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At least 415 records · Page 23Linked to original sources

Trace metal bioavailability: Modeling chemical and biological interactions of sediment-bound zinc

Extractable concentrations of sediment-bound Zn, as modified by the physicochemical form of the metal in the sediments, controlled Zn concentrations in the deposit-feeding bivalves Scrobicularia plana (collected from 40 stations in 17 estuaries in southwest England) and Macoma balthica (from 28 stations in San Francisco Bay). Over a wide range of concentrations, a significant correlation was found between ammonium acetate-soluble concentrations of Zn in sediments and Zn concentrations in Scrobicularia . This correlation was insufficiently precise to be of predictive value for Scrobicularia , and did not hold for Macoma over the narrower range of Zn concentrations observed in San Francisco Bay. Strong correlation of Zn concentrations in Scrobicularia and the bioavailability of sediment-bound Zn to Macoma with ratios of sorption substrate (oxides of iron and manganese, organic carbon, carbonates, humic materials) concentrations in sediments were found in both the English and San Francisco Bay study areas. These correlations were attributed to substrate competition for sorption of Zn within sediments, assuming: 1) competition for sorption of Zn was largely controlled by the relative concentrations of substrates present in the sediments and 2) the bioavailability of Zn to the deposit feeders was determined by the partitioning of Zn among the substrates. The correlations indicated that the availability of Zn to the bivalves increased when concentrations of either amorphous inorganic oxides or humic substances increased in sediments. Availability was reduced at increased concentrations of organic carbon and, in San Francisco Bay, ammonium acetate-soluble Mn. Concentrations of biologically available Zn in solution and low salinities may also have enhanced Zn uptake, although the roles of these variables were less obvious from the statistical analysis.

Book chapter

Monthly Strontium/Calcium oscillations in symbiotic coral aragonite: Biological effects limiting the precision of the paleotemperature proxy

In thermodynamic equilibrium with sea water the Sr/Ca ratio of aragonite varies predictably with temperature and the Sr/Ca ratio in coral have thus become a frequently used proxy for past Sea Surface Temperature (SST). However, biological effects can offset the Sr/Ca ratio from its equilibrium value. We report high spatial resolution ion microprobe analyses of well defined skeletal elements in the reef‐building coral Porites lutea that reveal distinct monthly oscillations in the Sr/Ca ratio, with an amplitude in excess of ten percent. The extreme Sr/Ca variations, which we propose result from metabolic changes synchronous with the lunar cycle, introduce variability in Sr/Ca measurements based on conventional sampling techniques well beyond the analytical precision. These variations can limit the accuracy of Sr/Ca paleothermometry by conventional sampling techniques to about 2°C. Our results may help explain the notorious difficulties involved in obtaining an accurate and consistent calibration of the Sr/Ca vs. SST relationship.

Geophysical Research Letters

A spatially explicit, empirical estimate of tree-based biological nitrogen fixation in forests of the United States

Quantifying human impacts on the nitrogen (N) cycle and investigating natural ecosystem N cycling depend on the magnitude of inputs from natural biological nitrogen fixation (BNF). Here, we present two bottom‐up approaches to quantify tree‐based symbiotic BNF based on forest inventory data across the coterminous United States and SE Alaska. For all major N‐fixing tree genera, we quantify BNF inputs using (1) ecosystem N accretion rates (kg N ha −1 yr −1 ) scaled with spatial data on tree abundance and (2) percent of N derived from fixation (%N dfa ) scaled with tree N demand (from tree growth rates and stoichiometry). We estimate that trees fix 0.30–0.88 Tg N yr −1 across the study area (1.4–3.4 kg N ha −1 yr −1 ). Tree‐based N fixation displays distinct spatial variation that is dominated by two genera, Robinia (64% of tree‐associated BNF) and Alnus (24%). The third most important genus, Prosopis , accounted for 5%. Compared to published estimates of other N fluxes, tree‐associated BNF accounted for 0.59 Tg N yr −1 , similar to asymbiotic (0.37 Tg N yr −1 ) and understory symbiotic BNF (0.48 Tg N yr −1 ), while N deposition contributed 1.68 Tg N yr −1 and rock weathering 0.37 Tg N yr −1 . Overall, our results reveal previously uncharacterized spatial patterns in tree BNF that can inform large‐scale N assessments and serve as a model for improving tree‐based BNF estimates worldwide. This updated, lower BNF estimate indicates a greater ratio of anthropogenic to natural N inputs, suggesting an even greater human impact on the N cycle.

