Spatial scaling of avian population dynamics: population abundance, growth rate, and variability

Synchrony in population fluctuations has been identified as an important component of population dynamics. In a previous study, we determined that local-scale (<15-km) spatial synchrony of bird populations in New England was correlated with synchronous fluctuations in lepidopteran larvae abundance and with the North Atlantic Oscillation. Here we address five questions that extend the scope of our earlier study using North American Breeding Bird Survey data. First, do bird populations in eastern North America exhibit spatial synchrony in abundances at scales beyond those we have documented previously? Second, does spatial synchrony depend on what population metric is analyzed (e.g., abundance, growth rate, or variability)? Third, is there geographic concordance in where species exhibit synchrony? Fourth, for those species that exhibit significant geographic concordance, are there landscape and habitat variables that contribute to the observed patterns? Fifth, is spatial synchrony affected by a species' life history traits? Significant spatial synchrony was common and its magnitude was dependent on the population metric analyzed. Twenty-four of 29 species examined exhibited significant synchrony in population abundance: mean local autocorrelation (rho)= 0.15; mean spatial extent (mean distance where rho=0) = 420.7 km. Five of the 29 species exhibited significant synchrony in annual population growth rate (mean local autocorrelation = 0.06, mean distance = 457.8 km). Ten of the 29 species exhibited significant synchrony in population abundance variability (mean local autocorrelation = 0.49, mean distance = 413.8 km). Analyses of landscape structure indicated that habitat variables were infrequent contributors to spatial synchrony. Likewise, we detected no effects of life history traits on synchrony in population abundance or growth rate. However, short-distance migrants exhibited more spatially extensive synchrony in population variability than either year-round residents or long-distance migrants. The dissimilarity of the spatial extent of synchrony across species suggests that most populations are not regulated at similar spatial scales. The spatial scale of the population synchrony patterns we describe is likely larger than the actual scale of population regulation, and in turn, the scale of population regulation is undoubtedly larger than the scale of individual ecological requirements.     

Multiple-locus heterozygosity,mortality, and growth in a cohort of Mytilus edulis

A positive correlation between the degree of individual heterozygosity (H) at five polymorphic enzyme loci and shell length was reported for a cohort of Mytilus edulis L. sampled in September, 1983, two months after settlement (Koehn and Gaffney, 1984). In the present study, the same cohort was resampled four and eight months after settlement in November, 1983, and April, 1984, respectively. Among individuals four months of age, there was a small, but statistically significant, negative correlation between H and shell length. In this sample, heterozygote deficiency in the cohort was less than that of any size group in the September sample. The reversal of the H-size relationship between September and November suggests that differential mortality, particularly among small, homozygous individuals, occurred in this period. For individuals eight months of age, there was no correlation between H and shell length, there was no heterozygote deficiency in the cohort and the average heterozygosity was the same as that of the largest individuals in September. At the youngest age, there was a positive relationship, between H and growth rate; as aging occurred, differential mortality altered the relationship between H and growth rate. viability differences among heterozygosity classes obscured the original effect of H on growth rate. If there is a positive relationship between heterozygosity and size, it can most likely be detected from sampling a large outbreeding population before differential mortality occurs and before energy is allocated to reproduction.     

Nonparametric MLE incorporation of heterogeneity and model testing into premarked cohort studies

We utilize mixture models and nonparametric maximum likelihood estimation to both develop a likelihood ratio test (lrt) for a common simplifying assumption and to allow heterogeneity within premarked cohort studies. Our methods allow estimation of the entire probability model and thus one can not only estimate many parameters of interest but one can also bootstrap from the estimated model to predict many things, including the standard deviations of estimators. Simulations suggest that our lrt has the appropriate protection for Type I error and often has good power. In practice, our lrt is important for determining the appropriateness of estimators and in examining if a simple design with only one capture period could be utilized for a future similar study.     

Demographic feedback between clonal growth and fragmentation in an invasive seaweed

