Annual Review of Ecology, Evolution, and Systematics - Volume 31, 2000

The inclusive fitness effect attributable to an allele can be divided into an effect on matrilineal kin when the allele is maternally derived and an effect on patrilineal kin when paternally derived. However, the allele is not subject to selection on its effects on patrilineal kin when maternally derived nor on its effects on matrilineal kin when paternally derived. As a result, natural selection may favor alleles with effects that differ, depending on the allele's parental origin. At autosomal loci, this process is predicted to lead to the silencing of alleles when inherited from one or the other parent. At X-linked loci subject to random X inactivation, the process is predicted to lead to quantitative differences of expression between maternal and paternal alleles but not to complete silencing of one allele. The implications of this theory and some challenges to the theory are reviewed.

Herbivory first evolved in terrestrial mammals during the late Cretaceous, ∼100 million years ago (Mya). Of the ∼35 ordinal-level clades of extinct or extant eutherian mammals from the New World, ∼24 have been adapted to herbivory in one form or another. Dental adaptations for specialized terrestrial browsing are first recognized during the early Cenozoic (Paleocene-Eocene). Mammalian herbivores adapted for grazing did not become widespread in the New World until the middle Cenozoic; it seems that this adaptation and the spread of grasslands occurred during the late Oligocene (30 Mya) in South America ∼10 million years earlier than in North America (20 Mya). Carbon isotopic evidence from fossil herbivore teeth indicates that C3 plants predominated until the late Miocene (∼8 Mya). Thereafter, C3 and C4 terrestrial communities diversified. Late Pleistocene extinctions ∼10,000 years ago decimated the diversity of mammalian herbivores, particularly those of larger body size.

Conservation in New Zealand is failing to halt an ongoing decline in biodiversity. Classical problems of ecosystem loss and fragmentation have largely been countered in some regions by reservation of 30% of total land area. Unsustainable harvesting of native biodiversity has stopped; indeed harvesting of terrestrial species is rare. In contrast, marine reserves do not cover even 1% of the managed area, and harvest of native species, some of it unsustainable, are a major industry. Introduced pests, especially mammals, are the overwhelming conservation problem. Legislation, management, and considerable public opinion is based on preservationist ideals that demand the sanctity of native land biodiversity. Considerable success in threatened species management, island eradications, and mainland control of pests is increasing opportunities for restoration. New legislation is increasingly built on concepts of sustainability and offers the opportunity for integrating conservation, use, and development. Realization of these opportunities requires greater understanding of the relative merits of preservation versus sustainability, the dynamics and costs of pest control, the need for ecosystem processes in addition to individual species, and the involvement of people, especially the rights of indigenous Maori. Understanding marine environments and linking attitudes to land and sea is also a challenge.

Recent theories regarding the evolution of predator-prey interactions is reviewed. This includes theory about the dynamics and stability of both populations and traits, as well as theory predicting how predatory and anti-predator traits should respond to environmental changes. Evolution can stabilize or destabilize interactions; stability is most likely when only the predator evolves, or when traits in one or both species are under strong stabilizing selection. Stability seems least likely when there is coevolution and a bi-directional axis of prey vulnerability. When population cycles exist, adaptation may either increase or decrease the amplitude of those cycles. An increase in the defensive ability of prey is less likely to produce evolutionary counter-measures in its partner than is a comparable increase in attack ability of the predator. Increased productivity may increase or decrease offensive and defensive adaptations. The apparent predominance of evolutionary responses of prey to predators over those of predators to prey is in general accord with equilibrium theory, but theory on stability may be difficult to confirm or refute. Recent work on geographically structured populations promises to advance our understanding of the evolution of predator-prey interactions.

