The species richness (diversity) of local plant and animal assemblages-biological communities-balances regional processes of species formation and geographic dispersal, which add species to communities, against processes of predation, competitive exclusion, adaptation, and stochastic variation, which may promote local extinction. During the past three decades, ecologists have sought to explain differences in local diversity by the influence of the physical environment on local interactions among species, interactions that are generally believed to limit the number of coexisting species. But diversity of the biological community often fails to converge under similar physical conditions, and local diversity bears a demonstrable dependence upon regional diversity. These observations suggest that regional and historical processes, as well as unique events and circumstances, profoundly influence local community structure. Ecologists must broaden their concepts of community processes and incorporate data from systematics, biogeography, and paleontology into analyses of ecological patterns and tests of community theory.
A latitudinal gradient in biodiversity has existed since before the time of the dinosaurs, yet how and why this gradient arose remains unresolved. Here we review two major hypotheses for the origin of the latitudinal diversity gradient. The time and area hypothesis holds that tropical climates are older and historically larger, allowing more opportunity for diversification. This hypothesis is supported by observations that temperate taxa are often younger than, and nested within, tropical taxa, and that diversity is positively correlated with the age and area of geographical regions. The diversification rate hypothesis holds that tropical regions diversify faster due to higher rates of speciation (caused by increased opportunities for the evolution of reproductive isolation, or faster molecular evolution, or the increased importance of biotic interactions), or due to lower extinction rates. There is phylogenetic evidence for higher rates of diversification in tropical clades, and palaeontological data demonstrate higher rates of origination for tropical taxa, but mixed evidence for latitudinal differences in extinction rates. Studies of latitudinal variation in incipient speciation also suggest faster speciation in the tropics. Distinguishing the roles of history, speciation and extinction in the origin of the latitudinal gradient represents a major challenge to future research.
George Gaylord Simpson famously postulated that much of life's diversity originated as adaptive radiations—more or less simultaneous divergences of numerous lines from a single ancestral adaptive type. However, identifying adaptive radiations has proven difficult due to a lack of broad‐scale comparative datasets. Here, we use phylogenetic comparative data on body size and shape in a diversity of animal clades to test a key model of adaptive radiation, in which initially rapid morphological evolution is followed by relative stasis. We compared the fit of this model to both single selective peak and random walk models. We found little support for the early‐burst model of adaptive radiation, whereas both other models, particularly that of selective peaks, were commonly supported. In addition, we found that the net rate of morphological evolution varied inversely with clade age. The youngest clades appear to evolve most rapidly because long‐term change typically does not attain the amount of divergence predicted from rates measured over short time scales. Across our entire analysis, the dominant pattern was one of constraints shaping evolution continually through time rather than rapid evolution followed by stasis. We suggest that the classical model of adaptive radiation, where morphological evolution is initially rapid and slows through time, may be rare in comparative data.
The present study proposes to reconcile the different spatial and temporal scales of regional species production and local constraint on species richness. Although interactions between populations rapidly achieve equilibrium and limit membership in ecological communities locally, these interactions occur over heterogeneous environments within large regions, where the populations of species are stably regulated through competition and habitat selection. Consequently, exclusion of species from a region depends on long‐term regional‐scale environmental change or evolutionary change among interacting populations, bringing species production and extinction onto the same scale and establishing a link between local and regional processes.
Ricklefs, Robert E. An Analysis of Nesting Mortality in Birds. Smithsonian Contributions to Zoology, 9:1-48. 1969.-This study was initiated to evaluate nesting mortality of birds as a feature of the environment and as a selective force in the evolution of reproductive strategies. Representative nesting-success data from the literature for most groups of birds were transformed into daily mortality rates to eliminate differences among species in the length of the nest cycle. These data are presented by taxonomic groupings and for passerines by geographical region and nest construction and placement. The strength and pattern of various mortality factors are described in detail. Predation, starvation, desertion, hatching failure, and adverse weather are the most prevalent factors, but nestsite competition, brood parasitism, and arthropod infestation may be important in some species. It is demonstrated that the various mortality factors can be identified by characteristic patterns of nesting losses involving differences in mortality rates between the egg and nestling periods and the within-nest component of mortality rates. Among Temperate Zone passerines, field-nesting and marsh-nesting species have the highest mortality rates while those species nesting in trees, especially in cavities, enjoy higher success. Starvation is prevalent in marsh and field species but desertion is more restricted to tree-nesting species. In general, arctic species have lower mortality rates and tropical species higher rates, although there is a similar gradient from arid to humid regions within the tropics. The relative abundance of a species is related directly to its mortality rate in arctic regions, but is not in temperate and tropical regions. Birds of prey generally have low mortality rates although starvation is often a major factor. Nesting losses in seabirds are caused primarily by crowded conditions in colonies and loss of eggs due to inadequate nest construction. Chick deaths come about primarily through their wandering away from parental care which is most common in the semiprecocial Charadriiformes. Precocial shorebirds and water birds enjoy higher egg success than ground-nesting passerines but game birds exhibit similar mortality rates. Little is known of the survival of precocial chicks after hatching except that mortality rates may be initially quite high and decrease with age. The fate of altricial birds after fledging is also poorly documented. It is postulated that interspecific differences in mortality rates are determined by evolutionarily acceptable levels of adult risk to lower mortality rates of offspring through parental care, adult adaptations of morphology and behavior for foraging which result in limitations on nesting adaptations, environmental unpredictability which reduces the effectiveness of adaptations, and-most import-the diversity of predators to which a species must adapt. Official publication date is handstamped in a limited number of initial copies and is recorded in the Institution's annual report, S...
Charles Darwin's travels on HMS Beagle taught him that islands are an important source of evidence for evolution. Because many islands are young and have relatively few species, evolutionary adaptation and species proliferation are obvious and easy to study. In addition, the geographical isolation of many islands has allowed evolution to take its own course, free of influence from other areas, resulting in unusual faunas and floras, often unlike those found anywhere else. For these reasons, island research provides valuable insights into speciation and adaptive radiation, and into the relative importance of contingency and determinism in evolutionary diversification.
An important issue in the study of biodiversity is the extent to which global patterns of species richness reflect large-scale processes and historical contingencies. Ecological interactions in local assemblages may constrain the number of species that can coexist, but differences in diversity in similar habitats within different regions (diversity anomalies) suggest that this limit is not firm. Variation in rate of species production could influence regional and perhaps local diversity independently of the ecological capacity of an area to support coexisting species, thereby creating diversity anomalies. Temperate Zone genera of plants that are disjunct between similar environments in eastern Asia and eastern North America (EAS-ENA) have twice as many species in Asia as in North America. Because lineages of these genera in Asia and North America are mostly sister pairs, they share a common history of adaptation and ecological relationship before disjunction. Thus, the diversity anomaly in EAS-ENA genera is not an artefact of taxon or habitat sampling but reflects differences in the net diversification (speciation-extinction) of the lineages in each of the continents. Here we propose that the most probable cause of the EAS-ENA anomaly in diversity is the extreme physiographical heterogeneity of temperate eastern Asia, especially compared with eastern North America, which in conjunction with climate and sea-level change has provided abundant opportunities for evolutionary radiation through allopatric speciation.
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