Many abundant plants, invertebrates, and seaweed are clonal, and this allows the formation of high-density aggregations, foraging, and the placement of modules into new space, and rapid rates of expansion. For these species, population density and rates of expansion are functions of recruitment of asexual modules and post-recruitment vegetative growth and survivorship. In this study, we provide the first experimental test of the relative importance of these two processes in determining the abundance of a clonal seaweed using Caulerpa taxifolia, an invasive green alga that spreads rapidly and reaches very high abundance. We asked two main questions: What is the relative importance to abundance (biomass) of vegetative stolon growth and fragment recruitment during expansion of established patches? Does greater fragment recruitment result in greater abundance in established patches? Vegetative growth of stolons underpinned patch expansion. Plots with stolons growing into them always had a greater abundance than plots where stolons were removed, even when fragment recruitment was increased. Greater recruitment only resulted in greater abundance when stolons were absent, a situation analogous to the establishment of new populations. Although post-recruitment processes were more important in determining abundance during patch expansion, there was greater ambient fragment recruitment when stolons were present compared to when they were absent, and as the abundance of C. taxifolia increased, demonstrating an important feedback between stolon growth, abundance, and fragment recruitment. In established patches, greater fragment recruitment over six months (six levels ranging from 0 to 480 recruits x m(-2) x mo(-1)) had no effect on biomass. Our experiments demonstrate that the rapid expansion and high abundance of invasive C. taxifolia are underpinned by post-recruitment vegetative growth and, during expansion, by a feedback between vegetative growth and asexual fragmentation.     

Demographic models inform selection of biocontrol agents for garlic mustard (Alliaria petiolata).

Nonindigenous invasive plants pose a major threat to natural communities worldwide. Biological control of weeds via selected introduction of their natural enemies can affect control over large spatial areas but also risk nontarget effects. To maximize effectiveness while minimizing risk, weed biocontrol programs should introduce the minimum number of host-specific natural enemies necessary to control an invasive nonindigenous plant. We used elasticity analysis of a matrix model to help inform biocontrol agent selection for garlic mustard (Alliaria petiolata (M. Bieb.) Cavara and Grande). The Eurasian biennial A. petiolata is considered one of the most problematic invaders of temperate forests in North America. Four weevil species in the genus Ceutorhynchus (Coleoptera: Curculionidae) are currently considered potential biocontrol agents. These species attack rosettes (C. scrobicollis), stems (C. roberti, C. alliariae), and seeds (C. constrictus) of A. petiolata. Elasticity analyses using A. petiolata demographic parameters from North America indicated that changes in the rosette-to-flowering-plant transition and changes in fecundity consistently had the greatest impact on population growth rate. These results suggest that attack by the rosette-feeder C. scrobicollis, which reduces overwintering survival, and seed or stem feeders that reduce seed output should be particularly effective. Model outcomes differed greatly as A. petiolata demographic parameters were varied within ranges observed in North America, indicating that successful control of A. petiolata populations may occur under some, but not all, conditions. Using these a priori analyses we predict: (1) rosette mortality and reduction of seed output will be the most important factors determining A. petiolata demography; (2) the root-crown feeder C. scrobicollis will have the most significant impact on A. petiolata demography; (3) releases of single control agents are unlikely to control A. petiolata across its full range of demographic variability; (4) combinations of agents that simultaneously reduce rosette survival and seed production will be required to suppress the most vigorous A. petiolata populations. These predictions can be tested using established long-term monitoring sites coupled with a designed release program. If demographic models can successfully predict biocontrol agent impact on invasive plant populations, a continued dialogue and collaboration between empirical and theoretical approaches may be the key to the development of successful biocontrol tactics for plant invaders in the future.     

Correlations between age, phenotype, and individual contribution to population growth in common terns

There have been numerous reports of changes in phenology, which are frequently attributed to environmental change. Age-dependent change in phenotypic traits, fledgling production, and the timing of events in the life cycle is also widespread. This means that changes in the age structure of a population could generate changes in phenology, which may be incorrectly attributed to environmental change or microevolution. Here, estimates of selection for arrival date, arrival mass, and laying date are compared when age is and is not corrected for. This is achieved using long-term individual-based data collected from a breeding colony of Common Terns (Sterna hirundo) and a novel fitness measure: individual contributions to population growth. The failure to correct for age generated deceptive estimates of selection in eight out of nine comparisons. In six out of nine comparisons, the direction of selection differed between age-corrected and uncorrected estimates. Persistent individual differences were detected: individuals remained within the same part of the phenotype distribution throughout life. The age-corrected estimates of selection were weak and explained little variation in fitness, suggesting that arrival date, arrival mass, and laying date are not under intense selection in this population. These results also demonstrate the importance of correcting for age when identifying factors associated with changes in seabird phenology.     

