Predicting the fundamental thermal niche of ectotherms.

Climate warming is predicted to increase mean temperatures and thermal extremes on a global scale. Because their body temperature depends on the environmental temperature, ectotherms bear the full brunt of climate warming. Predicting the impact of climate warming on ectotherm diversity and distributions requires a framework that can translate temperature effects on ectotherm life-history traits into population- and community-level outcomes. Here we present a mechanistic theoretical framework that can predict the fundamental thermal niche and climate envelope of ectotherm species based on how temperature affects the underlying life-history traits. The advantage of this framework is twofold. First, it can translate temperature effects on the phenotypic traits of individual organisms to population-level patterns observed in nature. Second, it can predict thermal niches and climate envelopes based solely on trait response data and, hence, completely independently of any population-level information. We find that the temperature at which the intrinsic growth rate is maximized exceeds the temperature at which abundance is maximized under density-dependent growth. As a result, the temperature at which a species will increase the fastest when rare is lower than the temperature at which it will recover from a perturbation the fastest when abundant. We test model predictions using data from a naturalized-invasive interaction to identify the temperatures at which the invasive can most easily invade the naturalized's habitat and the naturalized is most likely to resist the invasive. The framework is sufficiently mechanistic to yield reliable predictions for individual species and sufficiently broad to apply across a range of ectothermic taxa. This ability to predict the thermal niche before a species encounters a new thermal environment is essential to mitigating some of the major effects of climate change on ectotherm populations around the globe.

Temperature-dependent dispersal and ectotherm species' distributions in a warming world.

Dispersal is a crucial component of species' responses to climate warming. Warming-induced changes in species' distributions are the outcome of how temperature affects dispersal at the individual level. Yet, there is little or no theory that considers the temperature dependence of dispersal when investigating the impacts of warming on species' distributions. Here I take a first step towards filling this key gap in our knowledge. I focus on ectotherms, species whose body temperature depends on the environmental temperature, not least because they constitute the majority of biodiversity on the planet. I develop a mathematical model of spatial population dynamics that explicitly incorporates mechanistic descriptions of ectotherm life history trait responses to temperature. A novel feature of this framework is the explicit temperature dependence of all phases of dispersal: emigration, transfer and settlement. I report three key findings. First, dispersal, regardless of whether it is random or temperature-dependent, allows both tropical and temperate ectotherms to track warming-induced changes in their thermal environments and to expand their distributions beyond the lower and upper thermal limits of their respective climate envelopes. In the absence of dispersal mortality, warming does not alter these new distributional limits. Second, an analysis based solely on trait response data predicts that tropical ectotherms should be able to expand their distributions polewards to a greater degree than temperate ectotherms. Analysis of the dynamical model confirms this prediction. Tropical ectotherms have an advantage when moving to cooler climates because they experience lower within-patch and dispersal mortality, and their higher thermal optima and maximal birth rates allow them to take advantage of the warmer parts of the year. Previous theory has shown that tropical ectotherms are more successful in invading and adapting the temperate climates than vice versa. This study provides the key missing piece, by showing how temperature-dependent dispersal could facilitate both invasion and adaptation. Third, dispersal mortality does not affect the poleward expansion of ectotherm distributions. But, it prevents both tropical and temperate ectotherms from maintaining sink populations in localities that are too warm to be viable in the absence of dispersal. Dispersal mortality also affects species' abundance patterns, causing a larger decline in abundance throughout the range when species disperse randomly rather than in response to thermal habitat suitability. In this way, dispersal mortality can facilitate the evolution of dispersal modes that maximize fitness in warmer thermal environments.

Increase in heat tolerance following a period of heat stress in a naturally occurring insect species.

Climate warming is the defining environmental crisis of the 21st century. Elucidating whether organisms can adapt to rapidly changing thermal environments is therefore a crucial research priority. We investigated warming effects on a native Hemipteran insect (Murgantia histrionica) that feeds on an endemic plant species (Isomeris arborea) of the California coastal sage scrub. Experiments conducted in 2009 quantified the temperature responses of juvenile maturation rates and stage-specific and cumulative survivorship. The intervening decade has seen some of the hottest years ever recorded, with increasing mean temperatures accompanied by an increase in the frequency of hot extremes. Experiments repeated in 2021 show a striking change in the bugs' temperature responses. In 2009, no eggs developed past the second nymphal stage at 33°C. In 2021, eggs developed into reproductive adults at 33°C. Upper thermal limits for maturation and survivorship have increased, along with a decrease in mortality risk with increasing age and temperature, and a decrease in the temperature sensitivity of mortality with increasing age. While we cannot exclude the possibility that other environmental factors occurring in concert could have affected our findings, the fact that all observed trait changes are in the direction of greater heat tolerance suggests that consistent exposure to extreme heat stress may at least be partially responsible for these changes. Harlequin bugs belong to the suborder Heteroptera, which contains a number of economically important pests, biological control agents and disease carriers. Their differential success in withstanding warming compared to beneficial holometabolous insects such as pollinators may exacerbate the decline of beneficial insects due to other causes (e.g. pollution and pesticides) with potentially serious consequences on both biodiversity and ecosystem functioning.

