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Determinants of reproductive costs in the long-lived Black-legged Kittiwake: A multiyear experiment

We studied reproductive costs of Black-legged Kittiwakes ( Rissa tridactyla ) in Prince William Sound, Alaska (USA) by removing entire clutches from randomly selected nests over four successive years, and then contrasting survival and fecundity of adults from manipulated and unmanipulated nests in each subsequent year. To elucidate mechanisms that lead to the expression of reproductive costs, we simultaneously characterized several behavioral and physiological parameters among adults in the two treatment groups. We also examined naturally nonbreeding adults that previously bred to determine their survival and future nonbreeding probabilities. Food availability varied during the study, being generally poor at the onset, and improving in later years. Adult nest attendance and body condition (assessed late in the chick- rearing period) varied accordingly among years, and between adults raising chicks and adults that had their eggs removed. Adults from unmanipulated nests incurred significant survival costs in all years, although fecundity costs were strongly expressed in only one of four years. Neither survival nor fecundity costs were strongly influenced by body condition or food availability, and no difference in reproductive costs was detected between the sexes. Although unmanipulated breeders survived at lower rates than manipulated breeders due to costs of reproduction, their survival rates were elevated compared to those of natural nonbreeders, presumably due to differences in individual ability. These findings indicate that models of adult survival must consider not only an organism's reproductive state, but also the factors that lead to that state. Although body condition appeared to be weakly related to survival, it was insufficient to explain the full magnitude of survival costs observed. We suggest that other parameters that were found to differ between treatment groups (e.g., rates of energy turnover, baseline levels of stress, and patterns of allocating body reserves) may be important mechanistic determinants of reproductive costs in kittiwakes, and potentially other long-lived species. Future efforts should move beyond simple assessments of body condition and toward more integrated measures of physiological condition when attempting to identify factors that influence how long-lived species balance the costs and benefits of reproduction.

Alaska

Geographic structure of adelie penguin populations: overlap in colony-specific foraging areas

In an investigation of the factors leading to geographic structuring among Ade??lie Penguin (Pygoscelis adeliae) populations, we studied the size and overlap of colony-specific foraging areas within an isolated cluster of colonies. The study area, in the southwestern Ross Sea, included one large and three smaller colonies, ranging in size from 3900 to 135000 nesting pairs, clustered on Ross and Beaufort Islands. We used triangulation of radio signals from transmitters attached to breeding penguins to determine foraging locations and to define colony-specific foraging areas during the chick-provisioning period of four breeding seasons, 1997-2000. Colony populations (nesting pairs) were determined using aerial photography just after egg-laying; reproductive success was estimated by comparing ground counts of chicks fledged to the number of breeding pairs apparent in aerial photos. Foraging-trip duration, meal size, and adult body mass were estimated using RFID (radio frequency identification) tags and an automated reader and weighbridge. Chick growth was assessed by weekly weighing. We related the following variables to colony size: foraging distance, area, and duration; reproductive success; chick meal size and growth rate; and seasonal variation in adult body mass. We found that penguins foraged closest to their respective colonies, particularly at the smaller colonies. However, as the season progressed, foraging distance, duration, and area increased noticeably, especially at the largest colony. The foraging areas of the smaller colonies overlapped broadly, but very little foraging area overlap existed between the large colony and the smaller colonies, even though the foraging area of the large colony was well within range of the smaller colonies. Instead, the foraging areas of the smaller colonies shifted as that of the large colony grew. Colony size was not related to chick meal size, chick growth, or parental body mass. This differed from the year previous to the study, when foraging trips of the large colony were very long, parents lost mass, and chick meals were smaller. In light of existing data on prey abundance in neritic waters in Antarctica suggesting that krill are relatively evenly distributed and in high abundance in the Southern Ross Sea, we conclude that penguins depleted or changed the availability of their prey, that the degree of alteration was a function of colony size, and that the large colony affected the location (and perhaps ultimately the size) of foraging areas for the smaller colonies. It appears, therefore, that foraging dynamics play a role in the geographic structuring of colonies in this species. ?? 2004 by the Ecological Society of America.