Habitat structure,population abundance and the opportunity for selection on body weight in the amphipodEogammarus oclairi

The importance of oyster-shell habitat-characteristics (depth of the shell layer or degree of fragmentation) to the amphipodEogammarus oclairi were studied in Grays Harbor estuary (Washington, USA) in regard to their effect on local density ofE. oclairi, the risk of predation by juvenile Dungeness crabsCancer magister, and the opportunity for selection on body weight during pairing.E. oclairi was the most abundant macrofaunal species in the intertidal oyster-shell assemblages (density range = 20 to 8500 amphipods m^–2), and its density was positively correlated with the depth of the oyster-shell layer (r = 0.85,n = 30,p < 0.005). Field experiments showed that amphipod density was much lower when oyster shells were whole (x = 340.7 ± 72.1,n = 10) as opposed to fragmented [x = 41.4 ± 9.3,n = 10;t _c (Welch's approximatet-test) = 13.0,df = 9.3,p < 0.05]. Densities of a predator, juvenileC. magister, were not affected by depth of the shell layer nor by degree of fragmentation (whole shell,x = 14.6 ± 13.0 crab;fragmented shell,x = 18.7 ± 2.4 crab;t _c = 0.83,df = 9.7,p > 0.05). Predation rate on single amphipods by crabs did not differ between habitat types (whole shells vs fragmented shells), but mating pairs were consumed more often in the whole-shell treatment (whole shell,x = 0.9 ± 0.8, fragmented shell,x = 0.1 ± 0.3). There were no differences in the size of single amphipods (both sexes) consumed between treatments, or in the size of the paired males consumed. Field experiments showed that the opportunity for selection (i) on male body weight increased with increasing amphipod density which, in turn, increased with increasing degree of shell fragmentation (whole-shell treatment,i = 0.0014 ± 0.0002, fragmented-shell treatment,i = 0.3756 ± 0.0338). Large spatial and temporal fluctuations in population abundance complicate the evaluation of the importance of selection in determining traits such as body weight.     

Modelling plant resource allocation and growth partitioning in response to environmental heterogeneity

Plant morphological adjustment in response to spatial resource heterogeneity is an important factor that determines the outcomes of plant–plant and plant–environment interactions. In this study, a dynamic model of resource allocation and growth partitioning at the whole-plant level is presented. The aim is to suggest a mechanism by which plants are capable of modifying their resource allocation to favour the growth of their growing parts sited in resource-rich patches. In this model, an individual plant is treated as a population of relatively independent subunits competing for internal resources. The growth decision of individual shoot and root subunits depends on their local endogenous nutrient status. The allocation of nutrients to different shoot parts and root parts is determined by the structure of the vascular networks. No specific partitioning functions and driving coefficients are introduced in the model to coordinate resource allocation and growth partitioning at the whole-plant level. Vascular tissues acquire resources from the nutrient flow passing through them to grow and maintain their activities. The simulation results show that, based on simple rules of nutrient supply, transport and utilization, plants are able to integrate activities at the whole-plant level to allocate proportionally more growth to their growing parts in the most favourable positions.     

Renewable electricity policies,heterogeneity, and cost effectiveness

Renewable electricity policies promote investment in renewable electricity generators and have become increasingly common around the world. Because of intermittency and the composition of other generators in the power system, the value of certain renewable – particularly wind and solar – varies across locations and technologies. This paper investigates the implications of this heterogeneity for the cost effectiveness of renewable electricity policies. A simple model of the power system shows that renewable electricity policies cause different investment mixes. Policies also differ according to their effect on electricity prices, and both factors cause the cost effectiveness to vary across policies. We use a detailed, long-run planning model that accounts for intermittency on an hourly basis to compare the cost effectiveness for a range of policies and alternative parameter assumptions. The differences in cost effectiveness are economically significant, where broader policies, such as an emissions price, outperform renewable electricity policies.     

Emerging prion disease drives host selection in a wildlife population

Infectious diseases are increasingly recognized as an important force driving population dynamics, conservation biology, and natural selection in wildlife populations. Infectious agents have been implicated in the decline of small or endangered populations and may act to constrain population size, distribution, growth rates, or migration patterns. Further, diseases may provide selective pressures that shape the genetic diversity of populations or species. Thus, understanding disease dynamics and selective pressures from pathogens is crucial to understanding population processes, managing wildlife diseases, and conserving biological diversity. There is ample evidence that variation in the prion protein gene (PRNP) impacts host susceptibility to prion diseases. Still, little is known about how genetic differences might influence natural selection within wildlife populations. Here we link genetic variation with differential susceptibility of white-tailed deer to chronic wasting disease (CWD), with implications for fitness and disease-driven genetic selection. We developed a single nucleotide polymorphism (SNP) assay to efficiently genotype deer at the locus of interest (in the 96th codon of the PRNP gene). Then, using a Bayesian modeling approach, we found that the more susceptible genotype had over four times greater risk of CWD infection; and, once infected, deer with the resistant genotype survived 49% longer (8.25 more months). We used these epidemiological parameters in a multi-stage population matrix model to evaluate relative fitness based on genotype-specific population growth rates. The differences in disease infection and mortality rates allowed genetically resistant deer to achieve higher population growth and obtain a long-term fitness advantage, which translated into a selection coefficient of over 1% favoring the CWD-resistant genotype. This selective pressure suggests that the resistant allele could become dominant in the population within an evolutionarily short time frame. Our work provides a rare example of a quantifiable disease-driven selection process in a wildlife population, demonstrating the potential for infectious diseases to alter host populations. This will have direct bearing on the epidemiology, dynamics, and future trends in CWD transmission and spread. Understanding genotype-specific epidemiology will improve predictive models and inform management strategies for CWD-affected cervid populations.     

