Marine Macroecology
edited by Jon D. Witman and Kaustuv Roy
University of Chicago Press, 2009
Cloth: 978-0-226-90411-5 | Paper: 978-0-226-90412-2 | Electronic: 978-0-226-90414-6
DOI: 10.7208/chicago/9780226904146.001.0001


Pioneered in the late 1980s, the concept of macroecology—a framework for studying ecological communities with a focus on patterns and processes—revolutionized the field. Although this approach has been applied mainly to terrestrial ecosystems, there is increasing interest in quantifying macroecological patterns in the sea and understanding the processes that generate them. Taking stock of the current work in the field and advocating a research agenda for the decades ahead, Marine Macroecology draws together insights and approaches from a diverse group of scientists to show how marine ecology can benefit from the adoption of macroecological approaches.

Divided into three parts, Marine Macroecology first provides an overview of marine diversity patterns and offers case studies of specific habitats and taxonomic groups. In the second part, contributors focus on process-based explanations for marine ecological patterns. The third part presents new approaches to understanding processes driving the macroecolgical patterns in the sea. Uniting unique insights from different perspectives with the common goal of identifying and understanding large-scale biodiversity patterns, Marine Macroecology will inspire the next wave of marine ecologists to approach their research from a macroecological perspective.


Jon D. Witman is professor in the Department of Ecology and Evolutionary Biology at Brown University. Kaustuv Roy is professor in the Section of Ecology, Behavior and Evolution at the University of California, San Diego.


“This book is a wonderful tour de force introduction to marine macroecology through environments and across taxa. It is aimed at students and established ecologists, and it should strongly remind us that small-scale experimental ecology is not the only way to do sound ecological science.”