Global Biogeochemical Cycles

Prediction of unprecedented biological shifts in the global ocean

Impermanence is an ecological principle 1 but there are times when changes occur nonlinearly as abrupt community shifts (ACSs) that transform the ecosystem state and the goods and services it provides 2 . Here, we present a model based on niche theory 3 to explain and predict ACSs at the global scale. We test our model using 14 multi-decadal time series of marine metazoans from zooplankton to fish, spanning all latitudes and the shelf to the open ocean. Predicted and observed fluctuations correspond, with both identifying ACSs at the end of the 1980s 4,5,6,7 and 1990s 5,8 . We show that these ACSs coincide with changes in climate that alter local thermal regimes, which in turn interact with the thermal niche of species to trigger long-term and sometimes abrupt shifts at the community level. A large-scale ACS is predicted after 2014—unprecedented in magnitude and extent—coinciding with a strong El Niño event and major shifts in Northern Hemisphere climate. Our results underline the sensitivity of the Arctic Ocean, where unprecedented melting may reorganize biological communities 5,9 , and suggest an increase in the size and consequences of ACS events in a warming world.

Nature Climate Change

Global cycling and climate effects of aeolian dust controlled by biological soil crusts

Biological soil crusts (biocrusts) cover ~12% of the global land surface. They are formed by an intimate association between soil particles, photoautotrophic and heterotrophic organisms, and they effectively stabilize the soil surface of drylands. Quantitative information on the impact of biocrusts on the global cycling and climate effects of aeolian dust, however, is not available. Here, we combine the currently limited experimental data with a global climate model to investigate the effects of biocrusts on regional and global dust cycling under current and future conditions. We estimate that biocrusts reduce the global atmospheric dust emissions by ~60%, preventing the release of ~0.7 Pg dust per year. Until 2070, biocrust coverage is expected to be severely reduced by climate change and land-use intensification. The biocrust loss will cause an increased dust burden, leading to a reduction of the global radiation budget of around 0.12 to 0.22 W m−2, corresponding to about 50% of the total direct forcing of anthropogenic aerosols. This biocrust control on dust cycling and its climate impacts have important implications for human health, biogeochemical cycling and the functioning of the ecosystems, and thus should be considered in the modelling, mitigation and management of global change.

Nature Geoscience

A global dataset of terrestrial biological nitrogen fixation

Biological nitrogen fixation (BNF) is the main natural source of new nitrogen inputs in terrestrial ecosystems, supporting terrestrial productivity, carbon uptake, and other Earth system processes. We assembled a comprehensive global dataset of field measurements of BNF in all major N-fixing niches across natural terrestrial biomes derived from the analysis of 376 BNF studies. The dataset comprises 32 variables, including site location, biome type, N-fixing niche, sampling year, quantification method, BNF rate (kg N ha −1 y −1 ), the percentage of nitrogen derived from the atmosphere (%N dfa ), N fixer or N-fixing substrate abundance, BNF rate per unit of N fixer abundance, and species identity. Overall, the dataset combines 1,207 BNF rates for trees, shrubs, herbs, soil, leaf litter, woody litter, dead wood, mosses, lichens, and biocrusts, 152 herb %N dfa values, 1,005 measurements of N fixer or N-fixing substrate abundance, and 762 BNF rates per unit of N fixer abundance for a total of 424 species across 66 countries. This dataset facilitates synthesis, meta-analysis, upscaling, and model benchmarking of BNF fluxes at multiple spatial scales.

