Modelling coupled peach tree-aphid population dynamics and their control by winter pruning and nitrogen fertilization

Nitrogen fertilization and winter pruning are commonly used to control crop production in peach [Prunus persica (L.) Batsch] orchards. They are also known to affect the dynamics of Myzus persicae (Sulzer) (Homoptera: Aphididae) aphid populations via bottom-up regulation processes. Interactions between crops and pests can cause complex system behaviour in response to management practices. An integrated approach will therefore improve the understanding of the effects of these two cultural practices on aphid and peach performances.We developed a simulation model that describes the cultural control of interacting peach tree and aphid population dynamics. It uses the principles of common trophic models while gathering available knowledge and explicit assumptions on peach and aphid functioning and the effects of cultural practices.The model was able to qualitatively reproduce the system behaviour observed in the field. It accounted for actions and feedback such as stimulation of foliar growth by winter pruning, consecutive aphid population increase, subsequent damage to foliage, and partial compensatory growth of foliage. The model also reproduced low losses in fruit production due to aphid infestations. However, it called for further integration of ‘long-term’ effects. Analysis of the model showed the complexity of peach tree and aphid responses to leaf N × winter pruning interactions. Simulations indicated that fruit production losses remained low within a range of realistic values of leaf N and pruning intensity, whereas manipulating peach and aphid dynamics, their interactions and their relationships to practices could result in higher losses.The model is useful to evaluate the relevance of cultural practices for a bottom-up regulation of aphid dynamics in crop-pest management. After considering other control methods and fruit quality, it can be used to find a combination of practices that optimises trade-offs between fruit production and environmental conservation goals. A modelling approach that links crop growth and pest population dynamics and integrates management practice effects has strong potential for improving crop-pest management in an integrated crop production context.     

Irruptive population dynamics in Yellowstone pronghorn.

Irruptive population dynamics appear to be widespread in large herbivore populations, but there are few empirical examples from long time series with small measurement error and minimal harvests. We analyzed an 89-year time series of counts and known removals for pronghorn (Antilocapra americana) in Yellowstone National Park of the western United States during 1918-2006 using a suite of density-dependent, density-independent, and irruptive models to determine if the population exhibited irruptive dynamics. Information-theoretic model comparison techniques strongly supported irruptive population dynamics (Leopold model) and density dependence during 1918-1946, with the growth rate slowing after counts exceeded 600 animals. Concerns about sagebrush (Artemisia spp.) degradation led to removals of >1100 pronghorn during 1947-1966, and counts decreased from approximately 700 to 150. The best models for this period (Gompertz, Ricker) suggested that culls replaced intrinsic density-dependent mechanisms. Contrary to expectations, the population did not exhibit enhanced demographic vigor soon after the termination of the harvest program, with counts remaining between 100 and 190 animals during 1967 1981. However, the population irrupted (Caughley model with a one-year lag) to a peak abundance of approximately 600 pronghorn during 1982-1991, with a slowing in growth rate as counts exceeded 500. Numbers crashed to 235 pronghorn during 1992-1995, perhaps because important food resources (e.g., sagebrush) on the winter range were severely diminished by high densities of browsing elk, mule deer, and pronghorn. Pronghorn numbers remained relatively constant during 1996-2006, at a level (196-235) lower than peak abundance, but higher than numbers following the release from culling. The dynamics of this population supported the paradigm that irruption is a fundamental pattern of growth in many populations of large herbivores with high fecundity and delayed density-dependent effects on recruitment when forage and weather conditions become favorable after range expansion or release from harvesting. Incorporating known removals into population models that can describe a wide range of dynamics can greatly improve our interpretation of observed dynamics in intensively managed populations.     

Hidden process models for animal population dynamics.

