The commonly observed high diversity of trees in tropical rain forests and corals on tropical reefs is a nonequilibrium state which, if not disturbed further, will progress toward a low-diversity equilibrium community. This may not happen if gradual changes in climate favor different species. If equilibrium is reached, a lesser degree of diversity may be sustained by niche diversification or by a compensatory mortality that favors inferior competitors. However, tropical forests and reefs are subject to severe disturbances often enough that equilibrium may never be attained.

https://www.colby.edu/biology/BI358j/Readings/Diversity%20in%20rainforests%20and%20coral%20reefs.pdf

Diversity in tropical rain forests and coral reefs.

Humans are altering the composition of biological communities through a variety of activities that increase rates of species invasions and species extinctions, at all scales, from local to global. These changes in components of the Earth's biodiversity cause concern for ethical and aesthetic reasons, but they also have a strong potential to alter ecosystem properties and the goods and services they provide to humanity. Ecological experiments, observations, and theoretical developments show that ecosystem properties depend greatly on biodiversity in terms of the functional characteristics of organisms present in the ecosystem and the distribution and abundance of those organisms over space and time. Species effects act in concert with the effects of climate, resource availability, and disturbance regimes in influencing ecosystem properties. Human activities can modify all of the above factors; here we focus on modification of these biotic controls. The scientific community has come to a broad consensus on many aspects of the re- lationship between biodiversity and ecosystem functioning, including many points relevant to management of ecosystems. Further progress will require integration of knowledge about biotic and abiotic controls on ecosystem properties, how ecological communities are struc- tured, and the forces driving species extinctions and invasions. To strengthen links to policy and management, we also need to integrate our ecological knowledge with understanding of the social and economic constraints of potential management practices. Understanding this complexity, while taking strong steps to minimize current losses of species, is necessary for responsible management of Earth's ecosystems and the diverse biota they contain.

/pdf/effects-of-biodiversity-on-ecosystem-functioning-a-consensus-13j8vabcus.pdf

Effects of biodiversity on ecosystem functioning: a consensus of current knowledge

All ecosystems are exposed to gradual changes in climate, nutrient loading, habitat fragmentation or biotic exploitation. Nature is usually assumed to respond to gradual change in a smooth way. However, studies on lakes, coral reefs, oceans, forests and arid lands have shown that smooth change can be interrupted by sudden drastic switches to a contrasting state. Although diverse events can trigger such shifts, recent studies show that a loss of resilience usually paves the way for a switch to an alternative state. This suggests that strategies for sustainable management of such ecosystems should focus on maintaining resilience.

/pdf/catastrophic-shifts-in-ecosystems-2yv2mwzdwb.pdf

Catastrophic shifts in ecosystems.

This book is the second of two volumes in a series on terrestrial and marine comparisons, focusing on the temporal complement of the earlier spatial analysis of patchiness and pattern (Levin et al. 1993). The issue of the relationships among pattern, scale, and patchiness has been framed forcefully in John Steele’s writings of two decades (e.g., Steele 1978). There is no pattern without an observational frame. In the words of Nietzsche, “There are no facts… only interpretations.”

https://esajournals.onlinelibrary.wiley.com/doi/pdf/10.2307/1941447

The Problem of Pattern and Scale in Ecology: The Robert H. MacArthur Award Lecture

Species richness is a fundamental measurement of community and regional diversity, and it underlies many ecological models and conservation strategies. In spite of its importance, ecologists have not always appreciated the effects of abundance and sampling effort on richness measures and comparisons. We survey a series of common pitfalls in quantifying and comparing taxon richness. These pitfalls can be largely avoided by using accumulation and rarefaction curves, which may be based on either individuals or samples. These taxon sampling curves contain the basic information for valid richness comparisons, including category‐subcategory ratios (species-to-genus and species-toindividual ratios). Rarefaction methods ‐ both sample-based and individual-based ‐ allow for meaningful standardization and comparison of datasets. Standardizing data sets by area or sampling effort may produce very different results compared to standardizing by number of individuals collected, and it is not always clear which measure of diversity is more appropriate. Asymptotic richness estimators provide lower-bound estimates for taxon-rich groups such as tropical arthropods, in which observed richness rarely reaches an asymptote, despite intensive sampling. Recent examples of diversity studies of tropical trees, stream invertebrates, and herbaceous plants emphasize the importance of carefully quantifying species richness using taxon sampling curves.

