Tree adaptive growth (TAG) model: a life-history theory-based analytical model for post-thinning forest stand dynamics.

Background: Understanding stand dynamics is essential for predicting future wood supply and associated ecosystem services for sustainable forest management. The dynamics of natural stands can be characterized by age-dependent growth and yield models. However, dynamics in managed stands appear somewhat different from that of natural stands, especially with difficulties in explaining the phenomenon of post-thinning overcompensation, based upon some long-term observations. Though overcompensation is an ideal outcome for the forest sector, it had been largely treated as an outlier and thus ignored or dismissed as "out-of-the-ordinary". Methodology: We developed a life history theory-based, state-dependent model of Tree Adaptive Growth (TAG) to investigate this phenomenon and verified that overcompensation should be a common outcome in post-thinning forest stands when the stand growth over time is sigmoid shaped. TAG posits that individual trees will invest proportionately more into growth following thinning because it is evolutionarily adaptive to do so. Results: Our investigation of the model's behavior unearthed diverse stand growth patterns similar to that which is observed in the empirical datasets and predicted by a statistics-based Tree's Compensatory Growth (TreeCG) model. Conclusion: A simple, theory-driven, analytical model, TAG, can reproduce the diverse growth patterns in post-thinning stands and thus assist addressing silviculture-related issues. The model can be applied to various jurisdictions even without detailed regional growth and yield relationships and is capable of incorporating the effects of other time sensitive factors like fertilization, pruning, and climate change.

Modelling the stand dynamics after a thinning induced partial mortality: A compensatory growth perspective.

Introduction: With increasing forest areas under management, dynamics of managed stands have gained more attention by forest managers and practitioners. Improved understanding on how trees and forest stands would respond to different disturbances is required to predict the dynamics of managed stand.s. Partial mortality commonly occurs in stand development, and different response patterns of trees and stands to partial mortality would govern stand dynamics. Methods: To investigate the possible response patterns using existing knowledge of growth and yield relationships, we developed TreeCG model, standing for Tree's Compensatory Growth, a state-dependent individual tree-based forest growth model that simulates the compensatory growth of trees after experiencing a partial mortality. The mechanism behind the simulation is the redistribution of resources, including nutrients and space, freed from died trees to surviving trees. The developed new algorithm simplified the simulations of annual growth increments of individual trees over a long period of stand development. Results: The model was able to reproduce the forest growth patterns displayed in long-term precommercial thinning experiments. The simulated forest growth displayed the process of compensatory growth from under compensation, to compensation-induced-equality, and to overcompensation over time. Discussion: Our model can simulate stand growth trajectories after different partial harvest regimes at different times and intensities, thus support decisions in best partial harvest strategies. This generic model can be refined with given tree species and specific site conditions to predict stand dynamics after given partial mortality for any jurisdictions under management. The simulation reassembles growth trajectories of natural stands when no thinning is conducted.

Detecting Compensatory Growth in Silviculture Trials: Empirical Evidence From Three Case Studies Across Canada.

Compensatory growth (CG) appears common in biology and is defined as accelerated growth after experiencing a period of unfavorable conditions. It usually leads to an increase in biomass that may eventually equal or even surpass that of sites not experiencing disturbance. In forestry, with sufficient time the stand volume lost in a disturbance such as a thinning operation could match or even exceed those from undisturbed sites, respectively called exact and overcompensation. The forest sector could benefit from enhanced productivity and associated ecosystem services such as carbon storage through overcompensation. Therefore, detection of CG in different types of forests becomes important for taking advantage of it in forest management. However, compensatory growth has not been reported widely in forestry, partially due to the paucity of long-term observations and lack of proper indicators. Legacy forest projects can provide a suitable data source, though they may be originally designed for other purposes. Three case studies representing different data structures of silviculture trials are investigated to evaluate if compensatory growth is common in forest stands. Our results showed that compensatory growth occurred in all three cases, and thus suggested that the compensatory growth might indeed be common in forest stands. We found that the relative growth (RG) can serve as a universal indicator to examine stand-level compensatory growth in historical long-term silviculture datasets. When individual tree-based measurements are available, both volume and value-based indicators can be used in detecting compensatory growth, and lumber value-based indicators could be more sensitive in detecting overcompensation.

Ecology and Prediction of Compensatory Growth: From Theory to Application in Forestry.