Ecological Monographs

Hierarchical spatial models of abundance and occurrence from imperfect survey data

Many estimation and inference problems arising from large-scale animal surveys are focused on developing an understanding of patterns in abundance or occurrence of a species based on spatially referenced count data. One fundamental challenge, then, is that it is generally not feasible to completely enumerate ('census') all individuals present in each sample unit. This observation bias may consist of several components, including spatial coverage bias (not all individuals in the Population are exposed to sampling) and detection bias (exposed individuals may go undetected). Thus, observations are biased for the state variable (abundance, occupancy) that is the object of inference. Moreover, data are often sparse for most observation locations, requiring consideration of methods for spatially aggregating or otherwise combining sparse data among sample units. The development of methods that unify spatial statistical models with models accommodating non-detection is necessary to resolve important spatial inference problems based on animal survey data. In this paper, we develop a novel hierarchical spatial model for estimation of abundance and occurrence from survey data wherein detection is imperfect. Our application is focused on spatial inference problems in the Swiss Survey of Common Breeding Birds. The observation model for the survey data is specified conditional on the unknown quadrat population size, N(s). We augment the observation model with a spatial process model for N(s), describing the spatial variation in abundance of the species. The model includes explicit sources of variation in habitat structure (forest, elevation) and latent variation in the form of a correlated spatial process. This provides a model-based framework for combining the spatially referenced samples while at the same time yielding a unified treatment of estimation problems involving both abundance and occurrence. We provide a Bayesian framework for analysis and prediction based on the integrated likelihood, and we use the model to obtain estimates of abundance and occurrence maps for the European Jay (Garrulus glandarius), a widespread, elusive, forest bird. The naive national abundance estimate ignoring imperfect detection and incomplete quadrat coverage was 77 766 territories. Accounting for imperfect detection added approximately 18 000 territories, and adjusting for coverage bias added another 131 000 territories to yield a fully corrected estimate of the national total of about 227 000 territories. This is approximately three times as high as previous estimates that assume every territory is detected in each quadrat.

Ecological Monographs

Scaling the effects of moose browsing on forage distribution, from the geometry of plant canopies to landscapes

Landscape heterogeneity influences large herbivores by altering their feeding rates, but as herbivores attempt to maximize feeding rates they also create spatial heterogeneity by altering plant growth. Herbivore feeding rates thus provide a quantitative link between the causes and consequences of spatial heterogeneity in herbivore-dominated ecosystems. The fractal geometry of plant canopies determines both the density and mass of twigs available to foraging herbivores. These properties determine a threshold distance between plants (d * ) that distinguishes the mechanisms regulating herbivore intake rates. When d * is greater than the actual distance between plants (d), intake is regulated by the rate of food processing in the mouth. But when d * < d, intake is regulated by the rate at which the herbivore encounters new plants. Alterations to plant geometry due to past browsing could change the rate at which herbivores encounter and process bites of plant tissue, modify d * relative to d, and thus change intake rates and the distribution of mechanisms regulating it across landscapes. We measured changes in the geometry of aspen (Populus tremuloides) and balsam fir (Abies balsamea) saplings along gradients of moose browsing from 2001 to 2005 at Isle Royale National Park, Michigan, USA. For aspen saplings, fractal dimension of bite density, bite mass, and forage biomass responded quadratically to increasing moose browsing and were greatest at -3-4 g-g.m-2.yr" 1 consumption. For balsam fir, in contrast, these same measures declined steadily with increasing moose browsing. The different responses of plant canopies to increased browsing altered d * around plants. In summer, d * > d for aspen saplings at all prior consumption levels. Food processing therefore regulated summer moose feeding rates across our landscapes. In winter, changes in bite mass due to past browsing were sufficient to cause d * < d for aspen and balsam fir. Therefore, travel velocity and food processing jointly regulated intake rate during winter. Browsing-induced changes in the small-scale geometry of plant canopies can determine intake rate at larger spatial scales by changing d * relative to d and, hence, which mechanisms determine intake rate, essentially altering how herbivores sense the distribution of their food resources. ?? 2009 by the Ecological Society of America.

Ecological Monographs

Sensitivity of the carbon cycle in the Arctic to climate change

The recent warming in the Arctic is affecting a broad spectrum of physical, ecological, and human/cultural systems that may be irreversible on century time scales and have the potential to cause rapid changes in the earth system. The response of the carbon cycle of the Arctic to changes in climate is a major issue of global concern, yet there has not been a comprehensive review of the status of the contemporary carbon cycle of the Arctic and its response to climate change. This review is designed to clarify key uncertainties and vulnerabilities in the response of the carbon cycle of the Arctic to ongoing climatic change. While it is clear that there are substantial stocks of carbon in the Arctic, there are also significant uncertainties associated with the magnitude of organic matter stocks contained in permafrost and the storage of methane hydrates beneath both subterranean and submerged permafrost of the Arctic. In the context of the global carbon cycle, this review demonstrates that the Arctic plays an important role in the global dynamics of both CO 2 and CH 4 . Studies suggest that the Arctic has been a sink for atmospheric CO 2 of between 0 and 0.8 Pg C/yr in recent decades, which is between 0% and 25% of the global net land/ocean flux during the 1990s. The Arctic is a substantial source of CH 4 to the atmosphere (between 32 and 112 Tg CH 4 /yr), primarily because of the large area of wetlands throughout the region. Analyses to date indicate that the sensitivity of the carbon cycle of the Arctic during the remainder of the 21st century is highly uncertain. To improve the capability to assess the sensitivity of the carbon cycle of the Arctic to projected climate change, we recommend that (1) integrated regional studies be conducted to link observations of carbon dynamics to the processes that are likely to influence those dynamics, and (2) the understanding gained from these integrated studies be incorporated into both uncoupled and fully coupled carbon&ndash;climate modeling efforts.