Scientific Data

Precise determination of δ 88 Sr in rocks, minerals, and waters by double-spike TIMS: A powerful tool in the study of chemical, geologic, hydrologic and biologic processes

We present strontium isotopic ( 88 Sr/ 86 Sr and 87 Sr/ 86 Sr) results obtained by 87 Sr– 84 Sr double spike thermal ionization mass-spectrometry (DS-TIMS) for several standards as well as natural water samples and mineral samples of abiogenic and biogenic origin. The detailed data reduction algorithm and a user-friendly Sr-specific stand-alone computer program used for the spike calibration and the data reduction are also presented. Accuracy and precision of our δ 88 Sr measurements, calculated as permil (‰) deviations from the NIST SRM-987 standard, were evaluated by analyzing the NASS-6 seawater standard, which yielded δ 88 Sr = 0.378 ± 0.009‰. The first DS-TIMS data for the NIST SRM-607 potassium feldspar standard and for several US Geological Survey carbonate, phosphate, and silicate standards (EN-1, MAPS-4, MAPS-5, G-3, BCR-2, and BHVO-2) are also reported. Data obtained during this work for Sr-bearing solids and natural waters show a range of δ 88 Sr values of about 2.4‰, the widest observed so far in terrestrial materials. This range is easily resolvable analytically because the demonstrated external error (±SD, standard deviation) for measured δ 88 Sr values is typically ≤0.02‰. It is shown that the “true” 87 Sr/ 86 Sr value obtained by the DS-TIMS or any other external normalization method combines radiogenic and mass-dependent mass-fractionation effects, which cannot be separated. Therefore, the “true” 87 Sr/ 86 Sr and the δ 87 Sr parameter derived from it are not useful isotope tracers. Data presented in this paper for a wide range of naturally occurring sample types demonstrate the potential of the δ 88 Sr isotope tracer in combination with the traditional radiogenic 87 Sr/ 86 Sr tracer for studying a variety of biological, hydrological, and geological processes.

Journal of Analytical Atomic Spectrometry

Spatially robust estimates of biological nitrogen (N) fixation imply substantial human alteration of the tropical N cycle

Biological nitrogen fixation (BNF) is the largest natural source of exogenous nitrogen (N) to unmanaged ecosystems and also the primary baseline against which anthropogenic changes to the N cycle are measured. Rates of BNF in tropical rainforest are thought to be among the highest on Earth, but they are notoriously difficult to quantify and are based on little empirical data. We adapted a sampling strategy from community ecology to generate spatial estimates of symbiotic and free-living BNF in secondary and primary forest sites that span a typical range of tropical forest legume abundance. Although total BNF was higher in secondary than primary forest, overall rates were roughly five times lower than previous estimates for the tropical forest biome. We found strong correlations between symbiotic BNF and legume abundance, but we also show that spatially free-living BNF often exceeds symbiotic inputs. Our results suggest that BNF in tropical forest has been overestimated, and our data are consistent with a recent top-down estimate of global BNF that implied but did not measure low tropical BNF rates. Finally, comparing tropical BNF within the historical area of tropical rainforest with current anthropogenic N inputs indicates that humans have already at least doubled reactive N inputs to the tropical forest biome, a far greater change than previously thought. Because N inputs are increasing faster in the tropics than anywhere on Earth, both the proportion and the effects of human N enrichment are likely to grow in the future.

Proceedings of the National Academy of Sciences of

Climate change and physical disturbance cause similar community shifts in biological soil crusts

Biological soil crusts (biocrusts)—communities of mosses, lichens, cyanobacteria, and heterotrophs living at the soil surface—are fundamental components of drylands worldwide, and destruction of biocrusts dramatically alters biogeochemical processes, hydrology, surface energy balance, and vegetation cover. While there has been long-standing concern over impacts of 5 physical disturbances on biocrusts (e.g., trampling by livestock, damage from vehicles), there is also increasing concern over the potential for climate change to alter biocrust community structure. Using long-term data from the Colorado Plateau, USA, we examined the effects of 10 years of experimental warming and altered precipitation (in full-factorial design) on biocrust communities, and compared the effects of altered climate with those of long-term physical 10 disturbance (>10 years of replicated human trampling). Surprisingly, altered climate and physical disturbance treatments had similar effects on biocrust community structure. Warming, altered precipitation frequency [an increase of small (1.2 mm) summer rainfall events], and physical disturbance from trampling all promoted early successional community states marked by dramatic declines in moss cover and increased cyanobacteria cover, with more variable effects 15 on lichens. While the pace of community change varied significantly among treatments, our results suggest that multiple aspects of climate change will affect biocrusts to the same degree as physical disturbance. This is particularly disconcerting in the context of warming, as temperatures for drylands are projected to increase beyond those imposed by the climate treatments used in our study.