Hidden process models are a conceptually useful and practical way to simultaneously account for process variation in animal population dynamics and measurement errors in observations and estimates made on the population. Process variation, which can be both demographic and environmental, is modeled by linking a series of stochastic and deterministic subprocesses that characterize processes such as birth, survival, maturation, and movement. Observations of the population can be modeled as functions of true abundance with realistic probability distributions to describe observation or estimation error. Computer-intensive procedures, such as sequential Monte Carlo methods or Markov chain Monte Carlo, condition on the observed data to yield estimates of both the underlying true population abundances and the unknown population dynamics parameters. Formulation and fitting of a hidden process model are demonstrated for Sacramento River winter-run chinook salmon (Oncorhynchus tshawytsha).     

Impact of curve construction and community dynamics on the species-time relationship

The species-time relationship (STR) describes how the species richness of a community increases with the time span over which the community is observed. This temporal scaling provides insight into theoretical questions on species diversity patterns as well as applied questions on the appropriate time scale for biodiversity assessments. To better understand STRs, we discuss the methods used to construct STRs in the literature and derive the impact of curve construction on STR properties. Using vegetation data from Mount St. Helens, Washington, USA, we illustrate the sensitivity of the STR to construction under colonization-dominated dynamics. This study highlights the importance of considering the type of STR when interpreting, comparing, and applying STRs, particularly in disturbed or successional systems.     

Habitat-specific impacts of multiple consumers on plant population dynamics

Multiple consumers often attack seeds, seedlings, and adult plants, but their population-level consequences remain uncertain. We examined how insect and small mammal consumers influenced the demography and abundance of the perennial shrub, bush lupine (Lupinus arboreus). In grassland and dune habitats we established replicate experimental lupine populations in 81-m2 plots that were either protected from, or exposed to, herbivorous voles and granivorous mice (via fencing) and/or root feeding insects (via insecticide treatment). Populations were initiated with transplanted seedlings in 1999 and 2000. We followed the demography of these cohorts, subsequent generations, and the seed bank for 5.5 years. Voles and insects killed many seedlings in dune (1999 only) and grassland (1999 and 2000) habitats. After 2000, insects and voles had minimal effects on seedling or adult survival. Seed predation by granivorous mice, however, greatly depressed seedling recruitment, resulting in lower adult lupine abundance in control plots vs. those protected from rodents. In grasslands, initial effects of voles and insects on seedling survival produced large differences among treatments in adult plant density and the cumulative number of seeds produced throughout the experiment. Differences among grassland populations in seed rain, however, had little influence on the magnitude of seedling recruitment into this habitat. Instead, recruitment out of a preexisting seed bank compensated for the lack of seed production in populations exposed to consumers. Shading by dense adults in plots protected from consumers limited seedling establishment within these populations. Although differences among populations in cumulative seed rain did not influence adult establishment, populations protected from consumers accumulated substantially larger seed banks than controls. These results illustrate how density dependence, habitat-specific seed dynamics, and particular demographic impacts of consumers interact to shape plant population responses to consumers.     

Interactive effects of prey and p,p'-DDE on burrowing owl population dynamics.

We used population models to explore the effects of the organochlorine contaminant p,p'-DDE and fluctuations in vole availability on the population dynamics of Burrowing Owls (Athene cunicularia). Previous work indicated an interaction between low biomass of voles in the diet and moderate levels of p,p'-DDE in Burrowing Owl eggs that led to reproductive impairment. We constructed periodic and stochastic matrix models that incorporated three vole population states observed in the field: average, peak, and crash years. We modeled varying frequencies of vole crash years and a range of impairment of owl demographic rates in vole crash years. Vole availability had a greater impact on owl population growth rate than did reproductive impairment if vole populations peaked and crashed frequently. However, this difference disappeared as the frequency of vole crash years declined to once per decade. Fecundity, the demographic rate most affected by p,p'-DDE, had less impact on population growth rate than adult or juvenile survival. A life table response experiment of time-invariant matrices for average, peak, and crash vole conditions showed that low population growth under vole crash conditions was due to low adult and juvenile survival rates, whereas the extremely high population growth under vole peak conditions was due to increased fecundity. Our results suggest that even simple models can provide useful insights into complex ecological interactions. This is particularly valuable when temporal or spatial scales preclude manipulative experimental work in the field or laboratory.     