Ecological Monographs

On the specification of structural equation models for ecological systems

The use of structural equation modeling (SEM) is often motivated by its utility for investigating complex networks of relationships, but also because of its promise as a means of representing theoretical concepts using latent variables. In this paper, we discuss characteristics of ecological theory and some of the challenges for proper specification of theoretical ideas in structural equation models (SE models). In our presentation, we describe some of the requirements for classical latent variable models in which observed variables (indicators) are interpreted as the effects of underlying causes. We also describe alternative model specifications in which indicators are interpreted as having causal influences on the theoretical concepts. We suggest that this latter nonclassical specification (which involves another variable type—the composite) will often be appropriate for ecological studies because of the multifaceted nature of our theoretical concepts. In this paper, we employ the use of meta‐models to aid the translation of theory into SE models and also to facilitate our ability to relate results back to our theories. We demonstrate our approach by showing how a synthetic theory of grassland biodiversity can be evaluated using SEM and data from a coastal grassland. In this example, the theory focuses on the responses of species richness to abiotic stress and disturbance, both directly and through intervening effects on community biomass. Models examined include both those based on classical forms (where each concept is represented using a single latent variable) and also ones in which the concepts are recognized to be multifaceted and modeled as such. To address the challenge of matching SE models with the conceptual level of our theory, two approaches are illustrated, compositing and aggregation. Both approaches are shown to have merits, with the former being preferable for cases where the multiple facets of a concept have widely differing effects in the system and the latter being preferable where facets act together consistently when influencing other parts of the system. Because ecological theory characteristically deals with concepts that are multifaceted, we expect the methods presented in this paper will be useful for ecologists wishing to use SEM.

Ecological Monographs

Causes and implications of the correlation between forest productivity and tree mortality rates

At global and regional scales, tree mortality rates are positively correlated with forest net primary productivity (NPP). Yet causes of the correlation are unknown, in spite of potentially profound implications for our understanding of environmental controls of forest structure and dynamics and, more generally, our understanding of broad-scale environmental controls of population dynamics and ecosystem processes. Here we seek to shed light on the causes of geographic patterns in tree mortality rates, and we consider some implications of the positive correlation between mortality rates and NPP. To reach these ends, we present seven hypotheses potentially explaining the correlation, develop an approach to help distinguish among the hypotheses, and apply the approach in a case study comparing a tropical and temperate forest. Based on our case study and literature synthesis, we conclude that no single mechanism controls geographic patterns of tree mortality rates. At least four different mechanisms may be at play, with the dominant mechanisms depending on whether the underlying productivity gradients are caused by climate or soil fertility. Two of the mechanisms are consequences of environmental selection for certain combinations of life-history traits, reflecting trade-offs between growth and defense (along edaphic productivity gradients) and between reproduction and persistence (as manifested in the adult tree stature continuum along climatic and edaphic gradients). The remaining two mechanisms are consequences of environmental influences on the nature and strength of ecological interactions: competition (along edaphic gradients) and pressure from plant enemies (along climatic gradients). For only one of these four mechanisms, competition, can high mortality rates be considered to be a relatively direct consequence of high NPP. The remaining mechanisms force us to adopt a different view of causality, in which tree growth rates and probability of mortality can vary with at least a degree of independence along productivity gradients. In many cases, rather than being a direct cause of high mortality rates, NPP may remain high in spite of high mortality rates. The independent influence of plant enemies and other factors helps explain why forest biomass can show little correlation, or even negative correlation, with forest NPP.

Ecological Monographs

Decomposition of heterogeneous organic matterand its long-term stabilization in soils

Soil organic matter is a complex mixture of material with heterogeneous biological, physical, and chemical properties. Decomposition models represent this heterogeneity either as a set of discrete pools with different residence times or as a continuum of qualities. It is unclear though, whether these two different approaches yield comparable predictions of organic matter dynamics. Here, we compare predictions from these two different approaches and propose an intermediate approach to study organic matter decomposition based on concepts from continuous models implemented numerically. We found that the disagreement between discrete and continuous approaches can be considerable depending on the degree of nonlinearity of the model and simulation time. The two approaches can diverge substantially for predicting long-term processes in soils. Based on our alternative approach, which is a modification of the continuous quality theory, we explored the temporal patterns that emerge by treating substrate heterogeneity explicitly. The analysis suggests that the pattern of carbon mineralization over time is highly dependent on the degree and form of nonlinearity in the model, mostly expressed as differences in microbial growth and efficiency for different substrates. Moreover, short-term stabilization and destabilization mechanisms operating simultaneously result in long-term accumulation of carbon characterized by low decomposition rates, independent of the characteristics of the incoming litter. We show that representation of heterogeneity in the decomposition process can lead to substantial improvements in our understanding of carbon mineralization and its long-term stability in soils.