Persistence of tri-trophic interactions in seasonal environments.

The interplay between species interactions and environmental variation is well-understood for pairwise interactions but not for multi-trophic interactions. Understanding how such interactions persist in a thermally variable environment is particularly important given that most biodiversity on the planet consists of ectotherms whose body temperature depends on the environmental temperature. Here we present a trait-based mathematical framework for investigating how tri-trophic food chains persist in seasonal environments. We report two key findings. First, the persistence of the tri-trophic interaction is enhanced if species at upper trophic levels (e.g. top predators) are more cold-adapted than those at lower levels (e.g. basal resources) by virtue of lower thermal optima, wider response breadths and lower mortality within the favourable temperature range. The important implication is that the assembly and persistence of multi-trophic interactions requires that species at lower trophic levels be somewhat maladapted to their ambient thermal environment, as in the case of recent invasions. Second, differential sensitivity to thermally varying environments provides a mechanistic explanation for the conflict of interest between the intermediate consumer and top predator. The same cold-adaptations that increase the consumer's ability to increase when rare deter the predator's ability to do so. Thus, being well-adapted to its thermal environment makes the intermediate consumer better able to acquire resources and avoid predators. We predict that the hierarchy in cold-adaptation should constrain the number of trophic levels that can be supported in a given thermal environment, and that ectotherm food chain lengths should increase with increasing latitude because larger-amplitude seasonal fluctuations generate more opportunities for species to diverge in their thermal optima.

Latitudinal directionality in ectotherm invasion success.

A striking pattern, seen in both fossil and extant taxa, is that tropical ectotherms are better at invading temperate habitats than vice versa. This is puzzling because tropical ectotherms, being thermal specialists, face a harsher abiotic environment and competition from temperate residents that are thermal generalists. We develop a mathematical framework to address this puzzle. We find that (i) tropical ectotherms can invade temperate habitats if they have higher consumption rates and lower mortality during warmer summers, (ii) stronger seasonal fluctuations at higher latitudes create more temporal niches, allowing coexistence of tropical invaders and temperate residents, and (iii) temperate ectotherms' failure to invade tropical habitats is due to greater mortality rather than lower competitive ability. Our framework yields predictions about population-level outcomes of invasion success based solely on species' trait responses to temperature. It provides a potential ecological explanation for why the tropics constitute both a cradle and a museum of biodiversity.

Predicting phenological shifts in a changing climate.

Phenological shifts constitute one of the clearest manifestations of climate warming. Advanced emergence is widely reported in high-latitude ectotherms, but a significant number of species exhibit delayed, or no change in, emergence. Here we present a mechanistic theoretical framework that reconciles these disparate observations and predicts population-level phenological patterns based solely on data on temperature responses of the underlying life history traits. Our model, parameterized with data from insects at different latitudes, shows that peak abundance occurs earlier in the year when warming increases the mean environmental temperature, but is delayed when warming increases the amplitude of seasonal fluctuations. We find that warming does not necessarily lead to a longer activity period in high-latitude species because it elevates summer temperatures above the upper limit for reproduction and development. Our findings both confirm and confound expectations for ectotherm species affected by climate warming: an increase in the mean temperature is more detrimental to low-latitude species adapted to high mean temperatures and low-amplitude seasonal fluctuations; an increase in seasonal fluctuations is more detrimental to high-latitude species adapted to low mean temperatures and high-amplitude fluctuations.

Effects of warming on predator-prey interactions - a resource-based approach and a theoretical synthesis.

