Paul K. Dayton

Competition, Disturbance, and Community Organization: The Provision and Subsequent Utilization of Space in a Rocky Intertidal Community

An understanding of community structure should be based on evidence that the growth and regulation of the component populations in the community are affected in a predictable manner by natural physical disturbances and by interactions with other species in the community. This study presents an experimental evaluation of the effects of such disturbances and competitive interactions on populations of sessile organisms in the rocky intertidal community, for which space can be demonstrated to be the most important limiting resource. This research was carried out at eight stations on the Washington coastline which have been ranked according to an exposure/desiccation gradient and subjected to comparable manipulation and observation. Physical variables such as wave exposure, battering by drift logs, and desiccation have important effects on the distribution and abundance of many of the sessile species in the community. In particular, wave exposure and desiccation have a major influence on the distribution patterns of all the algae and of the anemone Anthopleura elegantissima. The probability of damage from drift logs is very high in areas where logs have accumulated along the intertidal. Log damage and wave exposure have complementary effects in the provision of free space in a mussel bed, as wave shock enlarges a patch created by log damage by wrenching the mussels from the substratum at the periphery of the bare patch. Competition for primary space results in clear dominance hierarchies, in which barnacles are dominant over algae. Among the barnacles, Balanus cariosus is dominant over both B. glandula and Chthamalus dalli; B. glandula is dominant over C. dalli. The mussel Mytilus californianus requires secondary space (certain algae, barnacles, or byssal threads) for larval settlement, but is capable of growing over all other sessile species and potentially is the competitive dominant of space in the community.

Ecosystem-Based Fishery Management

Ecosystem-based fishery management (EBFM) is a new direction for fishery management, essentially reversing the order of management priorities so that management starts with the ecosystem rather than a target species. EBFM aims to sustain healthy marine ecosystems and the fisheries they support. Pikitch et al . describe the potential benefits of implementation of EBFM that, in their view, far outweigh the difficulties of making the transition from a management system based on maximizing individual species.

EXPERIMENTAL EVALUATION OF ECOLOGICAL DOMINANCE IN A ROCKY INTERTIDAL ALGAL COMMUNITY

The mechanisms by which various species exert influence disproportionate to their abundance or mass on the structure of a lower intertidal algal community were evaluated experimentally. These functional roles were evaluated experimentally. These functional roles were evaluated by various controlled manipulations at seven stations along the Washington coastline ranked according to an exposure/desiccation gradient. The algae were divided into three categories: canopy species, which grow above the other species and apparently succeed in competitively dominating the light resources as demonstrated by algal blooms following their removal; obligate understory species, which die after the canopy species are removed; and fugitive species, which are quick to colonize new space. Ecological dominance was exerted in areas of moderate wave exposure by Hedophyllum sessile, which competitively displaces a large number of fugitive algal species and which furnishes a protected habitat for many obligate understory algae that die or defoliate after the removal of Hedophyllum. Hedophyllum loses this dominance in the most exposed areas, although such sites apparently represent its physiologically optimal habitat, because in these areas it is out—competed by Laminaria setchellii and Lessoniopsis littoralis. In these wave exposed habitats Lessoniopsis was demonstrated to exert a strong competitive dominance over all the other species in the association. The molluscan herbivores were not observed to express any measurable effects on the recruitment or survival of the algae. However, the echinoid Strongylocentrotus purpuratus often overexploits its prey and has a pronounced influence on most of the algal species. In this respect S. purpuratus enjoys an important community role singular among the many herbivores. Similarly, Pycnopodia helianthoides and Anthopleura xanthogrammica are disproportionately important carnivores, because their predation on Strongylocentrotus, clearing large areas of urchins, results in patches in which algal succession follows. The rate of algal succession following removal of the dominant algal species or of Strongylocentrotus is proportional to the degree of wave exposure. The Hedophyllum canopy recovery at the Eagle Point area of San Juan Island, a site exposed to relatively little wave action and thus high levels of desiccation, was relatively slow, with only 10%—26% cover reestablished after 3 yr. In contrast, Hedophyllum canopy developed up to 66% cover in only 1 yr in the exposed area of Waadah Island; it then quickly lost its dominance to Laminaria and Lessoniopsis. Algal succession in deeper Portage Head tidepools was found to be relatively slow with no clear dominance expressed after 5 yr.