Compensatory growth has been observed in forests, and it also appears as a common phenomenon in biology. Though it sometimes takes different names, the essential meanings are the same, describing the accelerated growth of organisms when recovering from a period of unfavorable conditions such as tissue damage at the individual level and partial mortality at the population level. Diverse patterns of compensatory growth have been reported in the literature, ranging from under-, to compensation-induced-equality, and to over-compensation. In this review and synthesis, we provide examples of analogous compensatory growth from different fields, clarify different meanings of it, summarize its current understanding and modeling efforts, and argue that it is possible to develop a state-dependent model under the conceptual framework of compensatory growth, aimed at explaining and predicting diverse observations according to different disturbances and environmental conditions. When properly applied, compensatory growth can benefit different industries and human society in various forms.

Forest Productivity Enhancement and Compensatory Growth: A Review and Synthesis.

This review and synthesis article attempts to integrate observations from forestry to contemporary development in related biological research fields to explore the issue of forest productivity enhancement and its contributions in mitigating the wood supply shortage now facing the forest sector. Compensatory growth has been clearly demonstrated in the long-term precommercial thinning and fertilization trial near the Shawnigan Lake, British Columbia, Canada. This phenomenon appears similar to many observations from other biological fields. The concept of compensatory growth can be applied to forest productivity enhancement through overcompensation, by taking advantage of theories and methods developed in other compensatory growth research. Modeling technology provides an alternative approach in elucidating the mechanisms of overcompensation, which could reveal whether the Shawnigan Lake case could be generalized to other tree species and regions. A new mitigation strategy for dealing with issues related to wood supply shortage could be formed through searching for and creating conditions promoting overcompensation. A forest growth model that is state dependent could provide a way of investigating the effect of partial harvest on forest growth trajectories and stand dynamics. Results from such a study could provide cost-effective decision support tools to practitioners.

State-dependent domicile leaving rates in Anopheles gambiae.

BACKGROUND: Transmission of Plasmodium greatly depends on the foraging behaviour of its mosquito vector (Anopheles spp.). The accessibility of blood hosts and availability of plant sugar (i.e., nectar) sources, together with mosquito energy state, have been shown to modulate blood feeding (and thus biting rates) of anopheline mosquitoes. In this study, the influence of mosquito starvation status and availability of nectar on the decision of female Anopheles gambiae mosquitoes to leave a bed net-protected blood host was examined. METHODS: Two small-scale mesocosm experiments were conducted using female mosquitoes starved for 0, 24 or 48 h, that were released inside a specially constructed hut with mesh-sealed exits and containing a bed net-protected human volunteer. Floral cues were positioned on one side of the hut or the other. Several biologically plausible exponential decay models were developed that characterized the emigration rates of mosquitoes from the huts. These varied from simple random loss to leaving rates dependent upon energy state and time. These model fits were evaluated by examining their fitted parameter estimates and comparing Akaike information criterion. RESULTS: Starved mosquitoes left domiciles at a higher rate than recently fed individuals however, there was no difference between 1- and 2-day-starved mosquitoes. There was also no effect of floral cue placement. The best fitting emigration model was one based on both mosquito energy state and time whereas the worst fitting model was one based on the assumption of constant leaving rates, independent of time and energy state. CONCLUSIONS: The results confirm that mosquito-leaving behaviour is energy-state dependent, and provide some of the first evidence of state-dependent domicile emigration in An. gambiae, which may play a role in malarial transmission dynamics. Employment of simple, first-principle, mechanistic models can be very useful to our understanding of why and how mosquitoes leave domiciles.

Impact of male alternative reproductive tactics on female costs of sexual conflict under variation in operational sex ratio and population density.

Sexual conflict over mating rate is both pervasive and evolutionarily costly. For females, the lifetime reproductive fitness costs that arise through interactions with potential mates will be influenced by the frequency of such interactions, and the fitness cost of each interaction. Both of these factors are likely to be influenced by variation in operational sex ratio (OSR) and population density. Variation in OSR- and density-dependent male alternative reproductive tactics (ARTs) may be particularly important if the fitness costs that females experience vary with the reproductive tactics that males express. Using a simple model, we consider several examples of OSR- and/or density-dependent variation in male ARTs and the frequency of male-female interactions, and find that variation in the expression of male ARTs has the potential to augment or diminish the costs of frequent male interactions for females. Accurately documenting variation in the expression of male ARTs and associated female fitness costs will benefit future work in this area.

