Interactions between organisms are a major determinant of the distribution and abundance of species. Ecology textbooks (e.g., Ricklefs 1984, Krebs 1985, Begon et al. 1990) summarise these important interactions as intra- and interspecific competition for abiotic and biotic resources, predation, parasitism and mutualism. Conspicuously lacking from the list of key processes in most text books is the role that many organisms play in the creation, modification and maintenance of habitats. These activities do not involve direct trophic interactions between species, but they are nevertheless important and common. The ecological literature is rich in examples of habitat modification by organisms, some of which have been extensively studied (e.g. Thayer 1979, Naiman et al. 1988).

Organisms as ecosystem engineers

Physical ecosystem engineers are organisms that directly or indirectly control the availability of resources to other organisms by causing physical state changes in biotic or abiotic materials. Physical ecosystem engineering by organisms is the physical modification, maintenance, or creation of habitats. Ecological effects of engineers on many other species occur in virtually all ecosystems because the physical state changes directly create nonfood resources such as living space, directly control abiotic resources, and indirectly modulate abiotic forces that, in turn, affect resource use by other organisms. Trophic interactions and resource competition do not constitute engineering. Engineering can have significant or trivial effects on other species, may involve the physical structure of an organism (like a tree) or structures made by an organism (like a beaver dam), and can, but does not invariably, have feedback effects on the engineer. We argue that engineering has both negative and positive effects on species richness and abundances at small scales, but the net effects are probably positive at larger scales encompassing engineered and nonengineered environments in ecological and evolutionary space and time. Models of the population dynamics of engineers suggest that the engineer/habitat equilibrium is often, but not always, locally stable and may show long-term cycles, with potential ramifications for community and ecosystem stability. As yet, data adequate to parameterize such a model do not exist for any engineer species. Because engineers control flows of energy and materials but do not have to participate in these flows, energy, mass, and stoichiometry do not appear to be useful in predicting which engineers will have big effects. Empirical observations suggest some potential generalizations about which species will be important engineers in which ecosystems. We point out some of the obvious, and not so obvious, ways in which engineering and trophic relations interact, and we call for greater research on physical ecosystem engineers, their impacts, and their interface with trophic relations.

Positive and negative effects of organisms as physical ecosystem engineers

The Devonian System of Euramerica contains at least 14 transgressive-regressive (T-R) cycles of eustatic origin. These are separated into three groups (or depophases) and from Carboniferous cycles by three prominent regressions. Twelve post-Lochkovian T-R cycles are recognized, and they commonly appear to result from abrupt deepening events followed by prolonged upward shallowing. Deepening events in the western United States (especially Nevada), western Canada, New York, Belgium, and Germany have been dated in the standard conodont zonation and are demonstrably simultaneous in several or all five regions. This synchroneity indicates control by eustatic sea-level fluctuations rather than by local or regional epeirogeny. Facies shifts in shelf sedimentary successions are more reliable indicators of the timing of sea-level fluctuations than are strandline shifts in the cratonic interior, because the latter are more influenced by local epeirogeny. Strandline shifts are most useful in estimating the relative magnitude for sea-level fluctuations.

Devonian facies progressions and the three prominent regressions are of a duration and an order of magnitude that could have been caused by episodes of growth and decay of Devonian oceanic ridge systems. The described T-R cycles could have formed in response to mid-plate thermal uplift and submarine volcanism. The latter process may have been a control on small-scale (1–5 m thick), upward-shallowing cycles within the major T-R cycles. Continental glaciation could have been a factor in sea-level fluctuations only in the Famennian and could not have been responsible for the Devonian facies progressions or the numerous T-R cycles.

The Frasnian extinctions were apparently cumulative rather than due to a single calamity. Two rapid sea-level rises occurred just before, and one at, the Frasnian-Famennian boundary. It is probable that this series of deepening events reduced the size of shallow-shelf habitats, caused repeated anoxic conditions in basinal areas, and drowned the reef ecosystems that had sustained the immensely diverse Devonian benthos.