Understanding seed physiology is central to reconstructing how angiosperms have evolved, to characterizing dormancy and germination regimes shared by suites of species, and to devising sound strategies for seed bank conservation, agriculture, and forestry. While species with dormant seeds have received the lion's share of attention, hundreds of plant species exhibit no seed dormancy and germinate either viviparously on the parent plant or shortly after release. Embryos of these recalcitrant and viviparous species cannot tolerate the maturation drying that is usually prerequisite to dormancy; such desiccation intolerance creates challenges for storing and preserving such embryos. I review the physiology, morphology, and ecology of these desiccation-intolerant, nondormant lineages. Differences in the production and function of plant hormones are implicated in the occurrence of recalcitrance and vivipary in plant families. Plant hormones are key regulators of seed physiology and simultaneously coordinate responses of the seedling and mature plant to their environment. Desiccation-intolerant embryos occur most commonly among species of wet or flooded environments and have evolved multiple times in disparate lineages. Natural selection in wetland environments simply may not eliminate these seed types or may select for changes in hormone physiology that simultaneously affect both maternal and embryonic tissues. Integrative data from ecological, genetic, and physiological studies are needed to elucidate evolutionary origins and maintenance of reproductive strategies in organisms.

Inbreeding depression is of major concern in the management and conservation of endangered species. Inbreeding appears universally to reduce fitness, but its magnitude and specific effects are highly variable because they depend on the genetic constitution of the species or populations and on how these genotypes interact with the environment. Recent natural experiments are consistent with greater inbreeding depression in more stressful environments. In small populations of randomly mating individuals, such as are characteristic of many endangered species, all individuals may suffer from inbreeding depression because of the cumulative effects of genetic drift that decrease the fitness of all individuals in the population. In three recent cases, introductions into populations with low fitness appeared to restore fitness to levels similar to those before the effects of genetic drift. Inbreeding depression may potentially be reduced, or purged, by breeding related individuals. However, the Speke's gazelle example, often cited as a demonstration of reduction of inbreeding depression, appears to be the result of a temporal change in fitness in inbred individuals and not a reduction in inbreeding depression.

     Down, July 17, 1870

 My Dear Lubbock,

 …In England and many parts of Europe the marriages of cousins are objected to from their supposed injurious consequences: but this belief rests on no direct evidence. It is therefore manifestly desirable that the belief should be either proved false, or should be confirmed, so that in this latter case the marriages of cousins might be discouraged…

  It is moreover, much to be wished that the truth of the often repeated assertion that consanguineous marriages lead to deafness and dumbness, blindness, &c, should be ascertained: and all such assertions could be easily tested by the returns from a single census.

     Believe me,

     Yours very sincerely,

     Charles Darwin

Cichlid fishes (Perciformes: Teleostei) found in the lakes of Africa have served as model systems for the study of evolution. The enormous number of species (1000 in Lake Malawi alone), the great diversity of trophic adaptations and behaviors, and the extreme rapidity of their divergence (<50,000 y for some faunas) single out these organisms as examples of evolution in progress. Because these fishes are confined to discrete lacustrine environments and their origination is bounded by geological features, these groups provide models with which to study evolution. We review theoretical studies and empirical research on the cichlid faunas of Africa to provide a synthetic overview of current knowledge of the evolutionary processes at work in this group. This view provides the critical information needed to formulate and test hypotheses that may permit discrimination among the diverse theories and models that have been advanced to explain the evolution of these fishes.

The composition and structure of the semiarid or desert grasslands of southwestern North America have changed over the past 150 y. Brushy or woody species in these communities have increased in density and cover. This increase in density of woody species is called brush encroachment because most of these species have been present in these communities at lower densities for thousands of years. The brushy or woody species were not introduced from other continents or from great distances. They are indigenous species that have increased in density or cover because of changes in local abiotic or biotic conditions. The brushy and woody plants are not the cause of these changes, but their increase is the result of other factors. The causes of changes that have led to the present woody-brushy composition of these semiarid grasslands has been difficult to determine. Warming of the climate seems to be a background condition, but the driving force seems to be chronic, high levels of herbivory by domestic animals. This herbivory has reduced the aboveground grass biomass, leading to the reduction of fine fuel and a concomitant reduction or complete elimination of grassland fires. This combination of factors favors the encroachment, establishment, survival and growth of woody plants. Less competition from grasses, dispersal of seeds of woody plants by domestic animals, and changes in rodent, lagomorph, and insect populations seem to modify the rate of change. Elevated levels of atmospheric CO2 are not necessary to explain shrub encroachment in these semiarid grasslands. The direction of future change is difficult to predict. The density of brushy and woody plants will probably increase as will the stature and number of species. However, if soil nutrients increase, woody legumes may be replaced by other brushy or woody species. Reversing the changes that have been going on for 150 y will be a difficult, long-term, and perhaps impossible, task.