State dependence, personality, and plants: light-foraging decisions in Mimosa pudica (L.).

Plants make foraging decisions that are dependent on ecological conditions, such as resource availability and distribution. Despite the field of plant behavioral ecology gaining momentum, ecologists still know little about what factors impact plant behavior, especially light-foraging behavior. We made use of the behavioral reaction norm approach to investigate light foraging in a plant species that exhibits rapid movement: Mimosa pudica. We explored how herbivore avoidance behavior in M. pudica (which closes its leaflets temporarily when disturbed) is affected by an individual's energy state and the quality of the current environment and also repeatedly tested the behavior of individuals from two seed sources to determine whether individuals exhibit a "personality" (i.e., behavioral syndrome). We found that when individuals are in a low-energy state, they adopt a riskier light-foraging strategy, opening leaflets faster, and not closing leaflets as often in response to a disturbance. However, when plants are in a high-energy state, they exhibit a plastic light-foraging strategy dependent on environment quality. Although we found no evidence that individuals exhibit behavioral syndromes, we found that individuals from different seed sources consistently behave differently from each other. Our results suggest that plants are capable of making state-dependent decisions and that plant decision making is complex, depending on the interplay between internal and external factors.

Making the best of a bad situation: host partial resistance and bypass of behavioral manipulation by parasites?

With few exceptions, parasitic manipulation dramatically reduces host fitness. That said, evidence of host resistance to behavior-manipulating parasites is scarce. Here, we suggest that the evolution of partial resistance, as well as bypass, to manipulation (PRM and BPM, respectively) represents new, seldom-explored options for parasitized hosts. Natural selection could favor hosts that partially resist certain manipulative dimensions to postpone their death and perform additional reproductive episodes (PRM). Alternatively, manipulated hosts may express novel traits that do not alter the manipulation per se but that alleviate its detrimental fitness consequences (BPM). If effective, PRM and BPM have many implications for the ecology and evolution of hosts and their parasites, especially the evolution of multidimensional manipulations.

On the evolution of omnivory in a community context.

Omnivory is extremely common in animals, yet theory predicts that when given a choice of resources specialization should be favored over being generalist. The evolution of a feeding phenotype involves complex interactions with many factors other than resource choice alone, including environmental heterogeneity, resource quality, availability, and interactions with other organisms. We applied an evolutionary simulation model to examine how ecological conditions shape evolution of feeding phenotypes (e.g., omnivory), by varying the quality and availability (absolute and relative) of plant and animal (prey) resources. Resulting feeding phenotypes were defined by the relative contribution of plants and prey to diets of individuals. We characterized organisms using seven traits that were allowed to evolve freely in different simulated environments, and we asked which traits are important for different feeding phenotypes to evolve among interacting organisms. Carnivores, herbivores, and omnivores all coexisted without any requirement in the model for a synergistic effect of eating plant and animal prey. Omnivores were most prevalent when ratio of plants and animal prey was low, and to a lesser degree, when habitat productivity was high. A key result of the model is that omnivores evolved through many different combinations of trait values and environmental contexts. Specific combinations of traits tended to form emergent trait complexes, and under certain environmental conditions, are expressed as omnivorous feeding phenotypes. The results indicate that relative availabilities of plants and prey (over the quality of resources) determine an individual's feeding class and that feeding phenotypes are often the product of convergent evolution of emergent trait complexes under specific environmental conditions. Foraging outcomes appear to be consequences of degree and type of phenotypic specialization for plant and animal prey, navigation and exploitation of the habitat, reproduction, and interactions with other individuals in a heterogeneous environment. Omnivory should not be treated as a fixed strategy, but instead a pattern of phenotypic expression, emerging from diverse genetic sources and coevolving across a range of ecological contexts.

Effects of larval density and feeding rates on larval life history traits in Anopheles gambiae s.s. (Diptera: Culicidae).

Mosquito larval habitat determines the fitness, survivorship, fecundity, and vector capacity of emerging adults. We manipulated larval density and food provisioning in the laboratory to determine their effects on a number of life history parameters of Anopheles gambiae. Larval mortality was positively correlated with larval density. In addition, increased density skewed sex ratios that favored females.

Effects of food, water depth, and temperature on diving activity of larval Anopheles gambiae sensu stricto: evidence for diving to forage.

