Robert T. Paine

Food Web Complexity and Species Diversity

It is suggested that local animal species diversity is related to the number of predators in the system and their efficiency in preventing single species from monopolizing some important, limiting, requisite. In the marine rocky intertidal this requisite usually is space. Where predators capable of preventing monopolies are missing, or are experimentally removed, the systems become less diverse. On a local scale, no relationship between latitude (10⚬ to 49⚬ N.) and diversity was found. On a geographic scale, an increased stability of annual production may lead to an increased capacity for systems to support higher-level carnivores. Hence tropical, or other, ecosystems are more diverse, and are characterized by disproportionately more carnivores.

Trophic Downgrading of Planet Earth

Until recently, large apex consumers were ubiquitous across the globe and had been for millions of years. The loss of these animals may be humankind’s most pervasive influence on nature. Although such losses are widely viewed as an ethical and aesthetic problem, recent research reveals extensive cascading effects of their disappearance in marine, terrestrial, and freshwater ecosystems worldwide. This empirical work supports long-standing theory about the role of top-down forcing in ecosystems but also highlights the unanticipated impacts of trophic cascades on processes as diverse as the dynamics of disease, wildfire, carbon sequestration, invasive species, and biogeochemical cycles. These findings emphasize the urgent need for interdisciplinary research to forecast the effects of trophic downgrading on process, function, and resilience in global ecosystems.

Food webs : linkage, interaction strength and community infrastructure

It seems particularly opportune to discuss food webs and evolving views on their structure here for both their genesis and first modern treatment (Elton 1927) and much of their later development (May 1973; Pimm & Lawton 1978) has a decidedly British accent to it. The central significance of webs is derived from the fact that the links between species are often easily identified and the resultant trophic scaffolding provides a tempting descriptor of community structure. If this structure is in any fashion related to the persistence of natural communities or their stability, however defined, then we are dealing with issues of vital ecological importance. Eltons views have admirably withstood the tests of time. They were especially useful to field biologists, and encouraged the assembly and organization of feeding data into networks of trophically bonded species or higher taxa. The early emphasis was on connectedness per se. Perhaps the first significant deviation from this theme was the development of the trophic dynamic viewpoint of Lindeman (1942) and all subsequent efforts to describe energy transfer and material flow through communities. A second departure, and one I believe to be conceptually richer, was the formalization of the view that web structure and community stability were related (MacArthur 1955). May (1973) in another landmark publication questioned this relationship and called attention to four primary web features: the number of species involved, the nature of their interconnections, the number of connections per species, and the intensity of interaction between web members. This focus has stimulated application to agroecosystems (Southwood & Way 1970), new interpretations of the number of trophic levels (Pimm & Lawton 1977), and a resurgence of interest in the significance of mutualism (Vance 1978). It has not been characterized by stunning breakthroughs, ecological stability remains a frustrating issue, and to a field ecologist, the ties between model and reality at times appear remote; All but ignored in these recent developments is an insightful recognition that trophic pathways might contribute little to ecosystem stability, and that the answers lie in the spatial patterning of the environment (Smith 1972). I wish to return to the basic observations on food webs as a naturalist and experimentalist, and employing an approach advocated by Sir Arthur Tansley (Godwin 1977), ask whether we are modelling their correct properties, and if not, what modifications might be made.

Challenges in the quest for keystones

Mary E. Power is a professor in the Department of Integrative Biology, University of California, Berkeley, CA 94720. David Tilman is a professor in the Department of Ecology, Evolution, and Behavior, University of Minnesota, St. Paul, MN 55108. James A. Estes is a wildlife biologist in the National Biological Service, Institute of Marine Science, University of California, Santa Cruz, CA 95064. Bruce A. Menge is a professor in the Department of Zoology, Oregon State University, Corvallis, OR 97331. William J. Bond is a professor doctor in the Department of Botany, University of Cape Town, Rondebosch 7700 South Africa. L. Scott Mills is an assistant professor in the Wildlife Biology Program, School of Forestry, University of Montana, Missoula, MT 59812. Gretchen Daily is Bing Interdisciplinary Research Scientist, Department of Biological Science, Stanford University, Stanford, CA 94305. Juan Carlos Castilla is a full professor and marine biology head in Facultad de Ciencias Biologicas, Pontificia Universidad Catolica de Chile, Casilla 114-D, Santiago, Chile. Jane Lubchenco is a distinguished professor in the Department of Zoology, Oregon State University, Corvallis, OR 97331. Robert T. Paine is a professor in the Department of Zoology, NJ-15, University of Washington, Seattle, WA 98195. ? 1996 American Institute of Biological Sciences. A keystone species is