Heterogeneous population growth, parental effects and genotype–environment interactions of a marine oligochaete

Cultures of asexually reproducing populations of the oligochaete Paranaislitoralis (Müller) collected from six different patches (3 to 50 m apart) on an intertidal mud flat in Flax Pond, New York, on two occasions, June and October 1993, showed significant differences among lines in life span, number of offspring produced, and in finite rate of increase (λ). Although growth rates were significantly lower in October than in June, they were always positive (λ > 1) in the laboratory cultures reared in field-collected sediment, while field data show that the densities of P. litoralis decreased sharply in summer and autumn from a seasonal high in early June. Cultures of worms reared at high densities without renewal of sediment crashed, and effects on individuals were irreversible: worms from late (declining) stages of population growth had a significantly higher mortality and lower reproduction than worms from earlier stages, also when transferred to high-quality food. Genetical analysis using RAPDs (random amplified polymorphic DNA) confirmed the existence of several clones of P. litoralis in our cultures. Experiments where parent and offspring were cultured in sediments of different qualities showed clone–environment interactions in the number of asexual offspring produced, but not in age at first reproduction. Clones also differed in that some showed significant parental effects of sediment quality on life-history characteristics while other clones did not. Our results indicate that P.litoralis populations in Flax Pond are not an example of a population subdivided into a set of permanent source and sink subpopulations, but rather an example of a continuously shifting mosaic of local growth conditions. Received: 21 April 1997 / Accepted: 3 September 1997     

Ontogenetic and sex-specific differences in density-dependent habitat selection of a marine fish population

The spatial dynamics of species are the result of complex interactions between density-independent and density-dependent sources of variability. Disentangling these two sources of variability has challenged ecologists working in both terrestrial and aquatic ecosystems. Using a novel spatially explicit statistical model, we tested for the presence of density-independent and density-dependent habitat selection in yellowfin sole (Limanda aspera) in the eastern Bering Sea. We found specificities in the density-dependent processes operating across ontogeny and particularly with gender. Density-dependent habitat expansion occurred primarily in females, and to a lesser degree in males. These patterns were especially evident in adult stages, while juvenile stages of both sexes exhibited a mix of different dynamics. Association of yellowfin sole with substrate type also varied by sex and to a lesser degree with size, with large females distributed over a wider range of substrates than males. Moreover, yellowfin sole expanded northward as cold subsurface waters retracted in summer, suggesting high sensitivity to arctic warming. Our findings illustrate how marginal habitats can play an important role in buffering density-dependent habitat expansion, with direct implications for resource management. Our spatially explicit modeling approach is effective in evaluating density-dependent spatial dynamics, and can easily be used to test similar hypotheses from a variety of aquatic and terrestrial ecosystems.     

Active density-dependent habitat selection in a controlled population of small mammals

Density-dependent habitat selection has numerous and far-reaching implications to population dynamics and evolutionary processes. Although several studies suggest that organisms choose and occupy high-quality habitats over poorer ones, definitive experiments demonstrating active selection, by the same individuals at the appropriate population scale, are lacking. We conducted a reciprocal food supplementation experiment to assess whether voles would first occupy a habitat receiving extra food, then change their preference to track food supplements moved to another habitat. Meadow voles, as predicted, were more abundant in food-supplemented habitat than in others. Density declined when food supplements ceased because the voles moved to the new habitat receiving extra food. Although males and females appeared to follow different strategies, meadow-vole densities reflected habitat quality because voles actively selected the best habitat available. It is thus clear that behavioral decisions on habitat use can motivate patterns of abundance, frequency, and gene flow that have widespread effects on subsequent evolution.     