Patch dynamics and stability of some California kelp communities

This paper considers three concepts of stability as they relate to the dynamics of distinctive patch types of algal canopy guilds in southern and central California kelp communities: (1) persistence of a patch through more than one generation of the dominant species, which was evaluated by using life tables and observations of patch borders; (2) inertia or the resistance of different patches to invasion or disturbance, which was evaluated by artificially enhancing gametophytes by transplanting sporogenic material, by removing canopy, and by evaluating some important disturbance processes; and (3) resilience or recoverability of a patch following a perturbation sufficient to allow invasion of different species, which was studied by defining some of the mechanisms of successful invasion or succession. By working in distinct habitats in southern (Pt. Loma and Santa Catalina Island) and central (Pt. Piedras Blancas) California, we could evaluate different types of physical stresses as they related to these stability concepts. Taller perennial canopy guilds were dominant competitors for light, but were more susceptible to physical wave stress. Dominance hierarchies in the competition for light appeared to be reversed in areas exposed to increasing wave stress. The main causes of mortality at Pt. Loma were entanglement with storm—dislodged Macrocystis plants and, in some areas, sea urchin grazing. Mortality in central California was due to winter storms. In most cases, distinct patches resisted invasion for >10 yr. The mechanisms of resistance involved (1) competition for light and, possibly, nutrients, and (2) limits to spore dispersal. When succession occurred, it was often mediated by many factors, including seasonality of spore production, which coincided with winter storm—related mortalities; mechanisms of kelp dispersal, which were most effective via drifting plants and fragments of fertile material held against the substrate by invertebrates; and survivorship of gametophytes and small sporophytes, which was influenced by local scour and grazing. Appropriate spatial scales, stability, and succession studies in these kelp communities were determined by the size of the disturbed area, which varied from the free space resulting from detachment of single plants to the free space resulting from catastrophies such as overgrazing or unusual storms. Temporal scales were influenced by seasonality of disturbance and algal reproductive condition and aperiodic episodes of cool, nutrient—rich water advected into the patch. There appeared to be conflicting morphological adaptations of the canopy guilds: exploitation of light was enhanced at higher canopy levels, whereas the lower canopy levels were better adapted to tolerate stress from wave surge. The adaptations of the algae appeared to form four distinct groups of tactics: (1) ruderals or plants, such as Nereocystis and Desmarestia, with opportunistic life histories; (2) kelps, such as Macrocystis, adapted to exploitative competition for light and nutrients; (3) kelps (Eisenia, Dictyoneurum) adapted to physical stress such as wave surge; and (4) those algae, such as corallines and Agarum, adapted to heavy grazing. Within any given area, the relative patch stability was determined by biological relationships; between areas, the patch stability patterns were attributable to physical differences.

Role of biological disturbance in maintaining diversity in the deep sea

Abstract This paper presents the hypothesis that the maintenance of high species diversity in the deep sea is more a result of continued biological disturbance than of highly specialized competitive niche diversification. Detrital food is the primary resource for most of the deep-sea species, but we suggest that in deposit feeding, most animals would consume available living particles as well as dead. We call this dominant life-style ‘cropping’. Predictable cropping pressure on smaller animals reduces the probability of their competitive exclusion and allows a high overlap in the utilization of food resources. Since cropping pressure is in part proportional to the abundance of the prey, proliferations of individual species are unlikely. Through time many species have accumulated in the deep sea because of speciation and immigration. Extinction rate is low because the biological and physical predictability of the environment has suppressed the possibility of population oscillations. Predictability in food supply for smaller deposit feeders is enhanced by the larger, mobile scavengers which consume and disperse large particles of food which fall to the ocean floor.

Mechanisms of contrast agent destruction

Various applications of contrast-assisted ultrasound, including blood vessel detection, perfusion estimation, and drug delivery, require controlled destruction of contrast agent microbubbles. The lifetime of a bubble depends on properties of the bubble shell, the gas core, and the acoustic waveform impinging on the bubble. Three mechanisms of microbubble destruction are considered: fragmentation, acoustically driven diffusion, and static diffusion. Fragmentation is responsible for rapid destruction of contrast agents on a time scale of microseconds. The primary characteristics of fragmentation are a very large expansion and subsequent contraction, resulting in instability of the bubble. Optical studies using a novel pulsed-laser optical system show the expansion and contraction of ultrasound contrast agent microbubbles with the ratio of maximum diameter to minimum diameter greater than 10. Fragmentation is dependent on the transmission pressure, occurring in over 55% of bubbles insonified with a peak negative transmission pressure of 2.4 MPa and in less than 10% of bubbles insonified with a peak negative transmission pressure of 0.8 MPa. The echo received from a bubble decorrelates significantly within two pulses when the bubble is fragmented, creating an opportunity for rapid detection of bubbles via a decorrelation-based analysis. Preliminary findings with a mouse tumor model verify the occurrence of fragmentation in vivo. A much slower mechanism of bubble destruction is diffusion, which is driven by both a concentration gradient between the concentration of gas in the bubble compared with the concentration of gas in the liquid, as well as convective effects of motion of the gas-liquid interface. The rate of diffusion increases during insonation, because of acoustically driven diffusion, producing changes in diameter on the time scale of the acoustic pulse length, thus, on the order of microseconds. Gas bubbles diffuse while they are not being insonified, termed static diffusion.

Antarctic Soft-Bottom Benthos in Oligotrophic and Eutrophic Environments

The benthos of the east and west sides of McMurdo Sound, Antarctica, is characterized by dramatically different infaunal assemblages. The eutrophic East Sound has higher infaunal densities than almost any other benthic assemblage in the world. In contrast, the oligotrophic West Sound, bathed by currents from beneath the Ross Ice Shelf, has patterns of mobile epifauna and low infauna density similar to bathyl deep-sea communities.