— Martin Zuschin, University of Vienna, Marine Ecology




- James W. Valentine
DOI: 10.7208/chicago/9780226904146.003.0001
[environmental framework, biodiversity, marine biodiversity, taxa, latitudinal diversity, gradient, sea]
This chapter reviews the environmental framework of the sea and its relation to major biodiversity patterns, and focuses on some of the leading hypotheses proposed to explain those relationships. The major biotic patterns are global or regional, and it is plausible to relate them to global or regional patterns of ecological parameters in the marine environment. Most hypotheses of major biodiversity patterns involve temperature, environmental heterogeneity, trophic resources, and biotic interactions. The chapter deals with such factors as they are distributed in larger environmental partitions, at levels from oceanic to subprovincial, in which correlates of local and regional diversities for distinctive faunal elements and ecological settings are examined in much more detail by workers with expertise in those systems. (pages 3 - 28)
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- William K. W. Li
DOI: 10.7208/chicago/9780226904146.003.0002
[microbial, plankton, diversity, macroecology, phytoplankton]
Patterns in ecology are not easily discerned. Some, such as the allometric scaling of abundance to body size, are clearly evident in the marine plankton. Others are not so obvious, but seem to emerge as boundary constraints when a large number of possibilities are considered. Shifts in the relative importance of bottom-up and top-down control, multiple resource competition, niche complementarity, and intermediate disturbance are examples indicating that biological processes in the pelagos share many characteristics with those elsewhere. Plankton communities are peculiarly shaped by the turbulent hydrodynamic environment, but the causal mechanisms underlying patterns of abundance, distribution, and diversity are general. It is useful to view the plankton, especially the microbial forms, as members at the low end of the space and time ranges in the ocean. In this way, it may be possible to assimilate plankton patterns into the commonalities of life. (pages 29 - 64)
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- Craig R. Mcclain, Michael A. Rex, Ron J. Etter
DOI: 10.7208/chicago/9780226904146.003.0003
[deep sea, macroecology, terrestrial systems, ecogeographic properties, ecological investigation, standing stock]
This chapter presents the basic macroecological features of the deep-sea fauna, including geographic variation in standing stock, species diversity, species ranges, and body size. It also explores the relationships among body size, diversity, and abundance, and between body size and metabolic rate. Whenever possible, the chapter compares these trends to those found in other ecosystems. It concentrates on the deep North Atlantic Ocean, which is by far the most intensively sampled region of the World Ocean. The emphasis is on communities that inhabit the soft-sediment habitats covering most of the seafloor. This chapter provides an excellent summary of biogeographic patterns in deep-sea chemosynthetic habitats. Many of the case studies in the chapter involve mollusks, because their taxonomy and biogeography are relatively well known, owing to the extensive published work of Philippe Bouchet, Anders Waren, John Allen, Howard Sanders, and their colleagues. While the data are limited and geographically restricted, they begin to provide a macroecological context for studying deep-sea assemblages. (pages 65 - 100)
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- Kaustuv Roy, Jon D. Witman
DOI: 10.7208/chicago/9780226904146.003.0004
[latitudinal diversity gradient, taxonomic richness, local richness, marine communities, regional species, local species, marine invertebrate, southern ocean]
Species diversity in the ocean changes along both latitude and longitude as well as with depth. The latitudinal diversity gradient (LDG), with high richness of species and higher taxa in the tropics and declining toward the poles, is considered to be one of the fundamental patterns of biological diversity on the planet. The presence of a latitudinal gradient in taxonomic richness is well established in groups such as marine mollusks, especially in the northern hemisphere, but the trend is less well documented for many other benthic invertebrates. This had led to the obvious question whether a tropical-polar cline in richness is a general pattern in the oceans, especially given the fact that several groups of benthic marine invertebrates show relatively high species richness in the higher latitudes of the southern ocean. It is known that the latitudinal cline in richness holds not just for well-studied invertebrate groups like mollusks but is also present in other groups ranging from crustaceans, bryozoans, epifaunal, invertebrates, and cephalopods to benthic foraminifera, gammaridean amphipods, and sabellid polychaetes. (pages 101 - 121)
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- Enrique Macpherson, Philip A. Hastings, D. Ross Robertson
DOI: 10.7208/chicago/9780226904146.003.0005
[marine environments, regional diversity, macroecological, marine fish, coral-reef fishes]
This chapter reviews current knowledge about patterns in the distribution and diversity of marine fish species in the context of the enormous spatial and temporal variation that results from oceanographic variability. This chapter also discusses processes commonly implicated in controlling the distribution and regional diversity of marine fishes. Broader study of the genetics of putative species, especially widespread coral reef species and open-ocean species may reveal significantly greater cryptic species diversity. In addition, recent analysis indicates no decline in the rate of morphospecies descriptions of tropical shore-fishes in recent decades. The chapter is also couched within the prevailing concept of species. Broader application of a phylogenetic species concept, recently advocated for coral-reef fishes, would result in an increase in overall species diversity and a concomitant decrease in average species range. (pages 122 - 152)
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- Bernabé Santelices, John J. Bolton, Isabel Meneses
DOI: 10.7208/chicago/9780226904146.003.0006
[macroalgal, global-scale pattern, latitudinal gradient, floras, biomass, marine algae]
This chapter discusses the evidence that allows for a first characterization of macroecological patterns in macroalgae. There is little doubt that macroalgal species richness does not always increase toward the tropics, as is the case for most kinds of macroscopic organisms in terrestrial and marine habitats. In all the floras examined, there is a tendency for species to have smaller mean latitudinal ranges in the areas of greater species richness, but this is independent of the latitudinal patterns of species richness. For the macroalgae, using coastline length as a surrogate for area, there is a significant correlation between coastline length and species richness. Both local species richness and the productivity and morphological effects of upwelled waters seem determined by regional factors as well as local interactions. Although latitudinal patterns of macroalgal morphologies can be traced in many areas, the explanations of such patterns remain elusive, but there is little understanding of the selective advantages of the different types of macroalgal morphologies. Latitudinal patterns of growth types, on the other hand, suggest increases in clonal species richness in warmer waters. (pages 153 - 192)
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- Sean Nee, Graham Stone
DOI: 10.7208/chicago/9780226904146.003.0007
[plankton, ocean metabolism, biodiversity, molecular evolution]
The paradox of the plankton is an apparently massive violation of the principle of competitive exclusion, or the “one species, one niche” requirement for coexistence. It is quite extraordinary, the extent to which this world view of homogenous water with little opportunity for niche diversification has been completely overturned by the discoveries of recent years, so it is appropriate to return for another look at the paradox of the plankton. In addition to the existence of many new discoveries, another reason to revisit the paradox is the current interest in the neutral theory of biodiversity, since such a theory was presented as a resolution of the paradox many years ago. The neutral theory of biodiversity was originally proposed as a candidate explanation of tropical tree diversity. The enormous tree diversity in the tropical forests seems to present a similar puzzle to the ecologist, as do the plankton. (pages 195 - 204)