Assessment of natural selection in a hybrid population of mussels: evaluation of exogenous vs endogenous selection models

We examined natural selection within a population of marine mussels, sampled in southwestern England in June 1991, containing a high frequency of hybrids between Mytilus edulis L. and M. galloprovincialis Lmk. This system is particularly tractable for the assessment of natural selection because hybridization is common and individual mussels can be aged, allowing changes in the frequency of hybrid genotypes among age classes to be determined. We show that strong viability selection occurs among hybrid genotypes which results in the virtual elimination of M. edulis–like genotypes from the population over a period of 3 years. Recombinant hybrid genotypes are intermediate in fitness, with M. edulis–like genotypes having a lower survival rate and M. galloprovincialis–like genotypes having a higher survival rate than genotypes of mixed ancestry. Since intermediate fitness for hybrid genotypes is inconsistent with endogenous selection models we conclude that the structure and position of this hybrid zone is probably generated by exogenous selection. This pattern of selection is a recurring feature of this hybrid population and likely occurs elsewhere in the hybrid zone. Selection against M. edulis–like genotypes appears to be offset by extensive immigration of larvae dispersed from pure populations of M. edulis. Received: 14 July 1997 / Accepted: 24 February 1998     

A world model of the growth in production and population

A model for the growth in GNP (gross national product) and population has been set up. On the basis of the present data it was found that the growth of a population is a function of GNP, and that GNP seems to follow a logistic growth.     

Spatial selection and inheritance: applying evolutionary concepts to population dynamics in heterogeneous space

Organisms in highly suitable sites generally produce more offspring, and offspring can inherit this suitability by not dispersing far. This combination of spatial selection and spatial inheritance acts to bias the distribution of organisms toward suitable sites and thereby increase mean fitness (i.e., per capita population increase). Thus, population growth rates in heterogeneous space change over time by a process conceptually analogous to evolution by natural selection, opening avenues for theoretical cross-pollination between evolutionary biology and ecology. We operationally define spatial inheritance and spatial selective differential and then combine these two factors in a modification of the breeder's equation, derived from simple models of population growth in heterogeneous space. The modified breeder's equation yields a conservative criterion for persistence in hostile environments estimable from field measurements. We apply this framework for understanding gypsy moth population persistence amidst abundant predators and find that the predictions of the modified breeder's equation match initial changes in population growth rate in independent simulation output. The analogy between spatial dynamics and natural selection conceptually links ecology and evolution, provides a spatially implicit framework for modeling spatial population dynamics, and represents an important null model for studying habitat selection.     

Age structure,growth rates,movement patterns and feeding in an estuarine population of the cardinalfish Apogon rueppellii

The biology of a population of the cardinalfish Apogon rueppellii has been studied over several years (1977–1983) in the Swan Estuary in south-western Australia, using ramples collected monthly from the shallows by beach seine and from various depths by otter trawl. While the life cycle of this species typically lasts for one year, at the end of which time the mean length is 50 to 60 mm, some individuals survive for a further year and attain lengths up to 104 mm. A. rueppellii shows a marked tendency to move offshore into deeper water during the winter months. This tendency is more pronounced in the 1+ than in the 0+ year class and in larger than smaller 0+ individuals. An inshore movement of A. rueppellii in the spring is followed by spawning and by oral brooding by the males, which leads to the recruitment of large numbers of a new 0+ year class on to the banks during the summer. The offshore movement is correlated with changes in salinity and temperature. The larger catches taken by otter trawl during the day than at night indicate that A. rueppellii exhibits a diel pattern of activity. Mean fecundity ranged from 70 in the 45 to 49 mm size class to 345 in the 90 to 94 mm size class. Measurements of fecundity and the number of oral-brooded eggs demonstrated that the majority of the eggs released by the female are collected and incubated by the males. Copepods are ingested in relatively greater amounts by small than by large A. rueppellii, whereas the reverse situation occurs with larger crustaceans, polychaetes and small fish. The presence of greater amounts of copepods in the diet during the day and of amphipods at night probably reflects the diel activity patterns of the prey.     

Stochastic growth reduces population fluctuations in Daphnia-algal systems

Deterministic, size-structured models are widely used to describe consumer-resource interactions. Such models typically ignore potentially large random variability in juvenile development rates. We present simple representations of this variability and show five approaches to calculating the model parameters for Daphnia pulex interacting with its algal food. Using our parameterized models of growth variability, we investigate the robustness of a recently proposed stabilizing mechanism for Daphnia populations. Growth rate variability increases the range of enrichments over which small-amplitude cycles or quasi-cycles occur, thus increasing the plausibility that the underlying mechanism contributes to the prevalence of small-amplitude cycles in the field and in experiments. More generally, our approach allows us to relate commonly available information on variance of development times to population stability.