential competitors. Two sets of NIS were distinguished. The first included abundant ascidians, serpulids, and bryozoans that are structuring components of early communities when favorable conditions exist (i.e. a lack of competitive autochthonous species). After settlement, these species are unable to develop in later-stage communities. The second set of NIS was composed of sabellid worms that settle in early and late communities but, unlike the other NIS, are able to persist and become dominant in late macrofouling communities independent of seasonal changes.     

Reduction of pollution impacts through the control of toxicant release rate must be site- and season-specific

The marine environment is commonly used for the deliberate disposal of industrial, mining and metabolic wastes. Managers will benefit from experimental work that identifies ways of reducing environmental impacts by varying the frequency and intensity of toxicant release. Using a field dosing technique we investigated the effects of three frequencies of pulse copper pollution event, and two intensities of copper dose, on developing assemblages of sessile marine invertebrates. The resulting impacts could then be compared in assemblages exposed to the same amount of toxicant via different disturbance regimes. The experiment was replicated simultaneously at three sites within Port Philip Bay, Victoria, Australia. Pulse pollution events altered assemblage composition through a direct negative effect on densities of large solitary ascidians. In response to the removal of the spatially dominant solitary ascidians, there were increases in recruitment of many different phyla, and in the abundance of older individuals of some serpulid and bryozoan taxa. Biodiversity, as measured by total species number and Shannon's H′, did not reflect the dramatic structural changes apparent within assemblages. If pulse pollution events had either a negative or positive effect on a species' density, then that effect was accentuated by increasing the intensity (strength) or the frequency of the pollution disturbance. Within populations, however, adult mortality might benefit new recruits of the same species through the freeing up of settlement space. In this case, the effects of copper pulses were evident as changes in the population size structure rather than the overall density of an organism. There were variable responses to manipulating the output rate of the toxicant between sites that appear to be driven by the rate of recovery of the dominant space occupiers. At one site there was a negligible effect of disturbance intensity and impacts could be minimized through reducing only the frequency of toxicant release. At two other sites both the intensity and frequency of disturbance determined the pollution effect and minimizing impacts was feasible only through an overall reduction in the amount of toxicant released. The management and reduction of pollution impacts through the control of toxicant release will require site- and season-specific modifications.     

Recruitment in invertebrates with short-lived larvae: the case of the bryozoan <Emphasis Type="Italic">Disporella hispida</Emphasis> (Fleming)

Temporal and spatial recruitment patterns of the cyclostome bryozoan Disporella hispida were monitored using settlement plates arranged along three benthic communities of an artificial reef at Blanes (NE Spain, NW Mediterranean). At the study site, the species mainly inhabits semi-obscure caves. By studying recruitment over one year I first inferred the larval release period for D. hispida and described its temporal occurrence in the communities and stations studied. Secondly, I attempted to determine whether the predicted restricted dispersal may account for the species' distribution at the study site. To this purpose, I compared the distribution of early recruits (15 days old) with that of adults. I also investigated environmental factors which may affect the extent of larval dispersal, and described the effects of post-recruitment processes occurring over a 4-month period. The brooding period, inferred from the study of early recruitment, was linked to spring-increments and autumn-decrements of water temperatures. Early recruits were distributed non-randomly in the communities and stations studied, being most abundant in the habitats where adults live. Strong hydrodynamic events seemed to modify this pattern, allowing recruitment out of the parental communities, and may hinder settlement. Post-recruitment mortality events were likely to prevent colonisation of habitats where the species do not live. Overall, philopatry and low post-recruitment mortality in the parental communities appeared to be the main mechanisms determining the distribution of D. hispida at the study site. Electronic Publication     

Experimental separation of effects of consumers on sessile prey in the low zone of a rocky shore in the Bay of Panama: direct and indirect consequences of food web complexity

The effects of predation by a diverse assemblage of consumers on community structure of sessile prey was evaluated in the low rocky intertidal zone at Taboguilla Island in the Bay of Panama. Four functional groups of consumers were defined: (1) large fishes, (2) small fishes and crabs, (3) herbivorous molluscs, and (4) predaceous gastropods, (l) and (2) included fast-moving consumers and (3) and (4) included slow-moving consumers. Experimental treatments were: no consumers deleted (all groups present), most combinations of deletions of single groups (i.e., one group absent, three present), pairs of groups deleted (two absent, two present), trios of groups deleted (three absent, one present), and the entire consumer assemblage deleted (all groups absent). Changes in abundance (percent cover) of crustose algae, solitary sessile invertebrates, foliose algae, and colonial sessile invertebrates were quantified periodically in 2–4 plots of each treatment from February 1977 to January 1980 after the initiation of the experiment in January 1977.