Anopheles gambiae larvae have frequently been observed to dive, but the ecology of this behavior has not been extensively examined. We manipulated food level, water depth, and temperature for individually-reared larvae and observed diving activity. Larvae dived more often under low food, which suggests that they dive to forage. There was only weak evidence for effects of water depth or temperature on diving. Experimental results are discussed in the context of energy budgets. Understanding larval ecology of this species is important for predicting how it will respond to environmental change. Further study is needed to assess the role that larval diving plays in both feeding ecology and thermal regulation of this and other medically important species.

Phenology of Dasineura oxycoccana (Diptera: Cecidomyiidae) on cranberry and blueberry indicates potential for gene flow.

Dasineura oxycoccana (Johnson) (Diptera: Cecidomyiidae) is a pest of cranberry, Vaccinium macrocarpon (Aiton) (Ericales: Ericaceae), and highbush blueberry, Vaccinium corymbosum (L.) (Ericales: Ericaceae), in North America. In British Columbia, Canada, D. oxycoccana was first found on highbush blueberry in 1991 and then on cranberry seven years later. Because many cranberry and highbush blueberry farms are adjacent to one another, we hypothesized that D. oxycoccana was moving from highbush blueberry onto cranberry. Cranberry and highbush blueberry differ in phenology, and adaptation to these different phenologies may result in host races or cryptic species on these two crops. We recognized the alternative hypothesis that D. oxycoccana had arrived as immature stages with cranberry vines imported from another region of North America. During spring and summer, we recorded the phenology of D. oxycoccana and the development of plant shoots from three cranberry and three highbush blueberry farms to determine whether the opportunity exists for successful movement of D. oxycoccana between the two crops. Our results show that D. oxycoccana from cranberry and highbush blueberry overlap in phenology for much of the season, indicating a high potential for movement and gene flow. However, differences were seen in number of larvae per shoot, location of pupae, and heat unit accumulation during larval development suggesting that instead there may be the potential for host race or cryptic species formation.

Energy-state dependent responses of Anopheles gambiae (Diptera: Culicidae) to simulated bednet-protected hosts.

In nature, Anopheles gambiae mosquitoes are found at various energy levels and such females must choose between seeking somatic energy from sugar sources and obtaining both somatic and gametic energy from blood hosts. We used a straight-tube olfactometer containing a simulated unobtainable blood host (human foot smell protected by a net) as well as a sugar source (honey odor). We assessed female probing rate and residence time at the net as a function of energy state (0, 24, 48, 72-h starved). In our trials, 0-h starved females showed low response to human odor, low probing rate, and residence time at the human odor site. By contrast, both 48 and 72-h individuals showed high response to foot odor, longer residence time, and higher probing rates. Seventy-two-h females also flew towards the honey source less often than other groups. Our findings suggest that managing sugar sources might be a viable strategy for influencing mosquito biting behavior.

Predator identity and the nature and strength of food web interactions.

1. Most trophic interaction theory assumes that all predators are an abstract form of risk to which prey respond in a quantitatively similar manner. This conceptualization can be problematic because recent empirical work demonstrates that variation in the responses of prey to different predators can play a key role in structuring communities and regulating ecosystem function. 2. Predator identity - the species specific response of prey to a predator - has been proposed as an ultimate mechanism driving the relative contribution of indirect effects in food webs; however few studies have explicitly tested this hypothesis. 3. This study explores the impact of predator identity on direct consumptive (CE) and non-consumptive effects (NCEs), and on the relative contribution of indirect, density and trait-mediated effects in trophic cascades within host-parasitoid communities. 4. We systematically compared the individual, host-parasitoid-plant interactions of two actively foraging parasitoid species with disparate foraging styles, one aggressive and one furtive, a common aphid host and plant. Our results demonstrate that the degree of risk aversion by prey to each particular predator species (i.e. predator identity) is a key factor driving the nature and strength of direct and indirect transmission pathways. 5. Both parasitoid species, in general, had a negative impact on plants. The magnitude of the aphid anti-predator dispersal response was positively correlated with plant infestation and plant damage. The qualitative effect of predator-induced infestation of new plants superseded the quantitative effects of predator-mediated reductions in aphid numbers. 6. The greatest indirect impact on plants was generated by the aggressively foraging parasitoid, and the strength of the aphids anti-predator response (a NCE) antagonistically traded-off with CEs due to an increased investment in attempting to capture risk-sensitized prey. In contrast, the furtive parasitoid did not elicit a strong anti-predator response, had little indirect impact on plants, but generated very high CEs due to the advantage of ovipositing into a sedentary prey population. 7. Our data suggest the responses of prey to different predatory cues may be an important mechanism driving the relative contribution of transmission pathways in trophic cascades. We conclude that predator identity is a key factor influencing the nature and strength of food web interactions.