Devonian eustatic fluctuations in Euramerica

Data on numbers of marine families within 91 metazoan classes known from the Phanerozoic fossil record are analyzed. The distribution of the 2800 fossil families among the classes is very uneven, with most belonging to a small minority of classes. Similarly, the stratigraphic distribution of the classes is very uneven, with most first appearing early in the Paleozoic and with many of the smaller classes becoming extinct before the end of that era. However, despite this unevenness, a Q-mode factor analysis indicates that the structure of these data is rather simple. Only three factors are needed to account for more than 90% of the data. These factors are interpreted as reflecting the three great “evolutionary faunas” of the Phanerozoic marine record: a trilobite-dominated Cambrian fauna, a brachiopod-dominated later Paleozoic fauna, and a mollusc-dominated Mesozoic-Cenozoic, or “modern,” fauna. Lesser factors relate to slow taxonomic turnover within the major faunas through time and to unique aspects of particular taxa and times.Each of the three major faunas seems to have its own characteristic diversity so that its expansion or contraction appears as being intimately associated with a particular phase in the history of total marine diversity. The Cambrian fauna expands rapidly during the Early Cambrian radiations and maintains dominance during the Middle to Late Cambrian “equilibrium.” The Paleozoic fauna then ascends to dominance during the Ordovician radiations, which increase diversity dramatically; this new fauna then maintains dominance throughout the long interval of apparent equilibrium that lasts until the end of the Paleozoic Era. The modern fauna, which slowly increases in importance during the Paleozoic Era, quickly rises to dominance with the Late Permian extinctions and maintains that status during the general rise in diversity to the apparent maximum in the Neogene. The increase in diversity associated with the expansion of each new fauna appears to coincide with an approximately exponential decline of the previously dominant fauna, suggesting possible displacement of each evolutionary fauna by its successor.

A factor analytic description of the phanerozoic marine fossil record

The composition of any environment or object is determined by a particular balance between material transport processes and chemical reactions within and around it. In the case of marine sedimentary deposits, the dominant agents of mass transport are often large bottom-dwelling animals that move particles and fluids during feeding, burrowing, tube construction, and irrigation. Such biogenic material transport has major direct and indirect effects on the composition of sediments and their overlying waters. In this chapter I review some of what is presently known about these effects, their implications for both chemical and biological properties of a deposit, and how they can be conceptualized in quantitative models.

The Effects of Macrobenthos on Chemical Properties of Marine Sediment and Overlying Water

During the Phanerozoic, the diversity of immobile suspension feeders living on the surface of soft substrata (ISOSS) declined significantly. Immobile taxa on hard surfaces and mobile taxa diversified. Extinction rates of ISOSS were significantly greater than in other benthos. These changes in the structure of benthic communities are attributed to increased biological disturbance of the sediment (bioturbation) by diversifying deposit feeders.

https://science.sciencemag.org/content/sci/203/4379/458.full.pdf

Biological bulldozers and the evolution of marine benthic communities.

and ratios in skeletal calcite of Mytilus trossulus: Covariation with metabolic rate, salinity, and carbon isotopic composition of seawater

Paleotemperatures have been widely deduced from skeletal 18 O/ 16 O ratios, but these are also dependent on salinity. Without an independent measure of salinity, 18 O/ 16 O ratios cannot provide accurate data on past temperature and climate. We grew marine mollusks (bivalves) in the field while real-time data on local seawater temperature and chemistry (Mg, Ca, δ 18 O, and salinity) were gathered. Here we show that for Mytilus trossulus, skeletal Mg/Ca ratios provide an accurate measure of temperature and that weekly sea-surface temperatures may be estimated with an apparent accuracy of approximately ± 1.5 °C. Thus, with analyses of both Mg/Ca and δ 18 O from the same specimen, it is possible to determine seawater temperature and δ 18 O.

Bivalve skeletons record sea-surface temperature and δ18O via Mg/Ca and 18O/16O ratios

Unlike other shell-enclosed marine invertebrates, articulate brachiopods are repellent to predators. Fish, sea stars, snails, and crabs all prefer bivalve molluscs such as mussels to articulates. The mussels tested are mobile and out-compete immobile articulates when space is limited. In subtidal field experiments, mussels alone and predators alone each reduced the survivorship of articulates. However, adding mussels to articulates in the presence of ambient predation increased brachiopod survivorship by diverting predation from the brachiopods to the mussels. Competition from mussels (or mussel-like bivalves) is a plausible cause of the post-Paleozoic decline of articulates.

https://science.sciencemag.org/content/sci/228/4707/1527.full.pdf

Brachiopods versus Mussels: Competition, Predation, and Palatability.

In the clastic Genesee Group of the Catskill delta, lateral changes of the fauna are believed to reflect onshore-offshore physicochemical gradients. A shoreward increase of infauna is interpreted as adaptation to increased environmental stress. Free immobile taxa were concentrated offshore, while vagile forms, presumably able to cope with shifting substrata, are dominant nearshore. A shorewards replacement of brachiopods by bivalves reflects the eurytopy and infaunal habits of the bivalves.



In Gencsee time, progradation was first rapid, then slow. The sequence is reversed in the superjacent Sonyea Group and the accompanying reversal of faunal patterns is strong evidence of faunal control by the rate of progradation. This indicates the hazardous nature of attempts to trace ‘community evolutioneditor’ using only a few studies from each period.

Charles W. Thayer

Papers

Biological bulldozers and the evolution of marine benthic communities.

and ratios in skeletal calcite of Mytilus trossulus: Covariation with metabolic rate, salinity, and carbon isotopic composition of seawater

Bivalve skeletons record sea-surface temperature and δ18O via Mg/Ca and 18O/16O ratios

Brachiopods versus Mussels: Competition, Predation, and Palatability.

Marine paleoecology in the Upper Devonian of New York