Ecological Monographs

Food-web dynamics in a large river discontinuum

Nearly all ecosystems have been altered by human activities, and most communities are now composed of interacting species that have not co-evolved. These changes may modify species interactions, energy and material flows, and food-web stability. Although structural changes to ecosystems have been widely reported, few studies have linked such changes to dynamic food-web attributes and patterns of energy flow. Moreover, there have been few tests of food-web stability theory in highly disturbed and intensely managed freshwater ecosystems. Such synthetic approaches are needed for predicting the future trajectory of ecosystems, including how they may respond to natural or anthropogenic perturbations. We constructed flow food webs at six locations along a 386-km segment of the Colorado River in Grand Canyon (Arizona, USA) for three years. We characterized food-web structure and production, trophic basis of production, energy efficiencies, and interaction-strength distributions across a spatial gradient of perturbation (i.e., distance from Glen Canyon Dam), as well as before and after an experimental flood. We found strong longitudinal patterns in food-web characteristics that strongly correlated with the spatial position of large tributaries. Above tributaries, food webs were dominated by nonnative New Zealand mudsnails (62% of production) and nonnative rainbow trout (100% of fish production). The simple structure of these food webs led to few dominant energy pathways (diatoms to few invertebrate taxa to rainbow trout), large energy inefficiencies (i.e., <20% of invertebrate production consumed by fishes), and right-skewed interaction-strength distributions, consistent with theoretical instability. Below large tributaries, invertebrate production declined ∼18-fold, while fish production remained similar to upstream sites and comprised predominately native taxa (80–100% of production). Sites below large tributaries had increasingly reticulate and detritus-based food webs with a higher prevalence of omnivory, as well as interaction strength distributions more typical of theoretically stable food webs (i.e., nearly twofold higher proportion of weak interactions). Consistent with theory, downstream food webs were less responsive to the experimental flood than sites closest to the dam. We show how human-induced shifts to food-web structure can affect energy flow and interaction strengths, and we show that these changes have consequences for food-web function and response to perturbations.

Arizona

A guide to Bayesian model selection for ecologists

The steady upward trend in the use of model selection and Bayesian methods in ecological research has made it clear that both approaches to inference are important for modern analysis of models and data. However, in teaching Bayesian methods and in working with our research colleagues, we have noticed a general dissatisfaction with the available literature on Bayesian model selection and multimodel inference. Students and researchers new to Bayesian methods quickly find that the published advice on model selection is often preferential in its treatment of options for analysis, frequently advocating one particular method above others. The recent appearance of many articles and textbooks on Bayesian modeling has provided welcome background on relevant approaches to model selection in the Bayesian framework, but most of these are either very narrowly focused in scope or inaccessible to ecologists. Moreover, the methodological details of Bayesian model selection approaches are spread thinly throughout the literature, appearing in journals from many different fields. Our aim with this guide is to condense the large body of literature on Bayesian approaches to model selection and multimodel inference and present it specifically for quantitative ecologists as neutrally as possible. We also bring to light a few important and fundamental concepts relating directly to model selection that seem to have gone unnoticed in the ecological literature. Throughout, we provide only a minimal discussion of philosophy, preferring instead to examine the breadth of approaches as well as their practical advantages and disadvantages. This guide serves as a reference for ecologists using Bayesian methods, so that they can better understand their options and can make an informed choice that is best aligned with their goals for inference.

Ecological Monographs

Using spatiotemporal statistical models to estimate animal abundance and infer ecological dynamics from survey counts