Macroevolution in the grasses has often involved change in the position (heterotopy) of developmental programs, possibly via ectopic gene expression. Heterotopy apparently has been involved in the evolution of unique epidermal morphology in the grasses and their sister genus, Joinvillea; in the origin of the grass flower and possibly in the spikelet as well; in the formation of unisexual flowers in the panicoid grasses, and in the repeated origin of C4 photosynthesis. Change in timing of development (heterochrony) may explain the novel morphology of the grass embryo. Changes in the structure and size of the nuclear genome correlate with phylogenetically informative cytogenetic characteristics. Most of the 10,000 species of grasses evolved tens of millions of years after the common ancestor of the family, indicating that the origin of novel morphologies did not lead to immediate radiation.

Tropical Asian rivers support a rich but incompletely known biota, including a host of fishes, a diverse array of benthic invertebrates, and an assemblage of mammals adapted to riverine wetlands. River ecology is dominated by flow seasonality imposed by monsoonal rains with profound consequences for fishes and zoobenthos. Information on life histories, feeding, and the trophic base of production of these animals is summarized. Widespread use of allochthonous foods by fishes and zoobenthos is apparent. Migration by fishes is often associated with breeding and results in seasonal occupation of different habitats. Riverine biodiversity is threatened by habitat degradation (pollution, deforestation of drainage basins), dams and flow regulation, as well as over-harvesting. Conservation efforts in tropical Asia are constrained by a variety of factors, including lack of ecological information, but the extent of public awareness and political commitment to environmental protection are likely determinants of the future of riverine biodiversity.

We summarize the influences of harvester ants of the genus Pogonomyrmex on communities and ecosystems. Because of nest densities, the longevity of nests, and the amount of seed harvested and soil handled, harvester ants have significant direct and indirect effects on community structure and ecosystem functioning. Harvester ants change plant species composition and diversity near their nests. These changes result from differential seed predation by the ants, their actions as seed dispersers and competitors with other granivores, and the favorable soil conditions they create through their digging. Their nest building creates islands of increased nutrient density. In some areas, the effects of their activities may be so pervasive that plant community structure is strongly influenced. Ant removal studies, which would reveal their total impact, have generally not been done. Granivore removals have been conducted in North America where ants are of lesser importance than small mammals, in contrast to other areas (except Israel) where ants are dominant granivores. We review the influence of harvester ants on their competitors, predators, and nest associates, and catalog the factors that influence their foraging patterns and consequently their local distribution.

The harvesting habit in ants since it was first scientifically confirmed by Moggridge has excited an exceptional degree of interest and surprise. But in truth, when one considers all the conditions, the wonder is that it is not more widely distributed.

     Henry Christopher McCook (138, p. 116)

Fossil plankton are difficult to identify but have formed a major component of most marine ecosystems throughout geological time. The earliest fossil heterotrophs include planktic forms, and subsequent adaptive innovations quickly appeared in the plankton; these include metazoans and animals with hard parts. Movement into the plankton occurred sporadically throughout geological history and seems to have been independent of any biological or environmental forcing mechanism. Subsequent radiations and extinctions in the cohort of plankton closely reflect events in the benthos. The diversity of zooplankton rose quickly during the early Paleozoic era, but low plankton diversity characterized the late Paleozoic. Significant radiations during early Mesozoic times led to an overall increase in diversity through the Phanerozoic eon. As the composition of the zooplankton has changed, so has their effect on biogeochemical cycles.

