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On the use of path analysis and related procedures for the investigation of ecological problems
No abstract available.
Mast flowering and semelparity in bamboos: The bamboo fire cycle hypothesis
No abstract available.
On incorporating fire into our thinking about natural ecosystems: A response to Saha and Howe
Ecologists long have had a fascination with fire impacts, although they have been slow to incorporate this ecological factor into serious thinking about the structure of communities and evolution of species (Bond and van Wilgen 1996). The remarks by Saha and Howe (2001, in this issue) illustrate some of the problems ecologists have in trying to apply fire to their thinking about natural ecosystems. Fire is commonly perceived qualitatively in terms of presence or absence, and the variation in frequency and intensity, as well as other components of the fire regime (e.g., fig. 1), is ignored. Often not considered is the fact that plant life histories are fine-tuned to particular fire regims, and in this regard, landscapes present a range of selective peaks and valleys, both figuratively and literally. Because landscape patterns affect the propagation of natural fires, it is imperative that the degree of human disturbance be considered (e.g. Marsh [1864] 1965; Gadgil and Guhu 1993) and the limitations of basing evolutionary arguments on anthropogenically derived landscapes be recognized (e.g., Janzen and Martin 1982).
Individual covariation in life-history traits: Seeing the trees despite the forest
We investigated the influence of age on survival and breeding rates in a long-lived species Rissa tridactyla using models with individual random effects permitting variation and covariation in fitness components among individuals. Differences in survival or breeding probabilities among individuals are substantial, and there was positive covariation between survival and breeding probability; birds that were more likely to survive were also more likely to breed, given that they survived. The pattern of age-related variation in these rates detected at the individual level differed from that observed at the population level. Our results provided confirmation of what has been suggested by other investigators: within-cohort phenotypic selection can mask senescence. Although this phenomenon has been extensively studied in humans and captive animals, conclusive evidence of the discrepancy between population-level and individual-level patterns of age-related variation in life-history traits is extremely rare in wild animal populations. Evolutionary studies of the influence of age on life-history traits should use approaches differentiating population level from the genuine influence of age: only the latter is relevant to theories of life-history evolution. The development of models permitting access to individual variation in fitness is a promising advance for the study of senescence and evolutionary processes.
Population dynamics and mutualism: Functional responses of benefits and costs
We develop an approach for studying population dynamics resulting from mutualism by employing functional responses based on density‐dependent benefits and costs. These functional responses express how the population growth rate of a mutualist is modified by the density of its partner. We present several possible dependencies of gross benefits and costs, and hence net effects, to a mutualist as functions of the density of its partner. Net effects to mutualists are likely a monotonically saturating or unimodal function of the density of their partner. We show that fundamental differences in the growth, limitation, and dynamics of a population can occur when net effects to that population change linearly, unimodally, or in a saturating fashion. We use the mutualism between senita cactus and its pollinating seed‐eating moth as an example to show the influence of different benefit and cost functional responses on population dynamics and stability of mutualisms. We investigated two mechanisms that may alter this mutualism's functional responses: distribution of eggs among flowers and fruit abortion. Differences in how benefits and costs vary with density can alter the stability of this mutualism. In particular, fruit abortion may allow for a stable equilibrium where none could otherwise exist.
Evolution of trophic transmission in parasites: Why add intermediate hosts?
Although multihost complex life cycles (CLCs) are common in several distantly related groups of parasites, their evolution remains poorly understood. In this article, we argue that under particular circumstances, adding a second host to a single-host life cycle is likely to enhance transmission (i.e., reaching the target host). For instance, in several situations, the propagules of a parasite exploiting a predator species will achieve a higher host-finding success by encysting in a prey of the target predator than by other dispersal modes. In such a case, selection should favor the transition from a singleto a two-host life cycle that includes the prey species as an intermediate host. We use an optimality model to explore this idea, and we discuss it in relation to dispersal strategies known among free-living species, especially animal dispersal. The model found that selection favored a complex life cycle only if intermediate hosts were more abundant than definitive hosts. The selective value of a complex life cycle increased with predation rates by definitive hosts on intermediate hosts. In exploring trade-offs between transmission strategies, we found that more costly trade-offs made it more difficult to evolve a CLC while less costly trade-offs between traits could favor a mixed strategy.