Catastrophic Storms, El Niño, and Patch Stability in a Southern California Kelp Community

Strong winter storms in southern California destroyed most of the canopy ofthe giant kelp Macrocystis pyrifera but not the patches of understory kelps in the Point Loma kelp forest near San Diego. Subsequent massive recruitment of Macrocystis juveniles and adults that survived the storms had low survival in the summer during the California El Ni�o of 1983. The combined disturbance may have long-lasting structural consequences for this community because, once established, the understory patches can resist invasion by Macrocystis.

Feeding Behavior of Asteroids and Escape Responses of their Prey in the Puget Sound Region

Observations were made with scuba on the diet and behavior of 18 species of undisturbed sea stars in their natural habitats along the shores of Washington state through all seasons. Some sea stars are specialists. Hippasteria spinosa feeds almost exclusively on a sea pen; Solaster stimpsoni eats holothurians; full—sized Orthasterias koehleri consume the venerid clam Humilaria; Solaster dawsoni eats its congener S. stimpsoni. Others show remarkably variable diets. In different habitats, Dermasterias imbricata specializes on either anemones, holothurians or sea pens, but within these habitats its diet is consistent throughout the year. The diet of Mediaster aequalis varies with both habitat and season. Pycnopodia helianthoides feeds on sea urchins on rocky substrata but digs clams from sand and cobble. Many other sea stars, including Luidia foliolata, Pteraster tesselatus, Pisater ochraceus, Evasterias troschelii and Leptasterias hexactis, are quite generalized in their diet and, though often demonstrating preferences in laboratory studies, will feed on a variety of prey determined largely by relative abundance of prey species in the particular habitat. The areal and seasonal variation in the diet of sea stars coupled with the reluctance of some to eat any prey in the laboratory makes extension of laboratory observations to the field diet suspect. Laboratory observations can be used to provide a more detailed understanding of field observations. Several previously undescribed behavioral mechanisms of food capture in asterioids and escape or defense responses or prey are described: Orthasterias pulls chips of shell off Humilaria until an opening is made, thus allowing exploitation of a prey species unavailable to other clam—eating asteroids. A number of sea stars, such as Luidia, Hippasteria, Mediaster, Pisaster brevispinus, Orthasterias and Pycnopodia dig into mud, sand or cobble substrata to obtain infaunal prey. The locomotory patterns of S. dawsoni and Crossaster papposus tend to allow encounter with their prey in such a way as to reduce the effectiveness of the escape response. A pushing response by Solaster stimpsoni and autotomy in Pycnopodia and Evasterias are thought to be effective responses to S. dawsoni predation. The similar swimming responses to two anemones, a nudibranch, and a holothurian are discussed in relation to asteroid predation. The avoidance responses that many invertebrates show to asteroids are correlated with predator—prey relationships. Two mechanisms may obscure this correlation. The very success of the response may effectively remove the responding organism from the predators diet. Thus scallops, although abundant, form a very inconspicuous portion of starfish diet. Also biochemical similarities between the predator and other organisms may make a prey species unable to distinguish only its predator. Thus Pycnopodia will move rapidly away from Crossaster and Solaster stimpsoni as well as from S. dawsoni, although the latter is its only asteroid predator. The similarities may be based either on a close taxonomic relationship with the predator, or on the diet of the other organism including species closely related to the responding prey species.

Principles for the Conservation of Wild Living Resources

We describe broadly applicable principles for the conservation of wild living resources and mechanisms for their implementation. These principles were engendered from three starting points. First, a set of principles for the conservation of wild living resources (Holt and Talbot 1978) required reexamination and updating. Second, those principles lacked mechanisms for implementation and consequently were not as effective as they might have been. Third, all conservation problems have scientific, economic, and social aspects, and although the mix may vary from problem to problem, all three aspects must be included in problem solving. We illustrate the derivation of, and amplify the meaning of, the principles, and discuss mechanisms for their implementation. The principles are: Principle I. Maintenance of healthy populations of wild living resources in perpetuity is inconsistent with unlimited growth of human consumption of and demand for those resources. Principle II. The goal of conservation should be to secure present and future options by maintaining biological diversity at genetic, species, population, and ecosystem levels; as a general rule neither the resource nor other components of the ecosystem should be perturbed beyond natural boundaries of variation. Principle III. Assessment of the possible ecological and sociological effects of resource use should precede both proposed use and proposed restriction or expansion of ongoing use of a resource. Principle IV. Regulation of the use of living resources must be based on understanding the structure and dynamics of the ecosystem of which the resource is a part and must take into account the ecological and sociological influences that directly and indirectly affect resource use. Principle V. The full range of knowledge and skills from the natural and social sciences must be brought to bear on conservation problems. Principle VI. Effective conservation requires understanding and taking account of the motives, interests, and values of all users and stakeholders, but not by simply averaging their positions. Principle VII. Effective conservation requires communication that is interactive, reciprocal, and continuous. Mechanisms for implementation of the principles are discussed.