Evolutionary physiology represents an explicit fusion of two complementary approaches: evolution and physiology. Stimulated by four major intellectual and methodological developments (explicit consideration of diverse evolutionary mechanisms, phylogenetic approaches, incorporation of the perspectives and tools of evolutionary genetics and selection studies, and generalization of molecular techniques to exotic organisms), this field achieved prominence during the past decade. It addresses three major questions regarding physiological evolution: (a) What are the historical, ecological, and phylogenetic patterns of physiological evolution? (b) How important are and were each of the known evolutionary processes (natural selection, sexual selection, drift, constraint, genetic coupling/hitchhiking, and others) in engendering or limiting physiological evolution? and (c) How do the genotype, phenotype, physiological performance, and fitness interact in influencing one another's future values? To answer these questions, evolutionary physiology examines extant and historical variation and diversity, standing genetic and phenotypic variability in populations, and past and ongoing natural selection in the wild. Also, it manipulates genotypes, phenotypes, and environments of evolving populations in the laboratory and field. Thus, evolutionary physiology represents the infusion of paradigms, techniques, and approaches of evolutionary biology, genetics, and systematics into physiology. The reciprocal infusion of physiological approaches into evolutionary biology and systematics can likewise have great value and is a future goal.

…each level [of biological integration] offers unique problems and insights, and….each level finds its explanations of mechanism in the levels below, and its significance in the levels above. George A. Bartholomew (7, p. 8)

The focus of most ideas on diversity maintenance is species coexistence, which may be stable or unstable. Stable coexistence can be quantified by the long-term rates at which community members recover from low density. Quantification shows that coexistence mechanisms function in two major ways: They may be (a) equalizing because they tend to minimize average fitness differences between species, or (b) stabilizing because they tend to increase negative intraspecific interactions relative to negative interspecific interactions. Stabilizing mechanisms are essential for species coexistence and include traditional mechanisms such as resource partitioning and frequency-dependent predation, as well as mechanisms that depend on fluctuations in population densities and environmental factors in space and time. Equalizing mechanisms contribute to stable coexistence because they reduce large average fitness inequalities which might negate the effects of stabilizing mechanisms. Models of unstable coexitence, in which species diversity slowly decays over time, have focused almost exclusively on equalizing mechanisms. These models would be more robust if they also included stabilizing mechanisms, which arise in many and varied ways but need not be adequate for full stability of a system. Models of unstable coexistence invite a broader view of diversity maintenance incorporating species turnover.

In large-herbivore populations, environmental variation and density dependence co-occur and have similar effects on various fitness components. Our review aims to quantify the temporal variability of fitness components and examine how that variability affects changes in population growth rates. Regardless of the source of variation, adult female survival shows little year-to-year variation [coefficient of variation (CV <10%)], fecundity of prime-aged females and yearling survival rates show moderate year-to-year variation (CV <20%), and juvenile survival and fecundity of young females show strong variation (CV >30%). Old females show senescence in both survival and reproduction. These patterns of variation are independent of differences in body mass, taxonomic group, and ecological conditions. Differences in levels of maternal care may fine-tune the temporal variation of early survival. The immature stage, despite a low relative impact on population growth rate compared with the adult stage, may be the critical component of population dynamics of large herbivores. Observed differences in temporal variation may be more important than estimated relative sensitivity or elasticity in determining the relative demographic impact of various fitness components.