Ecologists often fit models to survey data to estimate and explain variation in animal abundance. Such models typically require that animal density remains constant across the landscape where sampling is being conducted, a potentially problematic assumption for animals inhabiting dynamic landscapes or otherwise exhibiting considerable spatiotemporal variation in density. We review several concepts from the burgeoning literature on spatiotemporal statistical models, including the nature of the temporal structure (i.e., descriptive or dynamical) and strategies for dimension reduction to promote computational tractability. We also review several features as they specifically relate to abundance estimation, including boundary conditions, population closure, choice of link function, and extrapolation of predicted relationships to unsampled areas. We then compare a suite of novel and existing spatiotemporal hierarchical models for animal count data that permit animal density to vary over space and time, including formulations motivated by resource selection and allowing for closed populations. We gauge the relative performance (bias, precision, computational demands) of alternative spatiotemporal models when confronted with simulated and real data sets from dynamic animal populations. For the latter, we analyze spotted seal ( Phoca largha ) counts from an aerial survey of the Bering Sea where the quantity and quality of suitable habitat (sea ice) changed dramatically while surveys were being conducted. Simulation analyses suggested that multiple types of spatiotemporal models provide reasonable inference (low positive bias, high precision) about animal abundance, but have potential for overestimating precision. Analysis of spotted seal data indicated that several model formulations, including those based on a log-Gaussian Cox process, had a tendency to overestimate abundance. By contrast, a model that included a population closure assumption and a scale prior on total abundance produced estimates that largely conformed to our a priori expectation. Although care must be taken to tailor models to match the study population and survey data available, we argue that hierarchical spatiotemporal statistical models represent a powerful way forward for estimating abundance and explaining variation in the distribution of dynamical populations.

Ecological Monographs

State-space modeling to support management of brucellosis in the Yellowstone bison population

The bison ( Bison bison ) of the Yellowstone ecosystem, USA, exemplify the difficulty of conserving large mammals that migrate across the boundaries of conservation areas. Bison are infected with brucellosis ( Brucella abortus ) and their seasonal movements can expose livestock to infection. Yellowstone National Park has embarked on a program of adaptive management of bison, which requires a model that assimilates data to support management decisions. We constructed a Bayesian state-space model to reveal the influence of brucellosis on the Yellowstone bison population. A frequency-dependent model of brucellosis transmission was superior to a density-dependent model in predicting out-of-sample observations of horizontal transmission probability. A mixture model including both transmission mechanisms converged on frequency dependence. Conditional on the frequency-dependent model, brucellosis median transmission rate was 1.87 yr &minus;1 . The median of the posterior distribution of the basic reproductive ratio ( R 0 ) was 1.75. Seroprevalence of adult females varied around 60% over two decades, but only 9.6 of 100 adult females were infectious. Brucellosis depressed recruitment; estimated population growth rate &lambda; averaged 1.07 for an infected population and 1.11 for a healthy population. We used five-year forecasting to evaluate the ability of different actions to meet management goals relative to no action. Annually removing 200 seropositive female bison increased by 30-fold the probability of reducing seroprevalence below 40% and increased by a factor of 120 the probability of achieving a 50% reduction in transmission probability relative to no action. Annually vaccinating 200 seronegative animals increased the likelihood of a 50% reduction in transmission probability by fivefold over no action. However, including uncertainty in the ability to implement management by representing stochastic variation in the number of accessible bison dramatically reduced the probability of achieving goals using interventions relative to no action. Because the width of the posterior predictive distributions of future population states expands rapidly with increases in the forecast horizon, managers must accept high levels of uncertainty. These findings emphasize the necessity of iterative, adaptive management with relatively short-term commitment to action and frequent reevaluation in response to new data and model forecasts. We believe our approach has broad applications.

Wyoming

The ecology of methane in streams and rivers: Patterns, controls, and global significance

Streams and rivers can substantially modify organic carbon (OC) inputs from terrestrial landscapes, and much of this processing is the result of microbial respiration. While carbon dioxide (CO 2 ) is the major end-product of ecosystem respiration, methane (CH 4 ) is also present in many fluvial environments even though methanogenesis typically requires anoxic conditions that may be scarce in these systems. Given recent recognition of the pervasiveness of this greenhouse gas in streams and rivers, we synthesized existing research and data to identify patterns and drivers of CH 4 , knowledge gaps, and research opportunities. This included examining the history of lotic CH 4 research, creating a database of concentrations and fluxes (MethDB) to generate a global-scale estimate of fluvial CH 4 efflux, and developing a conceptual framework and using this framework to consider how human activities may modify fluvial CH 4 dynamics. Current understanding of CH 4 in streams and rivers has been strongly influenced by goals of understanding OC processing and quantifying the contribution of CH 4 to ecosystem C fluxes. Less effort has been directed towards investigating processes that dictate in situ CH 4 production and loss. CH 4 makes a meager contribution to watershed or landscape C budgets, but streams and rivers are often significant CH 4 sources to the atmosphere across these same spatial extents. Most fluvial systems are supersaturated with CH 4 and we estimate an annual global emission of 26.8 Tg CH 4 , equivalent to ~15-40% of wetland and lake effluxes, respectively. Less clear is the role of CH 4 oxidation, methanogenesis, and total anaerobic respiration to whole ecosystem production and respiration. Controls on CH 4 generation and persistence can be viewed in terms of proximate controls that influence methanogenesis (organic matter, temperature, alternative electron acceptors, nutrients) and distal geomorphic and hydrologic drivers. Multiple controls combined with its extreme redox status and low solubility result in high spatial and temporal variance of CH 4 in fluvial environments, which presents a substantial challenge for understanding its larger-scale dynamics. Further understanding of CH 4 production and consumption, anaerobic metabolism, and ecosystem energetics in streams and rivers can be achieved through more directed studies and comparison with knowledge from terrestrial, wetland, and aquatic disciplines.