Uncertainty analysis of transient population dynamics

Two types of demographic analyses, perturbation analysis and uncertainty analysis, can be conducted to gain insights about matrix population models and guide population management. Perturbation analysis studies how the perturbation of demographic parameters (survival, growth, and reproduction parameters) may affect the population projection, while uncertainty analysis evaluates how much uncertainty there is in population dynamic predictions and where the uncertainty comes from. Previously, both perturbation analysis and uncertainty analysis were conducted on the long-term population growth rate. However, the population may not reach its equilibrium state, especially when there is management by harvesting or hunting. Recently, there has been an increased interest in short-term transient dynamics, which can differ from asymptotic long-term dynamics. There are currently techniques to conduct perturbation analyses of short-term transient dynamics, but no techniques have been proposed for uncertainty analysis of such dynamics. In this study, we introduced an uncertainty analysis technique, the general Fourier Amplitude Sensitivity Test (FAST), to study uncertainties in transient population dynamics. The general FAST is able to identify the amount of uncertainty in transient dynamics and contributions by different demographic parameters. We applied the general FAST to a mountain goat (Oreamnos americanus) matrix population model to give a clear illustration of how uncertainty analysis can be conducted for transient dynamics arising from matrix population models.     

Recolonizing wolves and mesopredator suppression of coyotes: impacts on pronghorn population dynamics.

Food web theory predicts that the loss of large carnivores may contribute to elevated predation rates and, hence, declining prey populations, through the process of mesopredator release. However, opportunities to test predictions of the mesopredator release hypothesis are rare, and the extent to which changes in predation rates influence prey population dynamics may not be clear due to a lack of demographic information on the prey population of interest. We utilized spatial and seasonal heterogeneity in wolf distribution and abundance to evaluate whether mesopredator release of coyotes (Canis latrans), resulting from the extirpation of wolves (Canis lupus) throughout much of the United States, contributes to high rates of neonatal mortality in ungulates. To test this hypothesis, we contrasted causes of mortality and survival rates of pronghorn (Antilocapra americana) neonates captured at wolf-free and wolf-abundant sites in western Wyoming, USA, between 2002 and 2004. We then used these data to parameterize stochastic population models to heuristically assess the impact of wolves on pronghorn population dynamics due to changes in neonatal survival. Coyote predation was the primary cause of mortality at all sites, but mortality due to coyotes was 34% lower in areas utilized by wolves (P < 0.001). Based on simulation modeling, the realized population growth rate was 0.92 based on fawn survival in the absence of wolves, and 1.06 at sites utilized by wolves. Thus, wolf restoration is predicted to shift the trajectory of the pronghorn population from a declining to an increasing trend. Our results suggest that reintroductions of large carnivores may influence biodiversity through effects on prey populations mediated by mesopredator suppression. In addition, our approach, which combines empirical data on the population of interest with information from other data sources, demonstrates the utility of using simulation modeling to more fully evaluate ecological theories by moving beyond estimating changes in vital rates to analyses of population-level impacts.     

Interacting trophic forcing and the population dynamics of herring

Small pelagic fish occupy a central position in marine ecosystems worldwide, largely by determining the energy transfer from lower trophic levels to predators at the top of the food web, including humans. Population dynamics of small pelagic fish may therefore be regulated neither strictly bottom-up nor top-down, but rather through multiple external and internal drivers. While in many studies single drivers have been identified, potential synergies of multiple factors, as well as their relative importance in regulating population dynamics of small pelagic fish, is a largely unresolved issue. Using a statistical, age-structured modeling approach, we demonstrate the relative importance and influence of bottom-up (e.g., climate, zooplankton availability) and top-down (i.e., fishing and predation) factors on the population dynamics of Bothnian Sea herring (Clupea harengus) throughout its life cycle. Our results indicate significant bottom-up effects of zooplankton and interspecific competition from sprat (Sprattus sprattus), particularly on younger age classes of herring. Although top-down forcing through fishing and predation by grey seals (Halichoerus grypus) and Atlantic cod (Gadus morhua) also was evident, these factors were less important than resource availability and interspecific competition. Understanding key ecological processes and interactions is fundamental to ecosystem-based management practices necessary to promote sustainable exploitation of small pelagic fish.     