The impacts of airborne pollutants have been studied in only a few groups of soil animals, notably protozoans, nematodes, potworms, earthworms, mites, and collembolans. Pollutants in the form of acid depositions, which contain SO4²⁻, NOx, H⁺, heavy metals, and some organic compounds, are not homogeneously distributed on the landscape. Deposition patterns depend mainly on landscape configuration and plant cover. Airborne pollutants affect soil animals both directly and indirectly. Direct toxic effects are associated with uptake of free acidic water from the environment by some soil animals and with consumption of polluted food by others. Indirect effects are mediated primarily through disappearance or reduction of the food resources (microflora and microfauna) of soil animals, changes in organic matter content, and modification of microclimate. In the field, changes in competition among species are probably important factors that influence the soil animal community structure as well as the reactions of individual species to soil acidification or liming. The overall effect is a depauperation of soil with an attendant reduction in the rate of organic matter decomposition. We have provided five hypotheses, using soil fauna as indicators, to allow for quick evaluation of environmental changes caused by airborne pollutants.

In 1973, C. S. Holling introduced the word resilience into the ecological literature as a way of helping to understand the non-linear dynamics observed in ecosystems. Ecological resilience was defined as the amount of disturbance that an ecosystem could withstand without changing self-organized processes and structures (defined as alternative stable states). Other authors consider resilience as a return time to a stable state following a perturbation. A new term, adaptive capacity, is introduced to describe the processes that modify ecological resilience. Two definitions recognize the presence of multiple stable states (or stability domains), and hence resilience is the property that mediates transition among these states. Transitions among stable states have been described for many ecosystems, including semi-arid rangelands, lakes, coral reefs, and forests. In these systems, ecological resilience is maintained by keystone structuring processes across a number of scales, sources of renewal and reformation, and functional biodiversity. In practice, maintaining a capacity for renewal in a dynamic environment provides an ecological buffer that protects the system from the failure of management actions that are taken based upon incomplete understanding, and it allows managers to affordably learn and change.

Selective reporting—e.g., the preferential publication of results that are statistically significant, or consistent with theory or expectation—presents a challenge to meta-analysis and seriously undermines the quest for generalizations. Funnel graphs (scatterplots of effect size vs. sample size) help reveal the extent of selective reporting. They also allow the strength of biological effects to be judged easily, and they reaffirm the value of graphical presentations of data over statistical summaries.

Funnel graphs of published results, including: (a) sex-ratio variation in birds, (b) field estimates of heritabilities, and (c) relations between fluctuating asymmetry and individual attractiveness or fitness, suggest selective reporting is widespread and raise doubts about the true magnitude of these phenomena. Quasireplication—the “replication” of previous studies using different species or systems—has almost completely supplanted replicative research in ecology and evolution. Without incentives for formal replicative studies, which could come from changes to editorial policies, graduate training programs, and research funding priorities, the contract of error will continue to thwart attempts at robust generalizations.

“For as knowledges are now delivered, there is a kind of contract of error between the deliverer and the receiver: for he that delivereth knowledge desireth to deliver it in such a form as may be best believed, and not as may be best examined; and he that receiveth knowledge desireth rather present satisfaction than expectant inquiry; and so rather not to doubt than not to err: glory making the author not to lay open his weakness, and sloth making the disciple not to know his strength.”

      The Advancement of Learning, Francis Bacon, 1605 (8:170–171)

Biological invasions of marine habitats have been common, and many patterns emerge from the existing literature. In North America, we identify 298 nonindigenous species (NIS) of invertebrates and algae that are established in marine and estuarine waters, generating many “apparent patterns” of invasion: (a) The rate of reported invasions has increased exponentially over the past 200 years; (b) Most NIS are crustaceans and molluscs, while NIS in taxonomic groups dominated by small organisms are rare; (c) Most invasions have resulted from shipping; (d) More NIS are present along the Pacific coast than the Atlantic and Gulf coasts; (e) Native and source regions of NIS differ among coasts, corresponding to trade patterns. The validity of these apparent patterns remains to be tested, because strong bias exists in the data. Overall, the emergent patterns reflect interactive effects of propagule supply, invasion resistance, and sampling bias. Understanding the relative contribution of each component remains a major challenge for invasion ecology and requires standardized, quantitative measures in space and time that we now lack.