Space on this shore is normally dominated by crustose algae; foliose algae, solitary sessile invertebrates, and colonial sessile invertebrates are all rare. After deletion of all consumers, ephemeral green algae increased from 0 to nearly 70% cover. Thereafter, a succession of spatial dominants occurred, with peak abundances as follows: the foliose coralline alga Jania spp. by July 1977, the barnacle Balanus inexpectatus by April 1978, and the rock oyster Chama echinata by January 1980. Although no longer occupying primary rock space, Jania persisted as a dominant or co-dominant turf species (with the brown alga Giffordia mitchelliae and/or the hydrozoan Abietinaria sp.) by colonizing shells of sessile animals as they became abundant instead of the rock surface.

Multivariate analysis variance (MANOVA) indicated that the effect of each group was as follows. Molluscan herbivores grazed foliose algae down to the grazer-resistant, but competitively inferior algal crusts, altered the relative abundances of the crusts, and inhibited recruitment of sessile invertebrates. Predaceous gastropods reduced the abundance of solitary sessile animals. Small fishes and crabs, and large fishes reduced the cover of solitary and colonial sessile animals and foliose algae, although they were incapable of grazing the foliose algae down to the rock surface. Many of the effects of each consumer group on prey groups or species were indirect; some effects were positive and some were negative. The variety of these indirect effects was due to both consumer-prey interactions among the consumers, and competitive or commensalistic interactions among the sessile prey. Comparison of the sum of the effects of each of the single consumer groups (i.e., the sum of the effect observed in treatments with one group absent, three present) with the total effects of all consumers (i.e., the effect observed in the treatment with all groups absent) indicates that a “keystone” consumer was not present in this community. Rather, the impacts of the consumer groups were similar but, due to dietary overlap and compensatory changes among the consumers, not readily detected in deletions of single consumer groups. The normally observed dominance of space by crustose algae is thus maintained by persistent, intense predation by a diverse assemblage of consumers on potentially dominant sessile animals and foliose algae. The large difference in structure between this and temperate intertidal communities seems due to differences in degree, not kind of ecological processes which produce the structure.     

The non-native solitary ascidian Ciona intestinalis (L.) depresses species richness

Non-native ascidians are a dominant feature of many sessile marine communities throughout the world and may have negative effects on species diversity. We tested effects of the non-native Ciona intestinalis on the sessile invertebrate community in San Francisco Bay, where it occurs in dense aggregations. In particular, we compared species richness between PVC panels from which C. intestinalis were experimentally removed to panels with naturally dense C. intestinalis growth, using fouling panels of four sizes (between 49 cm² and 1177 cm²) to measure the effect of C. intestinalis recruitment on species-area relationships. We initially deployed 120 fouling panels (30 of each size) at a site known to have dense populations of C. intestinalis, assigning these to three different treatments: (1) Experimental removal, whereby new recruits of C. intestinalis were removed on a weekly basis, pulling panels out of the water for a short time period to do so; (2) Manipulated control, whereby panels were removed from the water each week (as in the experimental removal) but without C. intestinalis removal; and (3) Unmanipulated control, which remained in the water throughout the experiment. After 4 months, all of the panels were collected and analyzed to estimate species richness and relative abundance (percent cover) of sessile invertebrates and of C. intestinalis. Across all panels, species richness was negatively correlated with C. intestinalis abundance. The removal of C. intestinalis produced communities with significantly higher species richness than the controls. The overall species composition of treated and control panels was also distinctly different, with many species occurring more often in the absence of C. intestinalis, while others occurred more often on C. intestinalis-dominated panels. These data suggest that C. intestinalis both depress local species diversity and alter community assembly processes to fundamentally change sessile community composition.     

The Role of Habitat Complexity in Community Development Is Mediated by Resource Availability

Habitat complexity strongly affects the structure and dynamics of ecological communities, with increased complexity often leading to greater species diversity and abundance. However, habitat complexity changes as communities develop, and some species alter their environment to themselves provide habitat for other species. Most experimental studies manipulate basal substrate complexity, and while the importance of complexity likely changes during community development, few studies have examined the temporal dynamics of this variable. We used two experiments to quantify the importance of basal substrate complexity to sessile marine invertebrate community development through space and time. First, we compared effects of substrate complexity at 70 sites across ten estuaries. Sites differed in recruitment and community development rates, and after three months provided spatial variation in community development stage. Second, we tested for effects of substrate complexity at multiple times at a single site. In both experiments, complexity affected marine sessile invertebrate community composition in the early stages of community development when resource availability was high. Effects of complexity diminished through time as the amount of available space (the primary limiting resource) declined. Our work suggests the presence of a bare-space threshold, at which structural complexity of the basal substrate is overwhelmed by secondary biotic complexity. This threshold will be met at different times depending on local recruitment and growth rates and is likely to vary with productivity gradients.     