Laboratory growth and metabolism of the winter flounder Pseudopleuronectes americanus from hatching through metamorphosis at three temperatures

Growth and metabolism of the winter flounder Pseudopleuronectes americanus were studied in the laboratory at 2°, 5° and 8°C. Dry-weight determinations of growth demonstrated significant direct regressions of growth on temperature. Mean, daily specific growth rates were 10.1%/day at 80°C, 5.8%/day at 50°C, and 2.6%/day at 2°C. Time to metamorphosis was 49 days at 8°C and 80 days at 5°C. Larvae did not survive to metamorphosis at 2°C. Absolute values of routine metabolism expressed in μl of oxygen consumed regressed on body weight were best described by a third-degree polynomial. Larval routine metabolism increased from hatching to metamorphosis, at which time it declined before again increasing. Temperature directly affected routine metabolism. Metabolism on a unit-weight basis decreased with increasing size and was also directly influenced by temperature.     

Energy flow and population dynamics of the barnacle Balanus glandula

The energetics and population dynamics of a barnacle (Balanus glandula Darwin) population in British Columbia, Canada, were studied. Consumption, energy flow, production and mortality were 6844.6, 6667.0, 2896.5 and 2522.8 Kcal m^-2 year^-1, respectively. These energy flow and production values are among the highest for animal populations reported, and therefore strongly suggest the functional importance of E. glandula in littoral systems. The young age groups of the first-year settlements were most important in contributing to the energy flow, production and reproduction of the entire population. Most of the assimilated energy in the older age groups was used in respiration.     

Complex dynamics of the population with a simple age structure

The effects of the following modes of density-dependent control of population growth: density-dependent birth rate, adult survival rate, juvenile survival rate are compared based on the mathematical model of population dynamics. It is shown that the most efficient mechanisms limiting population size are decreasing with the growth of the adult population birth rate and/or the decreasing survival rate of the offspring with the increase in their number. However, these same mechanisms are responsible for oscillations of the population size and its chaotic change. The density-dependence of the adult survival rate is not efficient in constraining the population growth, but it can substantially limit the magnitude of oscillations of the population size.     

Biology and population dynamics of the intertidal isopod Cirolana harfordi

The distribution of Cirolana harfordi (Lockington) populations is determined largely by the availability of loose boulders on sandy beaches. The isopods swim out from under the rocks at high tide to feed. Their diet consists primarily of minute polychaetes and crustaceans. In addition, the isopods locate and utilize any available dead animal matter in the surf zone. Breeding occurs throughout most of the year, except for a brief lull in the fall. Females produce 1 or 2 broods of 18 to 68 young during their 2-year life-span. Marsupial incubation lasts 3 to 4 months. The population size and structure remained relatively constant from one year to the next. Mortality rates estimated for juveniles, males and females showed that newly emergent young and post-reproductive females suffer the greatest losses (up to 75% mortality per month).     

Non-linear biological responses to disturbance: consequences on population dynamics

We assessed how non-linear biological responses to environmental noise, or “noise filtering”, impact the spectra of density-dependent population dynamics, and the correlation between noise and population dynamics. The noise was assumed to affect population growth rate in a discrete-time population model by Hassell [J. Anim. Ecol. 44 (1975) 283–295] where the population growth rate was linked to the environment with an optimum type filter. When compared to unfiltered noise, the filtered noise can distort the stationary distribution of population values. The optimum type filter can make cyclic population dynamics more regular and low population values can become more frequent or rare depending on the strength of density dependence. Filtering can cause blue shifted and red shifted population dynamics and determine the strength of correlation between environmental noise and population size. In most cases, optimum type filtering makes linear correlation between population dynamics and noise weaker. The filter effect on population spectra and noise versus population correlation is sensitive to changes in population model parameters, the location where noise hits the filter, and noise colour.     