We theoretically explore consequences of warming for predator-prey dynamics, broadening previous approaches in three ways: we include beyond-optimal temperatures, predators may have a type III functional response, and prey carrying capacity depends on explicitly modelled resources. Several robust patterns arise. The relationship between prey carrying capacity and temperature can range from near-independence to monotonically declining/increasing to hump-shaped. Predators persist in a U-shaped region in resource supply (=enrichment)-temperature space. Type II responses yield stable persistence in a U-shaped band inside this region, giving way to limit cycles with enrichment at all temperatures. In contrast, type III responses convey stability at intermediate temperatures and confine cycles to low and high temperatures. Warming-induced state shifts can be predicted from system trajectories crossing stability and persistence boundaries in enrichment-temperature space. Results of earlier studies with more restricted assumptions map onto this graph as special cases. Our approach thus provides a unifying framework for understanding warming effects on trophic dynamics.

Evolution of Thermal Reaction Norms in Seasonally Varying Environments.

Thermal reaction norms of ectotherms exhibit a distinctive latitudinal pattern: the temperature at which performance is maximized coincides with the mean habitat temperature in tropical ectotherms but exceeds the mean temperature in temperate ectotherms. We hypothesize, on the basis of Jensen's inequality, that this pattern is driven by latitudinal variation in seasonal temperature fluctuations. We test this hypothesis with an eco-evolutionary model that integrates the quantitative genetics of reaction norm evolution with stage-structured population dynamics, which we parameterize with data from insects. We find that thermal optima of temperate and Mediterranean species evolve to exceed the mean habitat temperature if seasonal fluctuations are strong, while the thermal optimum of tropical species evolves to coincide with the mean habitat temperature if fluctuations are weak. Importantly, ecological dynamics can impose a constraint on reaction norm evolution. Tropical species cannot tolerate an increase in seasonal fluctuations at the high mean habitat temperature it experiences, while the temperate species cannot tolerate a reduction in seasonal fluctuations if the mean temperature is higher. In both cases, stochastic extinction during periods of low abundances precludes adaptation to a novel thermal environment. Our findings suggest a potential directionality in colonization success. Tropical ectotherms, because of their high thermal optima, can successfully colonize temperate habitats, while temperate ectotherms, because of their low optima, are less successful in colonizing tropical habitats.

Effects of temperature and resource variation on insect population dynamics: the bordered plant bug as a case study.

In species with complex life cycles, population dynamics result from a combination of intrinsic cycles arising from delays in the operation of negative density-dependent processes (e.g., intraspecific competition) and extrinsic fluctuations arising from seasonal variation in the abiotic environment. Abiotic variation can affect species directly through their life history traits and indirectly by modulating the species' interactions with resources or natural enemies.We investigate how the interplay between density-dependent dynamics and abiotic variability affects population dynamics of the bordered plant bug (Largus californicus), a Hemipteran herbivore inhabiting the California coastal sage scrub community. Field data show a striking pattern in abundance: adults are extremely abundant or nearly absent during certain periods of the year, leading us to predict that seasonal forcing plays a role in driving observed dynamics.We develop a stage-structured population model with variable developmental delays, in which fecundity is affected by both intra-specific competition and temporal variation in resource availability and all life history traits (reproduction, development, mortality) are temperature-dependent. We parameterize the model with experimental data on temperature-responses of life history and competitive traits and validate the model with independent field census data.We find that intra-specific competition is strongest at temperatures optimal for reproduction, which theory predicts leads to more complex population dynamics. Our model predicts that while temperature or resource variability interact with development-induced delays in self-limitation to generate population fluctuations, it is the interplay between all three factors that drive the observed dynamics. Considering how multiple abiotic factors interact with density-dependent processes is important both for understanding how species persist in variable environments and predicting species' responses to perturbations in their typical environment.

Selection on stability across ecological scales.

Much of the focus in evolutionary biology has been on the adaptive differentiation among organisms. It is equally important to understand the processes that result in similarities of structure among systems. Here, we discuss examples of similarities occurring at different ecological scales, from predator-prey relations (attack rates and handling times) through communities (food-web structures) to ecosystem properties. Selection among systemic configurations or patterns that differ in their intrinsic stability should lead generally to increased representation of relatively stable structures. Such nonadaptive, but selective processes that shape ecological communities offer an enticing mechanism for generating widely observed similarities, and have sparked new interest in stability properties. This nonadaptive systemic selection operates not in opposition to, but in parallel with, adaptive evolution.

A metric for quantifying the oscillatory tendency of consumer-resource interactions.