Resource allocation in offspring provisioning: An evaluation of the conditions favoring the evolution of matrotrophy
We used analytic and simulation models to determine the ecological conditions favoring evolution of a matrotrophic fish from a lecithotrophic ancestor given a complex set of trade‐offs. Matrotrophy is the nourishment of viviparous embryos by resources provided between fertilization and parturition, while lecithotrophy describes embryo nourishment provided before fertilization. In fishes and reptiles, embryo nourishment encompasses a continuum from solely lecithotrophic to primarily matrotrophic. Matrotrophy has evolved independently from lecithotrophic ancestors many times in many groups. We assumed matrotrophy increased the number of offspring a viviparous female could gestate and evaluated conditions of food availability favoring lecithotrophy or matrotrophy. The matrotrophic strategy was superior when food resources exceeded demand during gestation but at a risk of overproduction and reproductive failure if food intake was limited. Matrotrophic females were leaner during gestation than lecithotrophic females, yielding shorter life spans. Our models suggest that matrotrophic embryo nourishment evolved in environments with high food availability, consistently exceeding energy requirements for maintaining relatively large broods. Embryo abortion with some resorption of invested energy is a necessary preadaptation to the evolution of matrotrophy. Future work should explore trade‐offs of age‐specific mortality and reproductive output for females maintaining different levels of fat storage during gestation.
Invertebrate eggs can fly: Evidence of waterfowl-mediated gene flow in aquatic invertebrates
Waterfowl often have been assumed to disperse freshwater aquatic organisms between isolated wetlands, but no one has analyzed the impact of this transport on the population structure of aquatic organisms. For three cladocerans (Daphnia ambigua, Daphnia laevis, and Sida crystallina) and one bryozoan (Cristatella mucedo), we estimated the genetic distances between populations across North America using sequences of several mitochondrial DNA genes and genotypic frequencies at allozyme and microsatellite loci. Waterfowl movements across North America (estimated from band recovery data) explained a significant proportion of the gene flow occurring between populations across the continent for three of the four species, even after controlling for geographic distances between localities. The fourth species, S. crystallina, has propagules less likely to survive desiccation or ingestion by birds. Differences in the capacity to exploit bird-mediated transport are likely to have important consequences for the ecology of aquatic communities and the spread of invasive species.
The role of local populations within a landscape context: Defining and classifying sources and sinks
The interaction of local populations has been the focus of an increasing number of studies in the past 30 years. The study of source-sink dynamics has especially generated much interest. Many of the criteria used to distinguish sources and sinks incorporate the process of apparent survival (i.e., the combined probability of true survival and site fidelity) but not emigration. These criteria implicitly treat emigration as mortality, thus biasing the classification of sources and sinks in a manner that could lead to flawed habitat management. Some of the same criteria require rather restrictive assumptions about population equilibrium that, when violated, can also generate misleading inference. Here, we expand on a criterion (denoted ?contribution? or Cr) that incorporates successful emigration in differentiating sources and sinks and that makes no restrictive assumptions about dispersal or equilibrium processes in populations of interest. The metric Cr is rooted in the theory of matrix population models, yet it also contains clearly specified parameters that have been estimated in previous empirical research. We suggest that estimates of emigration are important for delineating sources and sinks and, more generally, for evaluating how local populations interact to generate overall system dynamics. This suggestion has direct implications for issues such as species conservation and habitat management.
Consumers limit the abundance and dynamics of a perennial shrub with a seed bank
For nearly 30 years, ecologists have argued that predators of seeds and seedlings seldom have population-level effects on plants with persistent seed banks and density-dependent seedling survival. We parameterized stage-based population models that incorporated density dependence and seed dormancy with data from a 5.5-year experiment that quantified how granivorous mice and herbivorous voles influence bush lupine (Lupinus arboreus) demography. We asked how seed dormancy and density-dependent seedling survival mediate the impacts of these consumers in dune and grassland habitats. In dune habitat, mice reduced analytical ?? (the intrinsic rate of population growth) by 39%, the equilibrium number of above-ground plants by 90%, and the seed bank by 98%; voles had minimal effects. In adjacent grasslands, mice had minimal effects, but seedling herbivory by voles reduced analytical ?? by 15% and reduced both the equilibrium number of aboveground plants and dormant seeds by 63%. A bootstrap analysis demonstrated that these consumer effects were robust to parameter uncertainty. Our results demonstrate that the quantitative strengths of seed dormancy and density-dependent seedling survival-not their mere existence-critically mediate consumer effects. This study suggests that plant population dynamics and distribution may be more strongly influenced by consumers of seeds and seedlings than is currently recognized. ?? 2006 by The University of Chicago.