Composite forces shape population dynamics of copepod crustaceans

Understanding the processes that control species abundance and distribution is a major challenge in ecology, yet for a large number of potentially important organisms, we know little about the biotic and abiotic factors that influence population size. One group of aquatic organisms that defies traditional demographic analyses is the Crustacea, particularly those with complex life cycles. We used likelihood techniques and information theoretics to evaluate a suite of models representing alternative hypotheses on factors controlling the abundance of two copepod crustaceans in a small, tropical floodplain lake. Quantitative zooplankton samples were collected at three stations in a Venezuelan floodplain lake from June through December 1984; the average sampling interval was two days. We constructed a series of models with stage structure that incorporated six biotic and abiotic covariates in various combinations to account for temporal changes in abundance of these target species and in their population growth rates. Our analysis produced several novel insights into copepod population dynamics. We found that multiple forces affected the abundance of particular stages, that these factors differed between species as well as among stages within each species, and that biotic processes had the largest effects on copepod population dynamics. Density dependence had a large effect on the survival of Oithona amazonica copepodites and on population growth rate of Diaptomus negrensis.     

Viability of embryos of the winter flounder Pseudopleuronectes americanus exposed to combinations of cadmium and salinity at selected temperatures

Developing eggs of the winter flounder Pseudopleuronectes americanus (Walbaum) were exposed to 9 combinations of cadmium and salinity at 5° and 10°C. Overall mean times to 50% hatch ranged from 7.7 days at 10°C to 17.9 days at 5°C. Mean percentages of total hatches ranged from 50 to 100% for all treatment combinations. Response-surface analyses indicate percentages of viable hatches were generally lowest at 10% S and highest in the 25 to 30% S range. Regression analyses of viable hatch data show that at both temperatures cadmium significantly influenced viable hatch in all experiments. Viable hatch was also significantly influenced by the linear and quadratic effect of salinity in both tests at 5°C and in 2 of 3 tests at 10°C. The interaction between cadmium and salinity also significantly affected viable hatch at 10°C.     

Impact of cannibalism on predator-prey dynamics: size-structured interactions and apparent mutualism

Direct and indirect interactions between two prey species can strongly alter the dynamics of predator-prey systems. Most predators are cannibalistic, and as a consequence, even systems with only one predator and one prey include two prey types: conspecifics and heterospecifics. The effects of the complex direct and indirect interactions that emerge in such cannibalistic systems are still poorly understood. This study examined how the indirect interaction between conspecific and heterospecific prey affects cannibalism and predation rates and how the direct interactions between both species indirectly alter the effect of the cannibalistic predator. I tested for these effects using larvae of the stream salamanders Eurycea cirrigera (prey) and Pseudotriton ruber (cannibalistic predator) by manipulating the relative densities of the conspecific and heterospecific prey in the presence and absence of the predator in experimental streams. The rates of cannibalism and heterospecific predation were proportional to the respective densities and negatively correlated, indicating a positive indirect interaction between conspecific and heterospecific prey, similar to "apparent mutualism." Direct interactions between prey species did not alter the effect of the predator. Although both types of prey showed a similar 30% reduction in night activity and switch in microhabitat use in response to the presence of the predator, cannibalism rates were three times higher than heterospecific predation rates irrespective of the relative densities of the two types of prey. Cumulative predation risks differed even more due to the 48% lower growth rate of conspecific prey. Detailed laboratory experiments suggest that the 3:1 difference in cannibalism and predation rate was due to the higher efficiency of heterospecific prey in escaping immediate attacks. However, no difference was observed when the predator was a closely related salamander species, Gyrinophilus porphyriticus, indicating that this difference is species specific. This demonstrates that cannibalism can result in the coupling of predator and prey mortality rates that strongly determines the dynamics of predator-prey systems.     

