Annual Review of Ecology, Evolution, and Systematics - Volume 35, 2004

Vertebrate dispersal of fruits and seeds is a common feature of many modern angiosperms and gymnosperms, yet the evolution and frequency of this feature in the fossil record remain unclear. Increasingly complex information suggests that (a) plants had the necessary morphological features for vertebrate dispersal by the Pennsylvanian, but possibly in the absence of clear vertebrate dispersal agents; (b) vertebrate herbivores first diversified in the Permian, and consistent dispersal relationships became possible; (c) the Mesozoic was dominated by large herbivorous dinosaurs, possible sources of diffuse, whole-plant dispersal; (d) simultaneously, several groups of small vertebrates, including lizards and, in the later Mesozoic, birds and mammals, could have established more specific vertebrate-plant associations, but supporting evidence is rudimentary; and (e) the diversification of small mammals and birds in the Tertiary established a consistent basis for organ-level interactions, allowing for the widespread occurrence of biotic dispersal in gymnosperms and angiosperms.

Many factors (climate warming, pollution, harvesting, introduced species) can contribute to disease outbreaks in marine life. Concomitant increases in each of these makes it difficult to attribute recent changes in disease occurrence or severity to any one factor. For example, the increase in disease of Caribbean coral is postulated to be a result of climate change and introduction of terrestrial pathogens. Indirect evidence exists that (a) warming increased disease in turtles; (b) protection, pollution, and terrestrial pathogens increased mammal disease; (c) aquaculture increased disease in mollusks; and (d) release from overfished predators increased sea urchin disease. In contrast, fishing and pollution may have reduced disease in fishes. In other taxa (e.g., sea grasses, crustaceans, sharks), there is little evidence that disease has changed over time. The diversity of patterns suggests there are many ways that environmental change can interact with disease in the ocean.

Bird song provides an unusually impressive illustration of vertebrate behavioral diversification. Research on bird song evolution traditionally focuses on factors that enhance song diversity, such as cultural transmission and sexual selection. Recent advances in the study of proximate mechanisms of vocal behavior, however, provide opportunities for studying mechanistic constraints on song evolution. The main goal of this review is to examine, from both conceptual and empirical perspectives, how proximate mechanisms might temper patterns of song evolution. We provide an overview of the two “substrates” of song evolution, memes and vocal mechanisms. We argue that properties of vocal mechanisms (control, production, and ontogeny) constrain vocal potential and may thus limit pathways of meme evolution. We then consider how vocal mechanisms may constrain song evolution under five scenarios of drift and selection and examine four specific song traits for which mechanistic constraints appear to counter the diversifying effects of sexual selection. These examples illustrate the interplay between meme evolution as a diversifying influence and proximate limitations as a barrier to song divergence. We conclude by suggesting that vocal mechanisms not only constrain song evolution but also can facilitate the evolution of novel vocal features.

Ecological indicators have widespread appeal to scientists, environmental managers, and the general public. Indicators have long been used to detect changes in nature, but the scientific maturation in indicator development primarily has occurred in the past 40 years. Currently, indicators are mainly used to assess the condition of the environment, as early-warning signals of ecological problems, and as barometers for trends in ecological resources. Use of ecological indicators requires clearly stated objectives; the recognition of spatial and tempor al scales; assessments of statistical variability, precision, and accuracy; linkages with specific stressors; and coupling with economic and social indicators. Legislatively mandated use of ecological indicators occurs in many countries worldwide and is included in international accords. As scientific advancements and innovation in the development and use of ecological indicators continue through applications of molecular biology, computer technology such as geographic information systems, data management such as bioinformatics, and remote sensing, our ability to apply ecological indicators to detect signals of environmental change will be substantially enhanced.

Deer have expanded their range and increased dramatically in abundance worldwide in recent decades. They inflict major economic losses in forestry, agriculture, and transportation and contribute to the transmission of several animal and human diseases. Their impact on natural ecosystems is also dramatic but less quantified. By foraging selectively, deer affect the growth and survival of many herb, shrub, and tree species, modifying patterns of relative abundance and vegetation dynamics. Cascading effects on other species extend to insects, birds, and other mammals. In forests, sustained overbrowsing reduces plant cover and diversity, alters nutrient and carbon cycling, and redirects succession to shift future overstory composition. Many of these simplified alternative states appear to be stable and difficult to reverse. Given the influence of deer on other organisms and natural processes, ecologists should actively participate in efforts to understand, monitor, and reduce the impact of deer on ecosystems.