Ecological Monographs

Interspecific competition among Hawaiian forest birds

The object of this study was to determine whether interspecific competition modified local geographic distribution, after taking into account the effect of habitat structure. The tendencies for 14 passerine birds to have positive or negative associations were examined, using 7861 sample points in seven native forests on the islands of Hawaii, Maui, and Kauai. All birds were at least partly insectivorous and were fairly common in forested areas, although some fed chiefly on nectar or fruit. Species—pairs were classified as primary or secondary potential competitors based on general dietary similarity. To evaluate the association between species and to account for the effect of individual species habitat preferences, partial correlations were computed for each species—pair in a study area from the simple correlations between the species and 26 habitat variables plus two quadratic terms to represent nonlinearity. The partial correlations represented a short—term ("instantaneous") assessment of the strength of competitive interactions, and did not reflect the accumulation of competitive displacement through time. Of 170 partial correlations in the analysis, only 10 indicated significant negative association. The general pattern was of positive association (76 significantly positive partials), which probably resulted from flocking and from attraction of birds to areas of resource superabundance. Two species showed consistent patterns of negative partial correlations over several adjacent study areas, the Japanese White—eye/Iiwi in montane Hawaii, and the Japanese White—eye/Elepaio in windward Hawaii; both patterns could be reasonably attributed to direct competition. Species—pairs were grouped by the native or exotic status of the component species. Native/exotic pairs had a significantly greater proportion of negative partial correlations (37%) than either native/native pairs (8%) or exotic/exotic pairs (0%). This pattern was consistent across the seven study areas and appeared to reflect the occurrence of interspecific competition along a broad and diffuse ecological "front" between a co—evolved native avifauna and recently introduced exotic species. The role of competition in the pattern was corroborated by the significantly higher proportion of negative partial correlations among species—pairs of primary potential competitors than among those of secondary potential competitors. Our results suggested that °47% of the primary potential competitors among native/exotic species—pairs may experience at least small depressions in local population density due to competition. Although the negative correlations were for the most part small (average negative r = 0.06), one species could eventually replace another as spatial displacement accumulated through time. The Japanese White—eye appeared to have a principal role in native/exotic interactions, with 62% of the partial correlations between it and native primary potential competitor species being negative. Noteworthy implications were that (1) it was important to account for the habitat responses of individual species when studying the role of interspecific competition in modifying small—scale geographic distribution; (2) competition was frequently sporadic in its geographic occurrence and in the species affected, thus supporting Wiens' (1977) theory of competition; and (3) as a consequence, the role of interspecific competition in modifying distribution may be difficult to detect statistically with small data sets.

Hawaii

Population differentiation along a flood frequency gradient: Physiological adaptations to flooding in Nyssa sylvatica

Throughout the southeastern United States the hardwood Nyssa sylvatica (sensu lato) is distributed along a soil moisture gradient from upland sites, which are never flooded, to floodplains, which are periodically flooded and drained to permanently flooded swamps. Population differentiation with respect to flood tolerance and related physiological attributes was investigated using 1—year—old seedlings grown in a greenhouse from seed collected along this gradient. Upland plants were very intolerant of flooded soils. Their root systems deteriorated, root respiration rates dropped and, after a year under such conditions, survival was poor and those that did remain were greatly stunted and had accumulated large concentrations of many nutrient elements. In contrast swamp plants were quite tolerant of flooded soils. Upon flooding, parts of the orginal root system were lost but new roots were initiated which had an increased capacity for alcoholic fermentation. Many of these new roots were more succulent, larger in diameter, and less branched than drained roots. Such succulent roots however were only a temporary response to short—term flooding; plants flooded for a year did not have such roots, rather the root system superficially resembled drained roots. Concomitant with this return to drained—like roots was an increase in internal oxygen transport to the roots and a drop in alcoholic fermentation. Floodplain plants under drained conditions allocated less biomass to roots than to shoots and had high respiration rates, traits similar to upland plants. Under flooded conditions they initiated new roots with medium respiration rates, allocated less biomass to roots than to shoots, significantly increased oxygen transport to the roots and had high survival, traits similar to swamp plants. Thus, the floodplain population produced a distinctly flood—tolerant phenotype; but not nearly as tolerant of flooded conditions as the swamp phenotype. Floodplain plants differed from swamp plants in transporting less oxygen to the roots under drained conditions, initiating fewer succulent—type roots and not accelerating alcoholic fermentation upon flooding and after a year under flooded conditions having less total biomass, less oxygen transport to the roots and a greater accumulation of Fe and Mn in the roots. The floodplain plants apparently have been selected to be similar to upland plants under drained conditions and swamp plants under flooding and one consequence of this is that their tolerance of flooded conditions is intermediate. It is argued that one of the more important trade—offs in adapting to flooded conditions is that high internal oxygen transport carries with it a ‘cost’ in terms of excessive water loss under water stress conditions.