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- James J. Leichter, Jon D. Witman
DOI: 10.7208/chicago/9780226904146.003.0008
[organism, communities, oceanographic influence, marine, ocean basin]
This chapter provides an overview of known and potential links between oceanographic forcing mechanisms, biological processes acting at scales of individual organisms to communities, and the resulting macroecological patterns. It provides an overview of a range of physical processes that may influence and modify macroecological patterns, and suggests promising avenues for future research. The chapter pays particular attention to phenomena acting on large spatial scales of ocean basins, as it is at these scales that macroecological patterns are most likely to occur and be recognized. The chapter suggests that it is time to broaden the perspective on oceanographic forcing of marine populations, communities, and ecosystems. (pages 205 - 226)
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- Steven D. Gaines, Sarah E. Lester, Ginny Eckert, Brian P. Kinlan, Rafe Sagarin, Brian Gaylord
DOI: 10.7208/chicago/9780226904146.003.0009
[dispersal distance, sea, marine systems, macroecological characteristics, geographic ranges]
Explorations of range edges have been historically somewhat divorced from studies of geographical variation in population size. The emerging data on patterns of abundance across entire species' ranges and modeling work predicting patterns of abundance under different dispersal and oceanographic scenarios both suggest that it could be fruitful to further study the potential for dispersal to influence geographic patterns of population size. In examining these three issues, the chapter stresses the value of a multifaceted approach to macroecological studies. These insights were gained by a combination of empirical documentation of large-scale patterns for different life history groups, modeling efforts, and critical examinations of mechanistic hypotheses. (pages 227 - 249)
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- Andrew Clarke
DOI: 10.7208/chicago/9780226904146.003.0010
[temperature, organism, surface waters, hemispheres, global pattern, marine macroecology]
Temperature is one of the most important physical factors affecting organisms. Temperature, typically combined with other environmental factors such as climate, has long been regarded as a key factor regulating the diversity of organisms, and from the time of the earliest naturalists the perceived favorableness of climate has been regarded as a key factor in determining how many and what kinds of organisms could live in a given place. Although many of these ideas were developed for the terrestrial realm, they have frequently been extended to the sea. Marine environments have many environmental features in common with the terrestrial realm, including movement, light, photoperiod, and temperature, but there are also some more specifically aquatic factors such as nutrient concentration and salinity. Most attention in this chapter, however, has been directed at temperature. (pages 250 - 278)
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- Sean R. Connolly
DOI: 10.7208/chicago/9780226904146.003.0011
[species richness, macroecological theory, marine species, diversity gradients, geographical variation, environmental mechanism, sea-level fluctuation]
The practice of attributing geographical variation in species richness to coincident variation in environmental conditions has a long history. In the marine realm, species richness gradients have been attributed to some of the same variables that have been invoked on land, such as average temperatures, seasonal climatic variability, habitat availability, and productivity. Explanations for particular patterns in the marine realm (e.g., longitudinal gradients in the Indo-Pacific) have also invoked mechanisms specific to those contexts, such as ocean currents, and vicariance events due to Plio-Pleistocene sea-level fluctuations. (pages 279 - 309)
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- Derek P. Tittensor, Boris Worm, Ransom A. Myers
DOI: 10.7208/chicago/9780226904146.003.0012
[macroecology, ecosystem, biodiversity, marine species, marine system, ocean]
The oceans have been used as an important source of food and materials for much of human history and prehistory, and until relatively recently were viewed as inexhaustible. Fishing has an effect at many levels in addition to that of the population and the species. Life-history changes, habitat alteration, and changes in community structure may also result, and these can all lead to detectable changes in the macroecology of the marine environment. This chapter takes a broad view of macroecology as being the effect of local and small-scale processes upon large-scale patterns in the marine environment, and the analysis and utilization of these large-scale statistical patterns to infer ecological change from local to global scales. The new tools and analytical processes that a macroecological viewpoint provides enable us to view the interactions and synergies between biological and ecological processes in multiple dimensions. (pages 310 - 338)
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- Jon D. Witman, Kaustuv Roy
DOI: 10.7208/chicago/9780226904146.003.0013
[macroecology, marine system, taxa, allometric scaling, marine invertebrates, phytoplankton, species diversity]
The goal of macroecology is to gain insight into large scale ecological phenomena mainly through rigorous statistical analyses of information on species abundance, diversity, body size, and range size. Over the last decade, such analyses have revealed a number of large-scale trends that hold across taxa and across the land-sea boundary. The allometric scaling relationship between body size and abundance has been quantified in a wide variety of groups, from higher plants to marine invertebrates and phytoplankton, although debate continues over whether there is a universal exponent. Species diversity of a variety of taxa, from plants and terrestrial vertebrates to corals and marine mollusks, has been shown to scale positively with available energy. Such statistical trends are being increasingly documented for both marine and terrestrial organisms. (pages 341 - 356)
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- Eric Sanford, Mark D. Bertness
DOI: 10.7208/chicago/9780226904146.003.0014
[macroecology, abiotic condition, biotic condition, marine species, latitudinal variation, species interaction]
Using small-scale field experiments, ecologists have demonstrated convincingly that species interactions often influence local community structure. Rigorously examining such large-scale processes, however, has created logistical and conceptual challenges. The developing discipline of macroecology has encouraged ecologists to address these challenges by recognizing the value of a broader spatial and temporal perspective. Among the most powerful tools to emerge has been the comparative-experimental approach. This approach uses identical experiments replicated along a broad spatial gradient to test the potential influence of changing abiotic and biotic conditions that are not amenable to manipulation. (pages 357 - 391)
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- Sean D. Connell, Andrew D. Irving
DOI: 10.7208/chicago/9780226904146.003.0015
[marine ecologists, macroecology, local-regional patterns, scale-dependent patterns, structured assessments, rocky coasts]
People are becoming increasingly aware that ecologists work on local patterns that are likely to represent the outcome of special and unique events incorporate variation from broad to local scales. For those interested in the discovery of detail, local patterns appear to have an infinite supply. For those interested in the existence of generalities, it is encouraging to observe that general patterns and responses can emerge from complexity on local scales. This realization, together with the need for a renewed effort for carefully planned sampling and experimentation across broad scales, suggests that there are opportunities to test some of the more interesting questions about the relative importance of processes across vast parts of the world's coast. (pages 392 - 418)
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