The oscillatory tendency of consumer-resource interactions is a key determinant of food-web persistence. Here, we develop a metric for quantifying oscillatory tendency that scales the positive feedback effects of saturating functional responses with the negative feedback effects of self-limitation. We use this metric to predict the oscillatory tendency of a pairwise interaction, tritrophic chain, and tritrophic web. This framework yields two key predictions. First, the oscillatory tendency of any food web increases with the number of trophic links with long handling times regardless of the magnitude of attack rates. Attack rates influence oscillatory tendency only when handling times are short. Second, the realized oscillatory tendency of a trophic link depends on how the product of the attack rate and handling time scales with the strength of self-limitation. Importantly, our metric allows calculations of the critical self-limitation strength at which a consumer-resource interaction moves from stable to oscillatory dynamics. Our data analysis reveals that the majority (77%) of interactions involve low attack rates and handling times, requiring only a modest level of self-limitation to suppress oscillations. Only 23% of the interactions exhibit a strong oscillatory tendency, consistent with previous findings, based on time-series data, that 30% of consumer-resource interactions in nature exhibit oscillations.

Effects of temperature on consumer-resource interactions.

Understanding how temperature variation influences the negative (e.g. self-limitation) and positive (e.g. saturating functional responses) feedback processes that characterize consumer-resource interactions is an important research priority. Previous work on this topic has yielded conflicting outcomes with some studies predicting that warming should increase consumer-resource oscillations and others predicting that warming should decrease consumer-resource oscillations. Here, I develop a consumer-resource model that both synthesizes previous findings in a common framework and yields novel insights about temperature effects on consumer-resource dynamics. I report three key findings. First, when the resource species' birth rate exhibits a unimodal temperature response, as demonstrated by a large number of empirical studies, the temperature range over which the consumer-resource interaction can persist is determined by the lower and upper temperature limits to the resource species' reproduction. This contrasts with the predictions of previous studies, which assume that the birth rate exhibits a monotonic temperature response, that consumer extinction is determined by temperature effects on consumer species' traits, rather than the resource species' traits. Secondly, the comparative analysis I have conducted shows that whether warming leads to an increase or decrease in consumer-resource oscillations depends on the manner in which temperature affects intraspecific competition. When the strength of self-limitation increases monotonically with temperature, warming causes a decrease in consumer-resource oscillations. However, if self-limitation is strongest at temperatures physiologically optimal for reproduction, a scenario previously unanalysed by theory but amply substantiated by empirical data, warming can cause an increase in consumer-resource oscillations. Thirdly, the model yields testable comparative predictions about consumer-resource dynamics under alternative hypotheses for how temperature affects competitive and resource acquisition traits. Importantly, it does so through empirically quantifiable metrics for predicting temperature effects on consumer viability and consumer-resource oscillations, which obviates the need for parameterizing complex dynamical models. Tests of these metrics with empirical data on a host-parasitoid interaction yield realistic estimates of temperature limits for consumer persistence and the propensity for consumer-resource oscillations, highlighting their utility in predicting temperature effects, particularly warming, on consumer-resource interactions in both natural and agricultural settings.

Effects of temperature on intraspecific competition in ectotherms.

Understanding how temperature influences population regulation through its effects on intraspecific competition is an important question for which there is currently little theory or data. Here we develop a theoretical framework for elucidating temperature effects on competition that integrates mechanistic descriptions of life-history trait responses to temperature with population models that realistically capture the variable developmental delays that characterize ectotherm life cycles. This framework yields testable comparative predictions about how intraspecific competition affects reproduction, development, and mortality under alternative hypotheses about the temperature dependence of competition. The key finding is that ectotherm population regulation in seasonal environments depends crucially on the mechanisms by which temperature affects competition. When competition is strongest at temperatures optimal for reproduction, effects of temperature and competition act antagonistically, leading to more complex dynamics than when competition is temperature independent. When the strength of competition increases with temperature past the optimal temperature for reproduction, effects of temperature and competition act synergistically, leading to dynamics qualitatively similar to those when competition is temperature independent. Paradoxically, antagonistic effects yield a higher population floor despite greater fluctuations. These findings have important implications for predicting effects of climate warming on population regulation. Synergistic effects of temperature and competition can predispose populations to stochastic extinction by lowering minimum population sizes, while antagonistic effects can increase the potential for population outbreaks through greater fluctuations in abundance.

Modeling oncolytic virotherapy: is complete tumor-tropism too much of a good thing?