Host-adapted parasitoids in biological control: does source matter?

It has been hypothesized that the success of a biological control introduction is, in part, dependent on the ability of the control agent to become established in its new environment or to its new population of hosts through local adaptation. Despite this, few studies have investigated the influence of the recent coevolutionary history of pest species and natural enemies on the efficacy of biological control agents, especially for agents that are mass-reared for release in agriculture. We investigate the evolutionary potential of a biological control agent Aphidius ervi to adapt to a key pest species, the foxglove aphid Aulacorthum solani, through components essential to the evolution of parasitoid virulence. We explored (1) the influence of genetic variation from natural source populations on the ability to parasitize natal and non-natal host species; (2) the heritability of key traits related to parasitoid fitness; and (3) the efficacy of parasitoid host-selection lines in a greenhouse system. Source populations maintained genetic variation in the ability to utilize natal and non-natal host species; however, only some of the traits sampled suggested local adaptation of parasitoid populations. The ability to parasitize a host was found to be genetically determined and strongly heritable, irrespective of host species. The greenhouse study demonstrated the potential of parasitoid selection lines to substantially increase performance of parasitoids for target pest species. This research provides insight into novel techniques that can be used to increase the quality of biological control agents through the development of lines of natural enemies adapted to particular pest species.

Size-mediated adaptive foraging: a host-selection strategy for insect parasitoids.

Foraging models are useful tools for generating predictions on predator-prey interactions, such as habitat or diet choice. However, the majority of studies attempting to explain adaptive behaviour using optimality criteria have assumed that there is no trait (e.g. size) variation among individual consumers or their prey. Hymenopteran parasitoids that attack the free-living stages of their host are an ideal system for studying the influence of body size on host selection because of the wide range of adult parasitoid sizes coupled with the defensive capabilities of their hosts. We report here our application of an experimentally parameterized host selection model to investigate the influence of parasitoid body size on the range of acceptable host instar classes. Using a demographic model, we compared the efficiency of parasitoids using an optimal host selection strategy against parasitoids using an indiscriminate host selection strategy over a range of different parasitoid body sizes. Net fitness accrual of parasitoids and the impact of host instar selection on aphid recruitment were assessed on different stage-structured aphid populations. Our results demonstrate that optimal host selection allows larger parasitoids to utilize a wider range of hosts. However, smaller parasitoids receive the greatest benefits from selecting hosts optimally by utilizing a restricted range of small, poorly defended hosts when they are abundant. We argue that the correlation between flexible host selection behaviour and adult body size may be a general phenomenon that applies to the majority of hymenopteran parasitoids that attack free-living, well-defended hosts. The potential of within-generation behavioural interactions to impact between-generation dynamics in host-parasitoid populations are discussed.

Host-range evolution in Aphidius parasitoids: fidelity, virulence and fitness trade-offs on an ancestral host.

The diversity of parasitic insects remains one of the most conspicuous patterns on the planet. The principal factor thought to contribute to differentiation of populations and ultimately speciation is the intimate relationship parasites share with hosts and the potential for disruptive selection associated with using different host species. Traits that generate this diversity have been an intensely debated topic of central importance to the evolution of specialization and maintenance of ecological diversity. A fundamental hypothesis surrounding the evolution of specialization is that no single genotype is uniformly superior in all environments. This "trade-off" hypothesis suggests that negative fitness correlations can lead to specialization on different hosts as alternative stable strategies. In this study we demonstrate a trade-off in the ability of the parasitoid, Aphidius ervi, to maintain a high level of fitness on an ancestral and novel host, which suggests a genetic basis for host utilization that may limit host-range expansion in parasitoids. Furthermore, behavioral evidence suggests mechanisms that could promote specialization through induced host fidelity. Results are discussed in the context of host-affiliated ecological selection as a potential source driving diversification in parasitoid communities and the influence of host species heterogeneity on population differentiation and local adaptation.