Ecological Monographs

Populations and home range relationships of the box turtle, Terrapene c. carolina (Linnaeus)

SUMMARY: A population study of the box turtle (Terrapene c. carolina Linnaeus) was made during the years 1944 to 1947 at the Patuxent Research Refuge, Maryland. A thirty acre area in well drained bottomland forest on the flood plain of the Patuxent River was selected for intensive study. Similarly forested land extended in all directions from the study plot. Markers were established at eighty-three foot intervals over the study plot for reference in recording locality data. Individuals were marked by filing notches in the marginal scutes according to a code system. There were 2109 collections of study area turtles. Records of collecting sites and turtle behavior showed that in the bottomlands habitat cover is utilized extensively during the day as well as at night. Turtles not actively moving about are almost always found in or around brush piles, heaps of debris, and tangles of vines and briars. Gully banks and woods openings are used for sunning. Turtles are occasionally found in the mud or water of the gullies. The commonest type of night retreat is a cavity constructed by the turtle in leaves, debris, or earth. These cavities, termed 'forms,' may be used only once, but are sometimes used repeatedly, often at intervals of several days or more. Different turtles sometimes use the same form on successive nights. Weather conditions most favorable to turtle activity are high humidity, warm sunny days, and frequent rains. The most unfavorable influences are low temperatures and drought. On most summer days there are some active turtles but individual turtles are not active every day. Periods of activity are alternated with periods of quiet even in favorable weather. This behavior is most pronounced in early spring and late fall when inactive days are often more numerous than active ones. Adult turtles occupy specific home ranges which they maintain from year to year. The turtles living in the study plot retained their ranges even through a flood that completely covered the area. Maximum home range diameters were determined by measurements of the mapped ranges of individual turtles. The average range of adult males was 330 feet, adult females 370 feet. The difference between male and female ranges was not statistically significant. There was no evidence of defense of territory. Ranges of turtles of all ages and both sexes overlapped grossly. Turtles were frequently found near each other and no antagonistic behavior was observed. A trail-laying device was developed in order to follow individual travel routes. The trailer consists of a light weight housing fastened to the turtle's back. It contains a spool of white thread that unwinds as the turtle moves, thus marking its exact route. Turtles selected for this more detailed study were followed with trailers for a total of 456 turtle days. Maps illustrating their travels are shown. Normal movements within the home range are characterized by, (1) turns, doublings, detours, and criss-crossing paths completely covering the area, (2) interspersion of fairly direct traverses of the home range, (3) frequently repeated travels over certain paths or routes. Trailer records and mapped collection records both show that the maximum limits of the home range are ordinarily reached within a few days or weeks. This general procedure is varied by some turtles to include intensive coverage of only one portion of the range at a time. Some turtles have two home ranges and travel between them at infrequent intervals. One turtle showing this behavior was followed with a trailer for 161 days during 1946 and 1947. Trips outside the home range are made by some turtles. These include egg laying trips by females as well as trips of unexplained nature made by both males and females. Turtles from other areas occasionally occur as transients in the study plot. The size of the population was estimated on the basis of collections during on

Ecological Monographs

Sea otters and kelp forests in Alaska: Generality and variation in a community ecological paradigm