The specific targeting of tumor cells by replication-competent oncolytic viruses is considered indispensable for realizing the potential of oncolytic virotherapy. Yet off-target infections by oncolytic viruses may increase virus production, further reducing tumor load. This ability may be critical when tumor-cell scarcity or the onset of an adaptive immune response constrain viral anti-tumoral efficacy. Here we develop a mathematical framework for assessing whether oncolytic viruses with reduced tumor-specificity can more effectively eliminate tumors while keeping losses to normal cell populations low. We find viruses that infect some normal cells can potentially balance the competing goals of tumor elimination and minimizing the effects on normal cell populations. Particularly when infected tissues can be regenerated, moderating rather than completely eliminating the ability of oncolytic viruses to infect and lyse normal cells could improve cancer treatment, with potentially fewer side-effects than conventional treatments such as chemotherapy.

The intrinsic growth rate as a predictor of population viability under climate warming.

1. Lately, there has been interest in using the intrinsic growth rate (rm) to predict the effects of climate warming on ectotherm population viability. However, because rm is calculated using the Euler-Lotka equation, its reliability in predicting population persistence depends on whether ectotherm populations can achieve a stable age/stage distribution in thermally variable environments. Here, we investigate this issue using a mathematical framework that incorporates mechanistic descriptions of temperature effects on vital rates into a stage-structured population model that realistically captures the temperature-induced variability in developmental delays that characterize ectotherm life cycles. 2. We find that populations experiencing seasonal temperature variation converge to a stage distribution whose intra-annual pattern remains invariant across years. As a result, the mean annual per capita growth rate also remains constant between years. The key insight is the mechanism that allows populations converge to a stationary stage distribution. Temperature effects on the biochemical processes (e.g. enzyme kinetics, hormonal regulation) that underlie life-history traits (reproduction, development and mortality) exhibit well-defined thermodynamical properties (e.g. changes in entropy and enthalpy) that lead to predictable outcomes (e.g. reduction in reaction rates or hormonal action at temperature extremes). As a result, life-history traits exhibit a systematic and predictable response to seasonal temperature variation. This in turn leads to temporally predictable temperature responses of the stage distribution and the per capita growth rate. 3. When climate warming causes an increase in the mean annual temperature and/or the amplitude of seasonal fluctuations, the population model predicts the mean annual per capita growth rate to decline to zero within 100 years when warming is slow relative to the developmental period of the organism (0.03-0.05°C per year) and to become negative, causing population extinction, well before 100 years when warming is fast (e.g. 0.1°C per year). The Euler-Lotka equation predicts a slower decrease in rm when warming is slow and a longer persistence time when warming is fast, with the deviation between the two metrics increasing with increasing developmental period. These results suggest that predictions of ectotherm population viability based on rm may be valid only for species with short developmental delays, and even then, only over short time-scales and under slow warming regimes.

Competition for benefits can promote the persistence of mutualistic interactions.

Mutualistic interactions are characterized by positive density-dependence that should cause interacting species to go extinct when rare. However, data show mutualistic interactions to be common and persistent. Previous theory predicts persistence provided that mutualistic species are regulated by factors external to the mutualistic interaction (e.g., limiting background resources). Empirical data suggest that competition for the benefits provided by mutualistic partners could be a source of negative density-dependence that allows for population regulation, but there is little, if any, theoretical exploration of this mechanism. Here we develop mathematical models to investigate whether competition for benefits alone can allow the persistence of obligate mutualistic interactions. We consider the role of trade-offs in persistence, specifically, trade-offs between benefits acquired versus given and between competition for access to partners (competitive ability) and benefit acquisition. We find that competition for benefits alone is sufficient to promote the persistence of pairwise interactions and the assembly of a three-species community module from an initially pairwise interaction. We find that a trade-off between benefits acquired versus given reduces opportunities for cheating (because a species that acquires significantly more benefits than it gives drives its partner extinct), while a trade-off between competitive ability and benefit acquisition facilitates persistence when it is weak, but constrains persistence when it is strong. When both trade-offs operate simultaneously, persistence requires that each species acquire sufficient benefits to avoid being cheated by its partners, but not so much that it loses its competitive ability. The key finding is that competition for benefits provides a biologically-realistic mechanism for the long-term persistence of mutualistic interactions and the assembly of complex community modules from initially pairwise interactions.

The biological control of disease vectors.