Effects of pulse versus steady recruitment on sessile marine communities

Variation in patterns of propagule establishment (recruitment) has important effects on population dynamics and the structure of some communities. Most experimental studies have varied recruitment by changing the nature of a single event early in community development, but recruitment can also vary from steady rates of arrival to highly episodic 'pulse' events, causing differences in the temporal spacing of individuals recruiting into patches. We examined whether two different temporal patterns of recruitment of sessile invertebrates affected temperate marine communities in southeastern Australia in two experiments that were run at different times at the same site and that manipulated several different species. Target species entered communities as either a single pulse of recruits within a 2-week period or steady input of the same total number of recruits over a longer time period (5-6 weeks). The pattern of recruitment had variable effects on community structure. The colonial ascidian Botryllus schlosseri did not have a strong influence on community structure whether it recruited in a single pulse or steadily. The cover of B. schlosseri was higher when recruitment occurred as a single pulse. In a second experiment, botryllid ascidians caused changes in the composition of communities when they recruited steadily compared to when they did not recruit or didemnids recruited, but caused no differences in communities when they recruited in a shorter pulse. In contrast, recruitment frequency of didemnid ascidians had little effect, though their presence/absence caused community differences. Though we found that different temporal recruitment patterns can alter community composition, the life history and ecology of particular taxa as well as differences in environmental background processes are likely to influence the strength of these effects.     

Benthic colonization and succession on temperate sublittoral rocky cliffs

Patterns of benthic colonization and succession were investigated on a temperate rocky cliff (Aegean Sea, Eastern Mediterranean). Cement and ceramic panels deployed on the rocky substratum at 25-30 m depth were sampled every 3 months over a 2-year period yielding 28 floral and 156 animal species. Diversity, cover and abundance had low values at short immersion periods and increased at the long ones. The vagile fauna responded to the increasing habitat complexity offered by the development of algal turfs and sessile species, and the community structure remained highly dynamic. Unlike the duration of immersion, the type of substratum didn't affect species colonization except for decapods that showed a preference for cement panels. Three to four stages of succession were recorded over immersion periods, according to vagile and sessile biota, respectively, during which species richness and abundance increased. The structure of the developed communities on both artificial materials differed from the natural algal-dominated benthic community of the same area, suggesting that the recovery of rocky shore communities on temperate cliffs is time consuming.     

Natural physical disturbance and predation: their importance in structuring a marine sessile community

Settling panels were used to determine the distribution of sessile organisms. Experimental and control cages were located at two study sites that were of different physical exposures. Twenty-nine species of sessile organisms were recorded for the two sites. Data on the presence or absence of each species, for each sample, were classified. The resulting hierarchy showed natural physical disturbance to be the major structuring factor in the community. Predation was of less importance in the physically stressed study site and had no effect on species distribution in the protected site.     

Development of a subtidal epifaunal community at the island of Helgoland

Epifaunal community development on plexiglas panels submerged in Helgoland Harbour (North Sea) was observed over one year. The course of colonization is described, and some data are presented on autecology, reporduction, and growth rate of particular species. About three months after initial settlement, conditions of coexistence in a mixed barnacle-ascidian community began to change increasingly due to heavy competition for space. The colonial speciesBotryllus schlosseri proved to be potentially dominant. Shortly before it attained monopolization by replacing barnacles (mainlyElminius modestus), a major physical disturbance eliminated the fast growing ascidian. The roles of physical factors, of biological interactions, and of historical events in community development are discussed in context with succession theory and other concepts evolved more recently. It is concluded that succession-like processes can occur in subtidal fouling communities, but there the existence of a globally stable climax is unlikely. Generally the concept of multistable points seems to be better applicable to marine ecosystems than that of succession in the classical sense. There is considerable need for further natural history observations and experiments as an empirical basis for current theory and modelling.     