Intertidal Landscapes: Disturbance and the Dynamics of Pattern

The mussel Mytilus californianus is a competitive dominant on wave—swept rocky intertidal shores. Mussel beds may exist as extensive monocultures; more often they are an everchanging mosaic of many species which inhabit wave—generated patches or gaps. This paper describes observations and experiments designed to measure the critical parameters of a model of patch birth and death, and to use the model to predict the spatial structure of mussel beds. Most measurements were made at Tatoosh Island, Washington, USA, from 1970—1979. Patch size ranged at birth from a single mussel to 38 m2; the distribution of patch sizes approximates the lognormal. Birth rates varied seasonally and regionally. At Tatoosh the rate of patch formation varied during six winters from 0.4—5.4% of the mussels removed per month. The disturbance regime during the summer and at two mainland sites was 5—10 times less. Annual disturbance patterns tended to be synchronous within 11 sites on one face of Tatoosh over a 10—yr interval, and over larger distances (16 km) along the coastline. The pattern was asynchronous, however, among four Tatoosh localities. Patch birth rate, and mean and maximum size at birth can be used as adequate indices of disturbance. Patch disappearance (death) occurs by three mechanisms. Very small patches disappear almost immediately due to a leaning response of the border mussels (0.2 cm/d). Intermediate—sized patches (<3.0 m2) are eventually obliterated by lateral movement of the peripheral mussels: estimates based on 94 experimental patches yield a mean shrinking rate of 0.05 cm/d from each of two principal dimensions. Depth of the adjacent mussel bed accounts for much of the local variation in closing rate. In very large patches, mussels must recruit as larvae from the plankton. Recovery begins at an average patch age of 26 mo; rate of space occupation, primarily due to individual growth, is 2.0—2.5%/mo. Winter birth rates suggest a mean turnover time (rotation period) for mussel beds varying from 8.1—34.7 yr, depending on the location. The minimal value is in close agreement with both observed and calculated minimal recovery times. Projections of total patch area, based on the model, are accurate to within 5% of the observed. Using a method for determining the age of patches, based on a growth curve of the barnacle Balanus cariosus, the model permits predictions of the age—size structure of the patch population. The model predicts with excellent resolution the distribution of patch area in relation to time since last disturbance. The most detailed models which include size structure within age categories are inconclusive due to small sample size. Predictions are food for large patches, the major determinants of environmental patterns, but cannot deal adequately with smaller patches because of stochastic effects. Colonization data are given in relation to patch age, size and intertidal position. We suggest that the reproductive season of certain long—lived, patch—dependent species is moulded by the disturbance regime. The necessary and vital connection between disturbance which generates spatial pattern and species richness in communities open to invasion is discussed.

Intertidal community structure

SummaryAlong exposed rocky intertidal shorelines of western North America the mussel Mytilus californianus exists as a characteristic, well-defined band. Measurements at Mukkaw Bay and Tatoosh Island, Washington State, suggest that the upper limit to distribution is constant. The lower limit is also predictably constant, as judged by photographs of the same areas taken up to 9 years apart. The band of mussels is formed by larval recruitment to a variety of substrates, especially the filamentous red alga Endocladia muricata. From the settlement site, if the mussels survive a series of predators including the starfish Pisaster ochraceus and a variety of carnivorous gastropods (Thais spp.), the mussles may be washed inward or migrate (be pushed) downward.When Pisaster was removed manually, the zonation pattern changed rapidly. Mussels advanced downward at Mukkaw Bay a vertical distance of 0.85 m in 5 years. No movement was observed on 2 adjacent control sites. At Tatoosh Island a maximum displacement of 1.93 m has been observed in 3 years; the slope there is 40°. Again, there was no change at control sites with Pisaster. At Mukkaw Bay over 25 species of invertebrates and benthic algae are excluded from occupancy of the primary substratum by mussels. The ecological dominance of mussels is discussed; predation is shown to enhance coexistence among potential competitors. A survival curve for Pollicipes polymerus indicates that the time course for interspecific competitive exclusion may be long (76 months). The clarity of the biological interrelationships and the constancy of pattern through time provide no support for the contention that intertidal communities are physically-controlled.