Ecological risks associated with the release of transgenic crops include nontarget effects of the crop and the escape of transgenes into wild populations. Nontarget effects can be of two sorts: (a) unintended negative effects on species that do not reduce yield and (b) greater persistence of the crop in feral populations. Conventional agricultural methods, such as herbicide and pesticide application, have large and well-documented nontarget effects. To the extent that transgenes have more specific target effects, transgenic crops may have fewer nontarget effects. The escape of transgenes into wild populations, via hybridization and introgression, could lead to increased weediness or to the invasion of new habitats by the wild population. In addition, native species with which the wild plant interacts (including herbivores, pathogens, and other plant species in the community) could be negatively affected by “transgenic-wild” plants. Conventional crop alleles have facilitated the evolution of increased weediness in several wild populations. Thus, some transgenes that allow plants to tolerate biotic and abiotic stress (e.g., insect resistance, drought tolerance) could have similar effects.

Mutualisms occur when interactions between species produce reciprocal benefits. However, the outcome of these interactions frequently shifts from positive, to neutral, to negative, depending on the environmental and community context, and indirect effects commonly produce unexpected mutualisms that have community-wide consequences. The dynamic, and context dependent, nature of mutualisms can transform consumers, competitors, and parasites into mutualists, even while they consume, compete with, or parasitize their partner species. These dynamic, and often diffuse, mutualisms strongly affect community organization and ecosystem processes, but the historic focus on pairwise interactions decoupled from their more complex community context has obscured their importance. In aquatic systems, mutualisms commonly support ecosystem-defining foundation species, underlie energy and nutrient dynamics within and between ecosystems, and provide mechanisms by which species can rapidly adjust to ecological variance. Mutualism is as important as competition, predation, and physical disturbance in determining community structure, and its impact needs to be adequately incorporated into community theory.

Species are routinely used as fundamental units of analysis in biogeography, ecology, macroevolution, and conservation biology. A large literature focuses on defining species conceptually, but until recently little attention has been given to the issue of empirically delimiting species. Researchers confronted with the task of delimiting species in nature are often unsure which method(s) is (are) most appropriate for their system and data type collected. Here, we review twelve of these methods organized into two general categories of tree- and nontree-based approaches. We also summarize the relevant biological properties of species amenable to empirical evaluation, the classes of data required, and some of the strengths and limitations of each method. We conclude that all methods will sometimes fail to delimit species boundaries properly or will give conflicting results, and that virtually all methods require researchers to make qualitative judgments. These facts, coupled with the fuzzy nature of species boundaries, require an eclectic approach to delimiting species and caution against the reliance on any single data set or method when delimiting species.

No one definition has as yet satisfied all naturalists; yet every naturalist knows vaguely what he means when he speaks of a species.

     Darwin (1859/1964)

Molecular tools have profoundly rearranged our understanding of metazoan phylogeny. Initially based on the nuclear small ribosomal subunit (SSU or 18S) gene, recent hypotheses have been corroborated by several sources of data (including the nuclear large ribosomal subunit, Hox genes, mitochondrial gene order, concatenated mitochondrial genes, and the myosin II heavy chain gene). Herein, the evidence supporting our current understanding is discussed on a clade by clade basis. Bilaterian animals consist of three clades: Deuterostomia, Lophotrochozoa, and Ecdysozoa. Each clade is supported by molecular and morphological data. Deuterostomia is smaller than traditionally recognized, consisting of hemichordates, echinoderms, chordates, and Xenoturbella (an enigmatic worm-like animal). Lophotrochozoa groups animals with a lophophore feeding apparatus (Brachiopoda, Bryozoa, and Phoronida) and trochophore larvae (e.g., annelids and mollusk), as well as several other recognized phyla (e.g., platyhelmin thes, sipunculans, nemerteans). Ecdysozoa comprises molting animals (e.g., arthropods, nematodes, tardigrades, priapulids), grouping together two major model organisms (Drosophila and Caenorhabditis) in the same lineage. Platyhelminthes do not appear to be monophyletic, with Acoelomorpha holding a basal position in Bilateria. Before the emergence of bilateral animals, sponges, ctenophorans, cnidarians, and placozoans split from the main animal lineage, but order of divergence is less than certain. Many questions persist concerning relationships within Ecdysozoa and Lophotrochozoa, poriferan monophyly, and the placement of many less-studied taxa (e.g., kinorhynchs, gastrotrichs, gnathostomulids, and entoprocts).

Local habitat and biological diversity of streams and rivers are strongly influenced by landform and land use within the surrounding valley at multiple scales. However, empirical associations between land use and stream response only varyingly succeed in implicating pathways of influence. This is the case for a number of reasons, including (a) covariation of anthropogenic and natural gradients in the landscape; (b) the existence of multiple, scale-dependent mechanisms; (c) nonlinear responses; and (d) the difficulties of separating present-day from historical influences. Further research is needed that examines responses to land use under different management strategies and that employs response variables that have greater diagnostic value than many of the aggregated measures in current use.