Possible latitudinal clines in Antarctic intertidal and subtidal zone communities encrusting ephemeral hard substrata

Aim Encrusting faunal communities on rocks were examined from Southern Ocean intertidal and subtidal (6 m) zones to investigate potential change with latitude. Location The site locations were South Georgia (54°S [sub-Antarctic]), Signy Island (60.5°S [maritime Antarctic]) and Adelaide Island (68°S [high Antarctic]). Methods The number of taxa, degree of colonization and bryozoan growth and mortality were measured. Results The communities present were relatively simple in terms of composition and interactions. Spirorbid polychaetes (Annelida) and cheilostomatid bryozoans were the principal components. The community simplicity and dominance of unitary (compared to colonial) fauna increased with latitude. Mean annual mortality of subtidal cheilostomatids also significantly increased with latitude. The number of bryozoan species and potential aspects of community complexity increased with latitude. The growth of the bryozoan Inversiula nutrix increased with latitude reversing the general trend of decreased growth rate. Conclusions These findings suggest that disturbance may significantly increase with latitude within the Southern Ocean. The apparent reversal of typical latitudinal growth trends is probably a result of classical island biogeography, with species decreasing away from a centre of high diversity.     

Effect of colony size on the competitive outcome of encrusting colonial organisms

The relative importance of colony size ratio of interacting species was studied in Tomioka Bay, Japan. Six encrusting colonial species belonging to the following three different taxonomic groups were tested: Ascidia (three species), Bryozoa (two) and Porifera (one). Colonies of these organisms were grown in the community of sessile organisms developed on plastic panels. Logistic regression analysis was carried out to determine the effect of size ratio on the competitive outcome of interacting colonies. The results between all possible combinations among these six species did not show a significant size effect in competitive outcome (i.e. a larger colony size did not always prove important in the success of a competitive interaction with smaller colonies of other species). On the contrary, competitive success depends on the types of species interacting. Certain species such asDidemnum moseleyi (ascidian) andHaliclona sp. (sponge), in spite of being smaller in colony size, won in competitive interactions with larger colonies of other species such asDiplosoma mitsukurii (ascidian) andWatersipora subovoidea (bryozoan). These results contradict the one reported earlier: that the larger the colony size, the more chance the colony will have to win in competitive interactions.     

The influence of resident adults on recruitment: a comparison to settlement

For species recruiting into established sessile communities, the adult colonies and individuals already present form a significant part of the environment and have the potential to alter both larval settlement rates and post-settlement mortality. Settlement rates can be reduced by predation on larvae, by the removal or addition of substratum space, or by stimulation or prohibition of larvae from settling on adjacent substratum. Once attached, the recruiting individual can still be influenced by predation or overgrowth by residents, by the added physical structure for firmer attachment, or by being camouflaged from motile predators. To examine those processes by which residents affect recruitment we exposed experimental substrata with three densities of adults of a single species at a site in eastern Long Island Sound, USA for a 1-wk period. Seven different species of common invertebrates were used in nine separate experiments. The major effect of most resident species was the usurpation of space and the restricting of recruitment to adjacent unoccupied areas. This was particularly true for resident ascidians and bryozoans, but less so for barnacles and oysters. In fact several species recruited in higher densities on or next to oysters and barnacles. Comparison to 1-day settlement experiments indicated that the encrusting ascidian species Diplosoma and possibly Botryllus reduced recruitment relative to settlement, probably by overgrowing newly-settled individuals. However, in the presence of most resident species, recruitment patterns were not greatly different from settlement patterns, indicating that the effects of the attached community on recruitment may result from influences on settlement.     

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.     

Plant community response to loss of large herbivores: comparing consequences in a South African and a North American grassland

Loss of biodiversity poses one of the greatest threats to natural ecosystems throughout the world. However, a comprehensive understanding of the impacts of species losses from upper trophic levels is still emerging. Here we compare the impacts of large mammalian herbivore species loss on grassland plant community structure and composition in a South African and North American grassland. Herbaceous plant communities were surveyed at sites without large mammalian herbivores present and at sites with a single species of herbivore present in both locations, and additionally at one site in South Africa with multiple herbivore species. At both the North American and South African locations, plant communities on sites with a single herbivore species were more diverse and species rich than on sites with no herbivores. At the multi-herbivore site in South Africa, plant diversity and richness were comparable to that of the single herbivore site early in the growing season and to the no herbivore site late in the growing season. Analyses of plant community composition, however, indicated strong differences between the multi-herbivore site and the single and no herbivore sites, which were more similar to each other. In moderate to high-productivity ecosystems with one or a few species of large herbivores, loss of herbivores can cause a significant decrease in plant diversity and richness, and can have pronounced impacts on grassland plant community composition. In ecosystems with higher herbivore richness, species loss may also significantly alter plant community structure and composition, although standard metrics of community structure may obscure these differences.