Compounded Perturbations Yield Ecological Surprises

ABSTRACT All species have evolved in the presence of disturbance, and thus are in a sense matched to the recurrence pattern of the perturbations. Consequently, disturbances within the typical range, even at the extreme of that range as defined by large, infrequent disturbances (LIDs), usually result in little long-term change to the systems fundamental character. We argue that more serious ecological consequences result from compounded perturbations within the normative recovery time of the community in question. We consider both physically based disturbance (for example, storm, volcanic eruption, and forest fire) and biologically based disturbance of populations, such as overharvesting, invasion, and disease, and their interactions. Dispersal capability and measures of generation time or age to first reproduction of the species of interest seem to be the important metrics for scaling the size and frequency of disturbances among different types of ecosystems. We develop six scenarios that describe communities that have been subjected to multiple perturbations, either simultaneously or at a rate faster than the rate of recovery, and appear to have entered new domains or “ecological surprises.” In some cases, three or more disturbances seem to have been required to initiate the changed state. We argue that in a world of ever-more-pervasive anthropogenic impacts on natural communities coupled with the increasing certainty of global change, compounded perturbations and ecological surprises will become more common. Understanding these ecological synergisms will be basic to environmental management decisions of the 21st century.

The Pisaster-Tegula Interaction: Prey Patches, Predator Food Preference, and Intertidal Community Structure

The herbivorous gastropod Tegula funebralis is not highly ranked in a food preference hierarchy of its major predator, the starfish Pisaster ochraceus, and exhibits a persistent broad overlap with it in the rocky intertidal zone at Mukkaw Bay, Washington. Observations on Tegula over a 5—yr period indicate that it settles high intertidally, lives there for 5—6 yr, and then tends to migrate lower into contact with Pisaster. Tegula lays down an annual growth line permitting it to be aged and a growth curve constructed. Analysis of relative growth and reproduction indicates that beyond a certain size (16 mm) large individuals perform less well in the upper than those in the lower intertidal zone. Pisaster consumes 25—28% of the adult Tegula per year in the area of spatial overlap, based on analysis of the age structure of 6—17 yr old Tegula, and by direct estimates of the percentage of the standing crop consumed annually. The relationship between Pisaster and sex ratio, relative energy limitation and reproductive output (fitness) of Tegula is discussed for three subpopulations. It is suggested that the implied results of the interaction is typical of that between a major predator and one of its less preferred prey. The prominent zonation exhibited by preferred prey, the observed intimacy of association of predator and less preferred prey, and the zoogeographic homogeneity of the Pacific rocky coastline community are discussed in relation to three intermeshing ecological processes.

Species Introduction in a Tropical Lake: A newly introduced piscivore can produce population changes in a wide range of trophic levels.

Probably in early 1967, a piscivore from South America, Cichla ocellaris, was introduced to Gatun Lake in the Panama Canal Zone. As this predator population spread through the lake, the initial effect was dramatic reductions in almost all secondary consumers. These species reductions produced, in turn, second- and third-order changes at other trophic levels of the ecosystem. The resulting changes in the lake community can be seen best by examining the general Gatun Lake food web. The decrease in numbers of the important planktivore Melaniris has resulted in changes within the zooplankton community, as illustrated by the cladoceran Ceriodaphnia. The tertiary-consumer populations, such as tarpon, black terns, kingfishers, and herons, formerly dependent on small fishes for food, appear less frequently in the Cichla areas of the lake. There has also been, possibly, a resurgence of the local mosquito populations (which are malaria vectors), caused by the reduction in the populations of insect-eating fishes. Even the primary producers may be affected by this introduction. Although at present the Gatun Lake ecosystem is undergoing rapid changes, we anticipate an eventual return to some form of equilibrium. However, it will be some time before we can evaluate the permanence or transience of the many changes produced in the trophic levels by the introduction of a single, top-level predator to this lake system.

Disaster, Catastrophe, and Local Persistence of the Sea Palm Postelsia palmaeformis

Two components of natural disturbance, its local intensity and frequency beyond a threshold level, limit a marine benthic alga to wave-swept shores. Transplant experiments indicate that the limited distribution is not due to physiological restriction. Instead, it requires predictable annual disturbance of moderate intensity for local persistence.