Richard W. Osman

Linking climate change and biological invasions: Ocean warming facilitates nonindigenous species invasions.

The spread of exotic species and climate change are among the most serious global environmental threats. Each independently causes considerable ecological damage, yet few data are available to assess whether changing climate might facilitate invasions by favoring introduced over native species. Here, we compare our long-term record of weekly sessile marine invertebrate recruitment with interannual variation in water temperature to assess the likely effect of climate change on the success and spread of introduced species. For the three most abundant introduced species of ascidian (sea squirt), the timing of the initiation of recruitment was strongly negatively correlated with winter water temperature, indicating that invaders arrived earlier in the season in years with warmer winters. Total recruitment of introduced species during the following summer also was positively correlated with winter water temperature. In contrast, the magnitude of native ascidian recruitment was negatively correlated with winter temperature (more recruitment in colder years) and the timing of native recruitment was unaffected. In manipulative laboratory experiments, two introduced compound ascidians grew faster than a native species, but only at temperatures near the maximum observed in summer. These data suggest that the greatest effects of climate change on biotic communities may be due to changing maximum and minimum temperatures rather than annual means. By giving introduced species an earlier start, and increasing the magnitude of their growth and recruitment relative to natives, global warming may facilitate a shift to dominance by nonnative species, accelerating the homogenization of the global biota.

The Establishment and Development of a Marine Epifaunal Community

Because of the nature of their substratum, the sessile invertebrate species of the marine epifaunal community living on rocks occur in discrete patterns of distribution. The rocks are finite patches or habitat islands with a limited space for colonization and growth. Such a system is ideal for studying the parameters affecting the distribution of species within a community. Also, because of the small size and immobility of the adults, the system is also ideal for studying the pattern of change in species composition and diversity within a community. This study used multiple series of manipulated experimental plates, which both duplicated natural rock surfaces and could be compared with samples of the rocks, to investigate the developmental and distributional processes of this community. Five major factors were found to be important to both the development of the community and its distribution on the rocks: (1) the selectivity of the metamorphosing larvae as to site of attachment; (2) the seasonal fluctuat...

Biodiversity, Invasion Resistance, and Marine Ecosystem Function: Reconciling Pattern and Process

A venerable generalization about community resistance to invasions is that more diverse communities are more resistant to invasion. However, results of experimental and observational studies often conflict, leading to vigorous debate about the mechanistic importance of diversity in determining invasion success in the field, as well as other eco- system properties, such as productivity and stability. In this study, we employed both field experiments and observational approaches to assess the effects of diversity on the invasion of a subtidal marine invertebrate community by three species of nonindigenous ascidians (sea squirts). In experimentally assembled communities, decreasing native diversity in- creased the survival and final percent cover of invaders, whereas the abundance of individual species had no effect on these measures of invasion success. Increasing native diversity also decreased the availability of open space, the limiting resource in this system, by buffering against fluctuations in the cover of individual species. This occurred because temporal patterns of abundance differed among species, so space was most consistently and completely occupied when more species were present. When we held diversity constant, but manipulated resource availability, we found that the settlement and recruitment of new invaders dramatically increased with increasing availability of open space. This suggests that the effect of diversity on invasion success is largely due to its effects on resource (space) availability. Apart from invasion resistance, the increased temporal stability found in more diverse communities may itself be considered an enhancement of ecosystem func- tion. In field surveys, we found a strong negative correlation between native-species richness and the number and frequency of nonnative invaders at the scale of both a single quadrat (25 3 25 cm), and an entire site (50 3 50 m). Such a pattern suggests that the means by which diversity affects invasion resistance in our experiments is important in determining the distribution of invasive species in the field. Further synthesis of mechanistic and ob- servational approaches should be encouraged, as this will increase our understanding of the conditions under which diversity does (and does not) play an important role in deter- mining the distribution of invaders in the field.

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.

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.

Local control of recruitment in an epifaunal community and the consequences to colonization processes

We found that recruitment, abundance, and dominance within two subtidal epifaunal communities in southern New England, USA persist year after year over large areas of the bottom. This long-term persistence in both dominance and recruitment is not expected in such an open system with disturbances continually creating open patches for recruiting larvae whose identity and abundances change both temporally and spatially. We suggest that the persistence results from strong local control of recruitment that overrides any variability in larval production and dispersal of species from outside a site. Although local dynamics that control persistence involve all life-stages, we found that intense predation on post-settlement individuals has drastic effects. This predation alters the relative abundances of recruits, prevents the invasion of some species, and allows others to dominate. In addition, epifaunal communities are often dominated by species producing short-lived, poorly dispersed larvae. The continued local recruitment of these species at a given site can contribute to the long-term persistence of dominants already present. Based on these observations, we suggest that a system of locally reproducing, self-sustaining populations coupled with strong local environmental differences (e.g. predation on recruits) limiting the invasion of other species may better represent some subtidal benthic communities than a system with widely-dispersed larvae, recruitment dominated by production outside the community, disturbance creating continual changes in dominance, and little long-term persistence.