In every respect, the valley rules the stream.

     H.B.N. Hynes (1975)

Studies of plant and animal assemblages from both the terrestrial and the marine fossil records reveal persistence for extensive periods of geological time, sometimes millions of years. Persistence does not require lack of change or the absence of variation from one occurrence of the assemblage to the next in geological time. It does, however, imply that assemblage composition is bounded and that variation occurs within those bounds. The principal cause for these patterns appears to be species-, and perhaps clade-level, environmental fidelity that results in long-term tracking of physical conditions. Other factors that influence persistent recurrence of assemblages are historical, biogeographic effects, the “law of large numbers,” niche differentiation, and biotic interactions. Much research needs to be done in this area, and greater uniformity is needed in the approaches to studying the problem. However, great potential also exists for enhanced interaction between paleoecology and neoecology in understanding spatiotemporal complexity of ecological dynamics.

Tropical forests are being lost at an alarming rate. Studies from various tropical locations report losses of forest birds as possibly direct or indirect results of deforestation. Although it may take a century for all the sensitive species to be extirpated from a site following habitat loss, species with larger or heavier bodies and those foraging on insects, fruits, or both are particularly extinction prone. Larger- or heavier-bodied species may occur at low densities, increasing their vulnerability to habitat alterations. Insectivores are vulnerable for reasons such as the loss of preferred microhabitats, poor dispersal abilities, and/or ground nesting habits that make them susceptible to predation. The lack of year-round availability of fruits may make survival in deforested or fragmented areas difficult for frugivores. Extirpation of large predators, superior competitors, pollinators, and seed dispersers may have repercussions for tropical ecosystem functioning. Large tropical reserves that adequately protect existing forest avifauna are needed. Sound ecological knowledge of tropical forest avifauna for biodiversity-friendly forest management practices is also needed but sorely lacking.

This review focuses on five key evolutionary issues pertaining to animal cognition, defined as the neuronal processes concerned with the acquisition, retention, and use of information. Whereas the use of information, or decision making, has been relatively well examined by students of behavior, evolutionary aspects of other cognitive traits that affect behavior, including perception, learning, memory, and attention, are less well understood. First, there is ample evidence for genetically based individual variation in cognitive traits, although much of the information for some traits comes from humans. Second, several studies documented positive association between cognitive abilities and performance measures linked to fitness. Third, information on the evolution of cognitive traits is available primarily for color vision and decision making. Fourth, much of the data on plasticity of cognitive traits appears to reflect nonadaptive phenotypic plasticity, perhaps because few evolutionary analyses of cognitive plasticity have been carried out. Nonetheless, several studies suggest that cognitive traits show adaptive plasticity, and at least one study documented genetically based individual variation in plasticity. Fifth, whereas assertions that cognition has played a central role in animal evolution are not supported by currently available data, theoretical considerations indicate that cognition may either increase or decrease the rate of evolutionary change.

Floral evolution has often been associated with differences in pollination syndromes. Recently, this conceptual structure has been criticized on the grounds that flowers attract a broader spectrum of visitors than one might expect based on their syndromes and that flowers often diverge without excluding one type of pollinator in favor of another. Despite these criticisms, we show that pollination syndromes provide great utility in understanding the mechanisms of floral diversification. Our conclusions are based on the importance of organizing pollinators into functional groups according to presumed similarities in the selection pressures they exert. Furthermore, functional groups vary widely in their effectiveness as pollinators for particular plant species. Thus, although a plant may be visited by several functional groups, the relative selective pressures they exert will likely be very different. We discuss various methods of documenting selection on floral traits. Our review of the literature indicates overwhelming evidence that functional groups exert different selection pressures on floral traits. We also discuss the gaps in our knowledge of the mechanisms that underlie the evolution of pollination syndromes. In particular, we need more information about the relative importance of specific traits in pollination shifts, about what selective factors favor shifts between functional groups, about whether selection acts on traits independently or in combination, and about the role of history in pollination-syndrome evolution.

Salamanders are cryptic and, though largely unrecognized as such, extremely abundant vertebrates in a variety of primarily forest and grassland environments, where they regulate food webs and contribute to ecosystem resilience-resistance (= stability) in several ways: (a) As mid-level vertebrate predators, they provide direct and indirect biotic control of species diversity and ecosystem processes along grazer and detritus pathways; (b) via their migrations, they connect energy and matter between aquatic and terrestrial landscapes; (c) through association with underground burrow systems, they contribute to soil dynamics; and (d) they supply high-quality and slowly available stores of energy and nutrients for tertiary consumers throughout ecological succession. Salamanders also can provide an important service to humans through their use as cost-effective and readily quantifiable metrics of ecosystem health and integrity. The diverse ecological roles of salamanders in natural areas underscore the importance of their conservation.