/pdf/quantifying-biodiversity-procedures-and-pitfalls-in-the-2ut2f3nw4g.pdf

Quantifying biodiversity: procedures and pitfalls in the measurement and comparison of species richness

The sequence of species observed after a relatively large space is opened up is a consequence of the following mechanisms. "Opportunist" species with broad dispersal powers and rapid growth to maturity usually arrive first and occupy empty space. These species cannot invade and grow in the presence of adults of their own or other species. Several alternative mechanisms may then determine which species replace these early occupants. Three models of such mechanisms have been proposed. The first "facilitation" model suggests that the entry and growth of the later species is dependent upon the earlier species "preparing the ground"; only after this can later species colonize. Evidence in support of this model applies mainly to certain primary successions and in heterotrophic succession. A second "tolerance" model suggests that a predictable sequence is produced by the existence of species that have evolved different strategies for exploiting resources. Later species will be those able to tolerate lower levels...

Mechanisms of succession in natural communities and their role in community stability and organization

Champion, H. G. 1936. A preliminary survey of the forest types of India and Burma. Indian Forest Records (Silviculture Series), Vol. 1, No. 1. Manson-Bahr, P. H. 1954. Manson's Tropical Diseases. 14 Ed., London, Cassell and Co. Nolte, A. 1955. Field observations on the daily routine and social behavior of common Indian monkeys, with special reference to the Bonnet monkey (Macaca radiata Geoff roy). Jour. Bombay Nat. Hist. Soc. 53: 177-184.

The Influence of Interspecific Competition and Other Factors on the Distribution of the Barnacle Chthamalus Stellatus

In a strictly defined sample of competition studies using controlled field experiments, covering 215 species and 527 experiments, competition was found in most of the studies, in somewhat more than half of the species, and in about two-fifths of the experiments. In most of these experiments interspecific competition was not distinguished from intraspecific competition. In the few studies in which the two were separated, interspecific competition was the stronger form in about onesixth of all experiments done. When competition was demonstrated, intraspecific competition was as strong or stronger than interspecific in three-quarters of the experiments. Some evidence from this literature survey suggests that negative results may be underrepresented, so that the absolute values of these figures may be too high. Since this bias should apply also to studies of all taxa, habitats, or other interactions it should not greatly affect estimates of the relative prevalence of competition. Since these estimates come fr...

On the prevalence and relative importance of interspecific competition: evidence from field experiments

"The balance of nature has been a background assumption in natural history since antiquity" (Egerton 1973, p. 322). This continues to be true today; some modern field ecologists, assuming that natural ecosystems are stable, have applied ideas of mathematical stability theory to the actual communities they are studying. We believe that, before one applies such theory to a natural population or community, one should first decide whether or not it is stable. Our aim here is to describe the sorts of evidence one would need to obtain from natural populations or communities in order to decide whether they are stable or persistent, as defined below. One aspect we shall stress in particular is whether any given real community exists in multiple stable states in different places at the same time or in the same place at different times (Sutherland 1974).

Joseph H. Connell

Papers

Diversity in tropical rain forests and coral reefs.

Mechanisms of succession in natural communities and their role in community stability and organization

The Influence of Interspecific Competition and Other Factors on the Distribution of the Barnacle Chthamalus Stellatus

On the prevalence and relative importance of interspecific competition: evidence from field experiments

On the Evidence Needed to Judge Ecological Stability or Persistence