Reproductive energetics,endocrinology and population dynamics of Nephtys caeca and N. hombergi

An investigation of gametogenetic cycles and spawning success in populations of Nephtys caeca and N. hombergi from the River Tyne estuary, North Shields, England, in relation to production of gonadotrophic (GH) and spawning hormones (SH) has been made, based on data collected from 1978–1984. The data show that a consistent pattern of gametogenesis and efficient spawning occurs in populations of N. caeca in north-east England, but that the reproductive cycle of N. hombergi is erratic. Premature oosorption and gametogenic failure in this latter species is associated with a low level of GH production. Following high levels of GH production, gravid females sometimes fail to spawn owing to the non-release of SH. The reproductive energetics and reproductive success of N. caeca and N. hombergi are described in relation to the observed patterns of endocrine activity and the consequent changes in population structure.     

Simulating the influences of various fire regimes on caribou winter habitat.

Caribou are an integral component of high-latitude ecosystems and represent a major subsistence food source for many northern people. The availability and quality of winter habitat is critical to sustain these caribou populations. Caribou commonly use older spruce woodlands with adequate terrestrial lichen, a preferred winter forage, in the understory. Changes in climate and fire regime pose a significant threat to the long-term sustainability of this important winter habitat. Computer simulations performed with a spatially explicit vegetation succession model (ALFRESCO) indicate that changes in the frequency and extent of fire in interior Alaska may substantially impact the abundance and quality of winter habitat for caribou. We modeled four different fire scenarios and tracked the frequency, extent, and spatial distribution of the simulated fires and associated changes to vegetation composition and distribution. Our results suggest that shorter fire frequencies (i.e., less time between recurring fires) on the winter range of the Nelchina caribou herd in eastern interior Alaska will result in large decreases of available winter habitat, relative to that currently available, in both the short and long term. A 30% shortening of the fire frequency resulted in a 3.5-fold increase in the area burned annually and an associated 41% decrease in the amount of spruce-lichen forest found on the landscape. More importantly, simulations with more frequent fires produced a relatively immature forest age structure, compared to that which currently exists, with few stands older than 100 years. This age structure is at the lower limits of stand age classes preferred by caribou from the Nelchina herd. Projected changes in fire regime due to climate warming and/or additional prescribed burning could substantially alter the winter habitat of caribou in interior Alaska and lead to changes in winter range use and/or population dynamics.     

Importance of life cycle events in the population dynamics of Gonyaulax tamarensis

Life cycle changes that allow populations of the toxic dinoflagellate Gonyaulax tamarensis Lebour to inhabit the benthos and the plankton alternately are important factors regulationg the initiation and decline of blooms in restricted embavments. When the dynamics of these estuarine populations were monitored during “bloom” and “non bloom” years, it was shown that: (1) each year, germination of benthie cysts inoculated the overlying waters during the vernal warming period, but a large residual population remained in the sediments throughout the blooms; (2) the resulting planktonic population began growth under suboptimal temperature conditions; (3) the populations developed from this inoculum through asexual reproduction until sexuality (and cyst formation) were induced; (4) encystment was not linked to any obvious environmental cue and occurred under apparently optimal conditions; and (5) an increase in the number of non-mitotic swimming cells (planozygotes, the precursors to dormant cysts) accompanied the rapid decline of the planktonic population. Thus encystment, in combination with hypothesized losses due to advection and grazing, contributed substantiatly to the decline of the vegetative cell population. We conclude that the encystment/excystment cycle temporally restricts the occurrence of the vegetative population and may not be optimized for rapid or sustained vegetative growth and bloom formation in shallow embayments. The factors that distinguish “bloom” from “non-bloom” years thus appear to be operating on the growth of the planktonic population.