PNAS

Decline in biological soil crust N-fixing lichens linked to increasing summertime temperatures

Across many global drylands, biocrusts form a protective barrier on the soil surface and fill many critical roles in these harsh yet fragile environments. Previous short-term research suggests that climate change and invasive plant introduction can damage and alter biocrust communities, yet few long-term observations exist. Using a globally unique long-term record of continuous biocrust surveys from a rare never-grazed, protected grassland on the US Colorado Plateau, we found lichen species diversity and cover to be negatively correlated with increasing summer air temperatures, while moss species showed more sensitivity to variation in precipitation and invasive grass cover. These results suggest that dryland systems may be at a critical tipping point where ongoing warming could result in biological soil crust degradation.

Proceedings of the National Academies of Science (

Overestimated natural biological nitrogen fixation translates to an exaggerated CO2 fertilization effect in Earth System Models

CO 2 fertilization of the terrestrial biosphere is limited by nitrogen. Biological nitrogen fixation (BNF) is the dominant natural nitrogen source to the terrestrial biosphere and can alleviate nitrogen limitation but is poorly constrained in Earth system models (ESMs). Here, we compare terrestrial BNF from an ensemble of ESMs of the 6th Coupled Model Intercomparison Project to a new global synthesis of observations across natural and agricultural biomes. We find that compared to observations, ESMs underestimate agricultural BNF but overestimate natural BNF in the present day by over 50%. Natural BNF is overestimated in the most productive ecosystems that contribute most to the terrestrial carbon sink (forests and grasslands). ESMs with different BNF representations yield a range of BNF responses to CO 2 enrichment. Some ESMs with phenomenological representations of BNF predict a natural BNF increase in response to a doubling of CO 2 that aligns with a meta-analysis of CO 2 enrichment experiments (31% increase) but fail to account for the substantial carbon cost of BNF. In contrast, ESMs with mechanistic representations of BNF account for its carbon cost as well as its regulation by nitrogen limitation but overestimate the BNF response to a doubling of CO 2 (135% increase). Overall, all current BNF representations in ESMs fall short of fully capturing its response to rising atmospheric CO 2 . Finally, we find a positive correlation between modeled present-day natural BNF and the CO 2 fertilization effect across ESMs, suggesting that overestimated natural BNF translates to an exaggerated CO 2 fertilization effect of approximately 11% in ESMs.

PNAS

Mapping biological soil crusts in a Hawaiian dryland

Historical and ongoing land use patterns in the Hawaiian Islands have degraded the Islands’ drylands, causing erosion and detrimentally affecting adjacent coastal marine ecosystems. Biological soil crust (biocrust) communities have been shown to increase soil stability in drylands worldwide, but their efficacy in mitigating soil erosion in Hawaiian drylands is largely unknown. Using a combination of field data and imagery collected by small unmanned aerial systems (sUAS), we mapped biocrusts and examined their influence on soil stability in the Kawaihae watershed, an erosion-prone dryland on leeward Hawai`i Island. We created classified maps of biocrust cover from imagery collected at three spatial resolutions (1.2, 2.1 and 2.8 cm/pixel) using the pixel-based Support Vector Machine (SVM) classifier and investigated the impacts of spatial resolution and biocrust level of development on classification accuracy. Our medium (2.1 cm) resolution image produced the highest overall classification accuracy when biocrust was treated as a single class (82.1%). We explored the spatial impacts of biocrusts on soil loss via sUAS-derived measurements of elevation change over a four-year time span. We found differences in soil loss among land cover types, but robustly quantifying these was a challenge, as much of the change fell below statistically significant limits of detection. We investigated the relationship between biocrust development and soil stability by conducting soil aggregate stability testing at the three biocrust levels of development (LODs) present at the study site. We found a significant increase in soil stability from soils without surface biocrusts (LOD score of 0) to those with biocrusts at any development level (LOD 1–3). Our research adds to the body of biocrust knowledge by presenting new information about biocrust distribution and soil stabilization capabilities in Hawaiian drylands. We also provide insights into the trade-offs between spatial resolution and classification accuracy for biocrust classification and land cover analysis.