Ecologists and evolutionary biologists are broadly interested in how the interactions among organisms influence their abundance, distribution, phenotypes, and genotypic composition. Recently, we have seen a growing appreciation of how multispecies interactions can act synergistically or antagonistically to alter the ecological and evolutionary outcomes of interactions in ways that differ fundamentally from outcomes predicted by pairwise interactions. Here, we review the evidence for criteria identified to detect community-based, diffuse coevolution. These criteria include (a) the presence of genetic correlations between traits involved in multiple interactions, (b) interactions with one species that alter the likelihood or intensity of interactions with other species, and (c) nonadditive combined effects of multiple interactors. In addition, we review the evidence that multispecies interactions have demographic consequences for populations, as well as evolutionary consequences. Finally, we explore the experimental and analytical techniques, and their limitations, used in the study of multispecies interactions. Throughout, we discuss areas in particular need of future research.

Spatial synchrony refers to coincident changes in the abundance or other time-varying characteristics of geographically disjunct populations. This phenomenon has been documented in the dynamics of species representing a variety of taxa and ecological roles. Synchrony may arise from three primary mechanisms:(a) dispersal among populations, reducing the size of relatively large populations and increasing relatively small ones; (b) congruent dependence of population dynamics on a synchronous exogenous random factor such as temperature or rainfall, a phenomenon known as the “Moran effect”; and (c) trophic interactions with populations of other species that are themselves spatially synchronous or mobile. Identification of the causes of synchrony is often difficult. In addition to intraspecific synchrony, there are many examples of synchrony among populations of different species, the causes of which are similarly complex and difficult to identify. Furthermore, some populations may exhibit complex spatial dynamics such as spiral waves and chaos. Statistical tests based on phase coherence and/or time-lagged spatial correlation are required to characterize these more complex patterns of spatial dynamics fully.

Edge effects have been studied for decades because they are a key component to understanding how landscape structure influences habitat quality. However, making sense of the diverse patterns and extensive variability reported in the literature has been difficult because there has been no unifying conceptual framework to guide research. In this review, we identify four fundamental mechanisms that cause edge responses: ecological flows, access to spatially separated resources, resource mapping, and species interactions. We present a conceptual framework that identifies the pathways through which these four mechanisms can influence distributions, ultimately leading to new ecological communities near habitat edges. Next, we examine a predictive model of edge responses and show how it can explain much of the variation reported in the literature. Using this model, we show that, when observed, edge responses are largely predictable and consistent. When edge responses are variable for the same species at the same edge type, observed responses are rarely in opposite directions. We then show how remaining variability may be understood within our conceptual frameworks. Finally, we suggest that, despite all the research in this area, the development of tools to extrapolate edge responses to landscapes has been slow, restricting our ability to use this information for conservation and management.

The evolutionary succession of marine photoautotrophs began with the origin of photosynthesis in the Archean Eon, perhaps as early as 3.8 billion years ago. Since that time, Earth's atmosphere, continents, and oceans have undergone substantial cyclic and secular physical, chemical, and biological changes that selected for different phytoplankton taxa. Early in the history of eukaryotic algae, between 1.6 and 1.2 billion years ago, an evolutionary schism gave rise to “green” (chlorophyll b–containing) and “red” (chlorophyll c–containing) plastid groups. Members of the “green” plastid line were important constituents of Neoproterozoic and Paleozoic oceans, and, ultimately, one green clade colonized land. By the mid-Mesozoic, the green line had become ecologically less important in the oceans. In its place, three groups of chlorophyll c–containing eukaryotes, the dinoflagellates, coccolithophorids, and diatoms, began evolutionary trajectories that have culminated in ecological dominance in the contemporary oceans. Breakup of the supercontinent Pangea, continental shelf flooding, and changes in ocean redox chemistry may all have contributed to this evolutionary transition. At the same time, the evolution of these modern eukaryotic taxa has influenced both the structure of marine food webs and global biogeochemical cycles.

We review the evidence of regime shifts in terrestrial and aquatic environments in relation to resilience of complex adaptive ecosystems and the functional roles of biological diversity in this context. The evidence reveals that the likelihood of regime shifts may increase when humans reduce resilience by such actions as removing response diversity, removing whole functional groups of species, or removing whole trophic levels; impacting on ecosystems via emissions of waste and pollutants and climate change; and altering the magnitude, frequency, and duration of disturbance regimes. The combined and often synergistic effects of those pressures can make ecosystems more vulnerable to changes that previously could be absorbed. As a consequence, ecosystems may suddenly shift from desired to less desired states in their capacity to generate ecosystem services. Active adaptive management and governance of resilience will be required to sustain desired ecosystem states and transform degraded ecosystems into fundamentally new and more desirable configurations.