Habitat selection responses of parents to offspring predation risk: An experimental test
The ability of nest predation to influence habitat settlement decisions in birds is widely debated, despite its importance in limiting fitness. Here, we experimentally manipulated nest predation risk across a landscape and asked the question, do migratory birds assess and respond to variation in nest predation risk when choosing breeding habitats? We examined habitat preference by quantifying the density and settlement date of eight species of migratory passerines breeding in areas with and without intact nest predator communities. We found consistently more individuals nesting in areas with reduced nest predation than in areas with intact predator assemblages, although predation risk had no influence on settlement or breeding phenology. Additionally, those individuals occupying safer nesting habitats exhibited increased singing activity. These findings support a causal relationship between habitat choice and nest predation risk and suggest the importance of nest predation risk in shaping avian community structure and breeding activity. ?? 2006 by The University of Chicago. All rights reserved.
Habitat connectivity and ecosystem productivity: implications from a simple model.
The import of resources (food, nutrients) sustains biological production and food webs in resource-limited habitats. Resource export from donor habitats subsidizes production in recipient habitats, but the ecosystem-scale consequences of resource translocation are generally unknown. Here, I use a nutrient-phytoplankton-zooplankton model to show how dispersive connectivity between a shallow autotrophic habitat and a deep heterotrophic pelagic habitat can amplify overall system production in metazoan food webs. This result derives from the finite capacity of suspension feeders to capture and assimilate food particles: excess primary production in closed autotrophic habitats cannot be assimilated by consumers; however, if excess phytoplankton production is exported to food-limited heterotrophic habitats, it can be assimilated by zooplankton to support additional secondary production. Transport of regenerated nutrients from heterotrophic to autotrophic habitats sustains higher system primary production. These simulation results imply that the ecosystem-scale efficiency of nutrient transformation into metazoan biomass can be constrained by the rate of resource exchange across habitats and that it is optimized when the transport rate matches the growth rate of primary producers. Slower transport (i.e., reduced connectivity) leads to nutrient limitation of primary production in autotrophic habitats and food limitation of secondary production in heterotrophic habitats. Habitat fragmentation can therefore impose energetic constraints on the carrying capacity of aquatic ecosystems. The outcomes of ecosystem restoration through habitat creation will be determined by both functions provided by newly created aquatic habitats and the rates of hydraulic connectivity between them.
Selection for rapid embryo development correlates with embryo exposure to maternal androgens among passerine birds
Greater offspring predation favors evolution of faster development among species. We hypothesized that greater offspring predation exerts selection on mothers to increase levels of anabolic androgens in egg yolks to achieve faster development. Here, we tested whether (1) concentrations of yolk androgens in passerine species were associated with offspring predation and (2) embryo and nestling development rates were associated with yolk androgen concentrations. We examined three androgens that increase in potency along the synthesis pathway: androstenedione (A4) to testosterone (T) to 5??- dihydrotestosterone (5??-DHT). Concentrations of none of these steroids were related to clutch size; only A4 was allometrically related to egg volume. Species that experience greater predation showed higher yolk concentrations of T and 5??-DHT. Higher concentrations of T and particularly 5??-DHT were strongly correlated with faster development during the embryo period and less so during the nestling period. Development rates were most strongly correlated with 5??-DHT, suggesting that potency increases along the androgen synthesis pathway and that effects are mediated by the androgen receptor pathway. These results are consistent with the hypothesis that selection for faster development by time-dependent offspring mortality may be achieved epigenetically by varying embryo exposure to maternal anabolic steroids. ?? 2007 by The University of Chicago. All rights reserved.