Hawaii

Development of a stock-recruitment model and assessment of biological reference points for the Lake Erie walleye fishery

We developed an updated stock–recruitment relationship for Lake Erie Walleye Sander vitreus using the Akaike information criterion model selection approach. Our best stock–recruitment relationship was a Ricker spawner–recruit function to which spring warming rate was added as an environmental variable, and this regression model explained 39% of the variability in Walleye recruitment over the 1978 through 2006 year-classes. Thus, most of the variability in Lake Erie Walleye recruitment appeared to be attributable to factors other than spawning stock size and spring warming rate. The abundance of age-0 Gizzard Shad Dorosoma cepedianum, which was an important term in previous models, may still be an important factor for Walleye recruitment, but poorer ability to monitor Gizzard Shad since the late 1990s could have led to that term failing to appear in our best model. Secondly, we used numerical simulation to demonstrate how to use the stock recruitment relationship to characterize the population dynamics (such as stable age structure, carrying capacity, and maximum sustainable yield) and some biological reference points (such as fishing rates at different important biomass or harvest levels) for an age-structured population in a deterministic way.

North American Journal of Fisheries Management

Earning their stripes: The potential of tiger trout and other salmonids as biological controls of forage fishes in a western reservoir

Maintaining a balance between predator and prey populations can be an ongoing challenge for fisheries managers, especially in managing artificial ecosystems such as reservoirs. In a high-elevation Utah reservoir, the unintentional introduction of the Utah Chub Gila atraria and its subsequent population expansion prompted managers to experimentally shift from exclusively stocking Rainbow Trout Oncorhynchus mykiss to also stocking tiger trout (female Brown Trout Salmo trutta × male Brook Trout Salvelinus fontinalis ) and Bonneville Cutthroat Trout O. clarkii utah (hereafter, Cutthroat Trout) as potential biological control agents. We measured a combination of diet, growth, temperature, and abundance and used bioenergetic simulations to quantify predator demand versus prey supply. Utah Chub were the predominant prey type for tiger trout, contributing up to 80% of the diet depending on the season. Utah Chub represented up to 70% of the total diet consumed by Cutthroat Trout. Although Utah Chub dominated the fish biomass in the reservoir, we still estimated abundances of 238,000 tiger trout, 214,000 Cutthroat Trout, and 55,000 Rainbow Trout. Consequently, when expanded to the population level of each predator, tiger trout and Cutthroat Trout consumed large quantities of Utah Chub on an annual basis: tiger trout consumed 508,000 kg (2,660 g/predator) of the standing prey population, and Cutthroat Trout consumed an estimated 322,000 kg (1,820 g/predator). The estimated combined consumption by Cutthroat Trout and tiger trout exceeded the estimate of Utah Chub annual production. As such, our results suggest that the high rates of piscivory exhibited by Cutthroat Trout and tiger trout in artificial lentic ecosystems are likely sufficient to effectively reduce the overall abundance of forage fishes and to prevent forage fishes from dominating fish assemblages. Collectively, this research provides the first documented findings on tiger trout ecology and performance, which will aid managers in designing and implementing the best stocking strategy to optimize sport fish performance, control undesirable forage fish, and enhance and maintain angler satisfaction.

Utah

Using interviews and biological sign surveys to infer seasonal use of forested and agricultural portions of a human-dominated landscape by Asian elephants in Nepal

Understanding how wide-ranging animals use landscapes in which human use is highly heterogeneous is important for determining patterns of human–wildlife conflict and designing mitigation strategies. Here, we show how biological sign surveys in forested components of a human-dominated landscape can be combined with human interviews in agricultural portions of a landscape to provide a full picture of seasonal use of different landscape components by wide-ranging animals and resulting human–wildlife conflict. We selected Asian elephants ( Elephas maximus ) in Nepal to illustrate this approach. Asian elephants are threatened throughout their geographic range, and there are large gaps in our understanding of their landscape-scale habitat use. We identified all potential elephant habitat in Nepal and divided the potential habitat into sampling units based on a 10 km by 10 km grid. Forested areas within grids were surveyed for signs of elephant use, and local villagers were interviewed regarding elephant use of agricultural areas and instances of conflict. Data were analyzed using single-season and multi-season (dynamic) occupancy models. A single-season occupancy model applied to data from 139 partially or wholly forested grid cells estimated that 0.57 of grid cells were used by elephants. Dynamic occupancy models fit to data from interviews across 158 grid cells estimated that monthly use of non-forested, human-dominated areas over the preceding year varied between 0.43 and 0.82 with a minimum in February and maximum in October. Seasonal patterns of crop raiding by elephants coincided with monthly elephant use of human-dominated areas, and serious instances of human–wildlife conflict were common. Efforts to mitigate human–elephant conflict in Nepal are likely to be most effective if they are concentrated during August through December when elephant use of human-dominated landscapes and human–elephant conflict are most common.