Multiscale patterns of spatial and temporal variation in density and population structure were used to evaluate the generality of a three—trophic—level cascade among sea otters (Enhydra lutris), invertebrate herbivores, and macroalgae in Alaska. The paradigm holds that where sea otters occur herbivores are rare and plants are abundant, whereas when sea otters are absent herbivores are relatively common and plants are rare. Spatial patterns were based on 20 randomly placed quadrats at 153 randomly selected sites distributed among five locations with and four locations without sea otters. Both sea urchin and kelp abundance differed significantly among locations with vs. without sea otters in the Aleutian Islands and southeast Alaska. There was little (Aleutian Islands) or no (southeast Alaska) overlap between sites with and without sea otters, in plots of kelp density against urchin biomass. Despite intersite variation in the abundance of kelps and herbivores, these analyses demonstrate that sea otter predation has a predictable and broadly generalizable influence on the structure of Alaskan kelp forests. The percent cover of algal turf and suspension feeder assemblages also differed significantly (although less dramatically) between locations with and without sea otters. Temporal variation in community structure was assessed over periods of from 3 to 15 yr at sites in the Aleutian Islands and southeast Alaska where sea otters were 1) continuously present, 2) continuously absent, or 3) becoming reestablished because of natural range expansion. Kelp and sea urchin abundance remained largely unchanged at most sites where sea otters were continuously present or absent, the one exception being at Torch Bay (southeast Alaska), where kelp abundance varied significantly through time and urchin abundance varied significantly among sites because of episodic and patchy disturbances. In contrast, kelp and sea urchin abundances changed significantly, and in the expected directions, at sites that were being recolonized by sea otters. Sea urchin biomass declined by 50% in the Aleutian Islands and by nearly 100% in southeast Alaska following the spread of sea otters into previously unoccupied habitats. In response to these different rates and magnitudes of urchin reduction by sea otter predation, increases in kelp abundance were abrupt and highly significant in southeast Alaska but much smaller and slower over similar time periods in the Aleutian Islands. The different kelp colonization rates between southeast Alaska and the Aleutian Islands appear to be caused by large—scale differences in echinoid recruitment coupled with size—selective predation by sea otters for larger urchins. The length of urchin jaws (correlated with test diameter, r 2 = 0.968) in sea otter scats indicates that sea urchins <15—20 mm test diameter are rarely eaten by foraging sea otters. Sea urchin populations in the Aleutian Islands included high densities of small individuals (<20 mm test diameter) at all sites and during all years sampled, whereas in southeast Alaska similarly sized urchins were absent from most populations during most years. Small (<30—35 mm test diameter) tetracycline—marked urchins in the Aleutian Islands grew at a maximum rate of °10 mm/yr; thus the population must have significant recruitment annually, or at least every several years. In contrast, echinoid recruitment in southeast Alaska was more episodic, with many years to perhaps decades separating significant events. Our findings help explain regional differences in recovery rates of kelp forests following recolonization by sea otters.

Alaska

Population dynamics and bioenergetics of a fossorial herbivore, Thomomys talpoides (Rodentia: Geomyidae), in a spruce-fir sere

Studies of the bioenergetics of the northern pocket gopher, Thomomys talpoides , are coupled with data on demography, activity budgets, and microclimates to model the energy requirements of individuals and populations in the Wasatch Mountains of Utah during 1976-1979. Metabolic rates during rest increased linearly with decreasing ambient temperature, but burrowing metabolic rates (16.3 mL O 2 • h -1 • g -9.75 ) were independent of both temperature and physical properties of the soil. Radio-telemetry studies indicated that free-ranging gophers are active =50% of each day. Conservative estimates of true energy consumption were calculated using estimates of habitat-specific minimum daily burrowing requirements. Rates of burrowing measured in the laboratory were either ∞ 0.0 or ∞ 2.0 cm/min. The low burrowing rate was observed when the soil was frozen or saturated with water, as would occur in the field in early winter and in spring, respectively. Gophers burrowed through soil at the study site at an average rate of ∞ 1.5 cm/min. Belowground food energy densities at gopher foraging depth declined from 24.6 to 3.2 J/cm 3 along a successional gradient (subalpine forb meadow to Engelmann spruce dominated forest). We conclude that individual gophers are food limited within the climax spruce seral stage. Further, daily energy costs associated with reproduction in females may exceed the belowground energy supply available in intermediate seral stages (aspen and subalpine fir). Reduction of burrowing rates for any reason will affect gophers in the late seral stages proportionately more than those resident in the meadow. The peak gopher densities recorded (from 62 individuals/ha in the meadow to 2 individuals/ha in spruce forest) support these inferences. Detailed demographic information was obtained only in the meadow seral stage. Adult survivorship was lower in winter than in summer and varied greatly between years (0.18-0.70 yr -1 ). Juvenile survivorship from weaning through the first year was comparable to adult annual rates. The fertility rate was 3.75 young • female -1 • yr -1 . The energy supply and demand analyses indicate that the growth of Thomomys talpoides populations in the early seral stages is seldom directly limited by the amount of food present. From our demographic, environmental, and autecological studies we conclude that stochastic events associated with weather affect energy acquisition (burrowing) rates, and thus survivorship. In montane environments, such events may prevent populations from attaining sizes at which territorial behavior would hypothetically limit further increases. The energy flow through the meadow population at moderate to high )1976-1977) densities (at least 1100 MJ • ha -1 • yr -1 ) indicates that pocket gophers are proficient energy movers relative to non-fossorial small mammals. Subalpine T. talpoides populations appear commonly to attain such densities. More than 30% of the annual primary productivity allocated to belowground parts of meadow forbs may be consumed by gophers.

Ecological Monographs