Forage nutritive quality in the Serengeti ecosystem: The roles of fire and herbivory
Fire and herbivory are important determinants of nutrient availability in savanna ecosystems. Fire and herbivory effects on the nutritive quality of savanna vegetation can occur directly, independent of changes in the plant community, or indirectly, via effects on the plant community. Indirect effects can be further subdivided into those occurring because of changes in plant species composition or plant abundance (i.e., quality versus quantity). We studied relationships between fire, herbivory, rainfall, soil fertility, and leaf nitrogen (N), phosphorus (P), and sodium (Na) at 30 sites inside and outside of Serengeti National Park. Using structural equation modeling, we asked whether fire and herbivory influences were largely direct or indirect and how their signs and strengths differed within the context of natural savanna processes. Herbivory was associated with enhanced leaf N and P through changes in plant biomass and community composition. Fire was associated with reduced leaf nutrient concentrations through changes in plant community composition. Additionally, fire had direct positive effects on Na and nonlinear direct effects on P that partially mitigated the indirect negative effects. Key mechanisms by which fire reduced plant nutritive quality were through reductions of Na-rich grasses and increased abundance of Themeda triandra, which had below-average leaf nutrients. ?? 2007 by The University of Chicago. All rights reserved.
The golden rule of reviewing
A major bottleneck in the time required to publish a scientific or scholarly paper is the speed with which reviews by peers are returned to journals. Peer review is a reciprocal altruistic system in which each individual may perform every task—editors, reviewers, and authors—at different times. Journals have no way to coerce reviewers to return their critiques faster. To greatly shorten the time to publication, all actors in this altruistic network should abide by the Golden Rule of Reviewing: review for others as you would have others review for you. Say yes to reviewing whenever your duties and schedule allow; provide a thorough, fair, and constructive critique of the work; and do it at your first opportunity regardless of the deadline.
How vegetation and sediment transport feedbacks drive landscape change in the Everglades and wetlands worldwide
Mechanisms reported to promote landscape self‐organization cannot explain vegetation patterning oriented parallel to flow. Recent catastrophic shifts in Everglades landscape pattern and ecological function highlight the need to understand the feedbacks governing these ecosystems. We modeled feedback between vegetation, hydrology, and sediment transport on the basis of a decade of experimentation. Results from more than 100 simulations showed that flows just sufficient to redistribute sediment from sparsely vegetated sloughs to dense ridges were needed for an equilibrium patterned landscape oriented parallel to flow. Surprisingly, although vegetation heterogeneity typically conveys resilience, in wetlands governed by flow/sediment feedbacks it indicates metastability, whereby the landscape is prone to catastrophic shifts. Substantial increases or decreases in flow relative to the equilibrium condition caused an expansion of emergent vegetation and loss of open‐water areas that was unlikely to revert upon restoration of the equilibrium hydrology. Understanding these feedbacks is critical in forecasting wetland responses to changing conditions and designing management strategies that optimize ecosystem services, such as carbon sequestration or habitat provision. Our model and new sensitivity analysis techniques address these issues and make it newly apparent that simply returning flow to predrainage conditions in the Everglades may not be sufficient to restore historic landscape patterns and processes.
Optimization of biomass composition explains microbial growth-stoichiometry relationships
Integrating microbial physiology and biomass stoichiometry opens far-reaching possibilities for linking microbial dynamics to ecosystem processes. For example, the growth-rate hypothesis (GRH) predicts positive correlations among growth rate, RNA content, and biomass phosphorus (P) content. Such relationships have been used to infer patterns of microbial activity, resource availability, and nutrient recycling in ecosystems. However, for microorganisms it is unclear under which resource conditions the GRH applies. We developed a model to test whether the response of microbial biomass stoichiometry to variable resource stoichiometry can be explained by a trade-off among cellular components that maximizes growth. The results show mechanistically why the GRH is valid under P limitation but not under N limitation. We also show why variability of growth rate-biomass stoichiometry relationships is lower under P limitation than under N or C limitation. These theoretical results are supported by experimental data on macromolecular composition (RNA, DNA, and protein) and biomass stoichiometry from two different bacteria. In addition, compared to a model with strictly homeostatic biomass, the optimization mechanism we suggest results in increased microbial N and P mineralization during organic-matter decomposition. Therefore, this mechanism may also have important implications for our understanding of nutrient cycling in ecosystems.