Ethology Ecology and Evolution

Chemical and biological status of lakes and streams in the upper midwest: assessment of acidic deposition effects

Many lakes in three areas in the Upper Midwest - northeastern Minnesota, northern Wisconsin, and the Upper Peninsula of Michigan - have low acid neutralizing capacity (ANC) and may be susceptible to change by acidic deposition. Northcentral Wisconsin and the Upper Peninsula of Michigan together contain about 150-300 acidic lakes (ANC ≤ 0), whereas none have been found in Minnesota. These acidic lakes are precipitation-dominated, Clearwater seepage lakes having small surface area, shallow depth, and low concentrations of dissolved organic carbon. The spatial distribution of these acidic lakes parallels a west to east gradient of increasing sulfate and hydrogen ion deposition. Several of these acidic lakes exhibit chemical characteristics and biological changes consistent with those observed elsewhere in waters reported to be acidified by acidic deposition. However, an hypothesis of recent lake acidification is not supported by analyses of either historical chemical data or diatom remains in lake sediments, and natural sources of acidity and alternative ecological processes have not been conclusively eliminated as causative factors. Streams in this three-state region have high ANC and appear to be insensitive to acidic deposition. The species richness and composition of lacustrine fish communities in the region are partly related to pH and associated chemical factors. Sport fishes considered acid-sensitive and of primary concern with regard to acidification include walleye, smallmouth bass, and black crappie. The fishery in at least one lake, Morgan Lake in Wisconsin (pH 4.6), may have declined because of acidification. Given the general lack of quantitative fishery data for acidic Wisconsin and Michigan lakes, however, more general conclusions concerning impacts or the absence of impacts of acidification on the region's fishery resources are not possible.

Minnesota;Wisconsin;Michigan

A synthesis of the biology and ecology of sculpin species in the Laurentian Great Lakes and implications for the adaptive capacity of the benthic ecosystem

The Laurentian Great Lakes have experienced recent ecosystem changes that could lead to reductions in adaptive capacity and ultimately a loss of biodiversity and production throughout the food web. Observed changes in Great Lakes benthic communities include declines of native species and widespread success of invasive species like dreissenid mussels in all but Lake Superior. Understanding the ecology of native benthic deepwater preyfish and the reasons for their declines is important for predicting future losses in adaptive capacity and diversity, as well as managing the Great Lakes ecosystem to avoid such losses. Native sculpin species ( Cottus bairdii , C. cognatus , C. ricei , Myoxocephalus thompsonii ) historically were among the most abundant of the Great Lakes native deepwater benthic preyfish community and are an important link between offshore benthic and pelagic food webs. With one exception, these species have declined in abundance throughout the Great Lakes in recent years, but relatively little is known about their biology and ecology. This review synthesizes the available knowledge for the Great Lakes sculpin species and provides suggestions for future research efforts, which include understanding reproductive ecology and spawning behavior, connectivity and dispersal of populations, early life history, and influences of interactions with native and non-native species.

Great Lakes

Optimal timing in biological processes

A general approach for obtaining solutions to a class of biological optimization problems is provided. The general problem is one of determining the appropriate time to take some action, when the action can be taken only once during some finite time frame. The approach can also be extended to cover a number of other problems involving animal choice (e.g., mate selection, habitat selection). Returns (assumed to index fitness) are treated as random variables with time-specific distributions, and can be either observable or unobservable at the time action is taken. In the case of unobservable returns, the organism is assumed to base decisions on some ancillary variable that is associated with returns. Optimal policies are derived for both situations and their properties are discussed. Various extensions are also considered, including objective functions based on functions of returns other than the mean, nonmonotonic relationships between the observable variable and returns; possible death of the organism before action is taken; and discounting of future returns. A general feature of the optimal solutions for many of these problems is that an organism should be very selective (i.e., should act only when returns or expected returns are relatively high) at the beginning of the time frame and should become less and less selective as time progresses. An example of the application of optimal timing to a problem involving the timing of bird migration is discussed, and a number of other examples for which the approach is applicable are described.

American Naturalist