Vector-borne diseases are common in nature and can have a large impact on humans, livestock and crops. Biological control of vectors using natural enemies or competitors can reduce vector density and hence disease transmission. However, the indirect interactions inherent in host-vector disease systems make it difficult to use traditional pest control theory to guide biological control of disease vectors. This necessitates a conceptual framework that explicitly considers a range of indirect interactions between the host-vector disease system and the vector's biological control agent. Here we conduct a comparative analysis of the efficacy of different types of biological control agents in controlling vector-borne diseases. We report three key findings. First, highly efficient predators and parasitoids of the vector prove to be effective biological control agents, but highly virulent pathogens of the vector also require a high transmission rate to be effective. Second, biocontrol agents can successfully reduce long-term host disease incidence even though they may fail to reduce long-term vector densities. Third, inundating a host-vector disease system with a natural enemy of the vector has little or no effect on reducing disease incidence, but inundating the system with a competitor of the vector has a large effect on reducing disease incidence. The comparative framework yields predictions that are useful in developing biological control strategies for vector-borne diseases. We discuss how these predictions can inform ongoing biological control efforts for host-vector disease systems.

A framework for elucidating the temperature dependence of fitness.

Climate warming is predicted to cause large-scale extinctions, particularly of ectothermic species. A striking difference between tropical and temperate ectotherms is that tropical species experience a mean habitat temperature that is closer to the temperature at which fitness is maximized (T(opt)) and an upper temperature limit for survival (T(max)) that is closer to T(opt) than do temperate species. Thus, even a small increase in environmental temperature could put tropical ectotherms at high risk of extinction, whereas temperate ectotherms have a wider temperature cushion. Although this pattern is widely observed, the mechanisms that produce it are not well understood. Here we develop a mathematical framework to partition the temperature response of fitness into its components (fecundity, mortality, and development) and test model predictions with data for insects. We find that fitness declines at high temperatures because the temperature responses of fecundity and mortality act in opposite ways: fecundity decreases with temperature when temperatures exceed the optimal range, whereas mortality continues to increase. The proximity of T(opt) to T(max) depends on how the temperature response of development mediates the interaction between fecundity and mortality. When development is highly temperature sensitive, mortality exceeds reproduction only after fecundity has started to decline with temperature, which causes fitness to decline rapidly to zero when temperatures exceed T(opt). The model correctly predicts empirically observed fitness-temperature relationships in insects from different latitudes. It also suggests explanations for the widely reported phenological shifts in many ectotherms and the latitudinal differences in fitness responses.

The role of transient dynamics in biological pest control: insights from a host-parasitoid community.

1. Identifying natural enemies that can maintain pests at low abundances is a priority in biological control. Here, we show that experiments combined with models generate new insights into identifying effective control agents prior to their release in the field. Using a host-parasitoid community (the harlequin bug and its egg parasitoids) as a model system, we report three key findings. 2. The interplay between the host's self-limitation and the parasitoids' saturating functional response causes the long-term (steady-state) outcomes for pest suppression to differ from those of short-term (transient) dynamics. When the bug's self-limitation is moderately strong, the parasitoid with the higher attack rate and conversion efficiency (Ooencyrtus) achieves greater host suppression in the long term, but its longer handling time causes long periods of transient dynamics during which the bug can reach high abundances; when the bug's self-limitation is weak, host fluctuations amplify over time and Ooencyrtus fails at host suppression altogether. In contrast, the parasitoid with the lower attack rate and conversion efficiency but the shorter handling time (Trissolcus) induces only weak transient fluctuations of short duration and can maintain the host at low abundances regardless of the strength of the bug's self-limitation. 3. Release of multiple enemy species can compromise host suppression if an enemy that induces stronger transient fluctuations excludes one that induces weaker fluctuations. For instance, Ooencyrtus excludes Trissolcus despite having a longer handling time because of its higher conversion efficiency. The model correctly predicts the time to exclusion observed in experiments, suggesting that it captures the key biological features of the host-parasitoid interaction. 4. Intraspecific interference reduces long-term pest suppression but improves short-term pest control by reducing the magnitude and duration of transient fluctuations. 5. These results highlight the importance of transient dynamics in pest suppression. Pests are unlikely to be strongly self-limited because they attack crop monocultures. Hence, pest fluctuations are likely to dominate short-term dynamics even when the long-term outcome is a stable equilibrium. The tendency to induce strong transient fluctuations (e.g. through a long handling time) is therefore a crucial consideration when identifying effective pest control agents.