The Influence of Seasonality and Stability on the Species Equilibrium

The species equilibrium theory has proven to be a useful framework in which to analyze the distribution and abundance of species, especially for islands and habitats distributed as islands. In this study, the distribution of marine epifaunal invertebrates on rocks has been interpreted using this model. The relationship between area and the number of species present is sustained within this community. However, the increase in the number of species with increasing area appears to result from increasing immigration rather than decreasing extinction, as predicted by the model. Two factors have a major effect on the species equilibrium: seasonality and substrate stability. The seasonality of the climate causes the immigration rate to change seasonally. Colonization curves are dependent on the time of initial immersion of the substrate and vary in shape. However, with time, all converge. A simple model which allows immigration to fluctuate seasonally is developed. It predicts both the initial difference in the ...

Stronger predation in the tropics shapes species richness patterns in marine communities

Species interactions are widely assumed to be stronger at lower latitudes, but surprisingly few experimental studies test this hypothesis, and none ties these processes to observed patterns of species richness across latitude. We report here the first experimental field test that predation is both stronger and has a disproportionate effect on species richness in the tropics relative to the temperate zone. We conducted predator-exclusion experiments on communities of sessile marine invertebrates in four regions, which span 32 degrees latitude, in the western Atlantic Ocean and Caribbean Sea. Over a three-month timescale, predation had no effect on species richness in the temperate zone. In the tropics, however, communities were from two to over ten times more species-rich in the absence of predators than when predators were present. While micro-and macro-predators likely compete for the limited prey resource in the tropics, micropredators alone were able to exert as much pressure on the invertebrate communities as the full predator community. This result highlights the extent to which exposure to even a subset of the predator guild can significantly impact species richness in the tropics. Patterns were consistent in analyses that included relative and total species abundances. Higher species richness in the absence of predators in the tropics was also observed when species occurrences were pooled across two larger spatial scales, site and region, demonstrating a consistent scaling relationship. These experimental results show that predation can both limit local species abundances and shape patterns of regional coexistence in the tropics. When preestablished diverse tropical communities were then exposed to predation for different durations, ranging from one to several days, species richness was always reduced. These findings confirmed that impacts of predation in the tropics are strong and consistent, even in more established communities. Our results offer empirical support for the long-held prediction that predation pressure is stronger at lower latitudes. Furthermore, we demonstrate the magnitude to which variation in predation pressure can contribute to the maintenance of tropical species diversity.

Patterns of species diversity; fact or artifact?

The pattern observed in nature has often served as a compelling motivation for seeking the underlying processes which are assumed to control the pattern. Spatial and temporal patterns of species diversity are the most commonly observed and described from the study of recent and paleocommunities. These patterns include those found within a local or discrete region such as succession and changes along an environmental gradient and interregional or global patterns such as latitudinal diversity gradients and comparisons of the deep-sea and the continental shelf faunas. Many hypotheses have been advanced to explain these patterns in terms of differences in the biology of species adapted to different or changing environments. However, a more simple explanation is possible: that these patterns do not result from any specific biologic processes. We examine this possibility and show that succession can result from probabilistic immigration and local extinction of species, diversity gradients can result from probabilistic disturbance in naturally patchy environments, and that latitudinal gradients can be simply a function of the shape of the earth. As long as such diversity patterns can be explained independent of any specific biologic processes, they offer no test of the importance of these processes, either ecologically or evolutionarily.

Pathways of arsenic uptake and incorporation in estuarine phytoplankton and the filter-feeding invertebrates Eurytemora affinis, Balanus improvisus and Crassostrea virginica

Arsenic uptake from water and from phytoplankton was followed in the copepod Eurytemora affinis and the barnacle Balanus improvisus collected from the Patuxent River estuary, Chesapeake Bay, eastern coast of the USA in 1987, and in the oyster Crassostrea virginica obtained from a hatchery on the shore of Chesapeake Bay in 1987. Dissolved arsenic was readily taken up by phytoplankton and by shell material of B. improvisus and C. virginica; however, no dissolved arsenic was incorporated into the invertebrate tissues. When E. affinis, B. improvisus and C. virginica were fed phytoplankton containing elevated arsenic contents, significant arsenic incorporation occurred. Juvenile B. improvisus incorporated relatively more arsenic than adults of all three species. Compared to the 100 to 200% increase in arsenic content by phytoplankton exposed to dissolved arsenic, the 25 to 50% increase in these invertebrate species via trophic transfer is relatively small. Even though the trophic pathway for arsenic transfer is the major one for higher trophic levels within an ecosystem, the potential for direct arsenic impact to trophic levels other than phytoplankton appears to be minimal.