Spatial distribution of limited resources and local density regulation in juvenile Atlantic salmon

1. Spatial heterogeneity of resources may influence competition among individuals and thus have a fundamental role in shaping population dynamics and carrying capacity. In the present study, we identify shelter opportunities as a limiting resource for juvenile Atlantic salmon (Salmo salar L.). Experimental and field studies are combined in order to demonstrate how the spatial distribution of shelters may influence population dynamics on both within and among population scales. 2. In closed experimental streams, fish performance scaled negatively with decreasing shelter availability and increasing densities. In contrast, the fish in open stream channels dispersed according to shelter availability and performance of fish remaining in the streams did not depend on initial density or shelters. 3. The field study confirmed that spatial variation in densities of 1-year-old juveniles was governed both by initial recruit density and shelter availability. Strength of density-dependent population regulation, measured as carrying capacity, increased with decreasing number of shelters. 4. Nine rivers were surveyed for spatial variation in shelter availability and increased shelter heterogeneity tended to decrease maximum observed population size (measured using catch statistics of adult salmon as a proxy). 5. Our studies highlight the importance of small-scale within-population spatial structure in population dynamics and demonstrate that not only the absolute amount of limiting resources but also their spatial arrangement can be an important factor influencing population carrying capacity.     

Shelters and their use by fishes on fringing coral reefs

Coral reef fish density and species richness are often higher at sites with more structural complexity. This association may be due to greater availability of shelters, but surprisingly little is known about the size and density of shelters and their use by coral reef fishes. We quantified shelter availability and use by fishes for the first time on a Caribbean coral reef by counting all holes and overhangs with a minimum entrance diameter ≥3 cm in 30 quadrats (25 m(2)) on two fringing reefs in Barbados. Shelter size was highly variable, ranging from 42 cm(3) to over 4,000,000 cm(3), with many more small than large shelters. On average, there were 3.8 shelters m(-2), with a median volume of 1,200 cm(3) and a total volume of 52,000 cm(3) m(-2). The number of fish per occupied shelter ranged from 1 to 35 individual fishes belonging to 66 species, with a median of 1. The proportion of shelters occupied and the number of occupants increased strongly with shelter size. Shelter density and total volume increased with substrate complexity, and this relationship varied among reef zones. The density of shelter-using fish was much more strongly predicted by shelter density and median size than by substrate complexity and increased linearly with shelter density, indicating that shelter availability is a limiting resource for some coral reef fishes. The results demonstrate the importance of large shelters for fish density and support the hypothesis that structural complexity is associated with fish abundance, at least in part, due to its association with shelter availability. This information can help identify critical habitat for coral reef fishes, predict the effects of reductions in structural complexity of natural reefs and improve the design of artificial reefs.     

Inter- and intra-specific patterns of density dependence and population size variability in Salmoniformes

Population dynamics are typically affected by a combination of density-independent and density-dependent factors, the latter of which have been conceptually and theoretically linked with how variable population sizes are over time—which in turn has been tied to how prone populations are to extinction. To address evidence for the occurrence of density dependence and its relationship with population size variability (pv), we quantified each of these for 126 populations of 8 species of Salmoniformes. Using random-effects models, we partitioned variation in the strength of density dependence and the magnitude of pv between and within species and estimated the correlation of density dependence and population size variability at both the between- and within-species levels. We found that variation in the strength of density dependence was predominately within species (I ² = 0.47). In contrast, variation in population size variability was distributed both between and within species (I ² = 0.40). Contrary to theoretical and conceptual expectations, the strength of density dependence and the magnitude of population size variability were positively correlated at the between species level (r = 0.90), although this estimate had 95 % credibility intervals (Bayesian analogues to confidence intervals) that overlapped zero. The within-species correlation between density dependence and population size variability was not distinguishable from zero. Given that density dependence for Salmoniformes was highly variable within species, we next determined the joint effects of intrinsic (density-dependent) and extrinsic (density-independent) factors on the population dynamics of a threatened salmonid, the Lahontan cutthroat trout (Oncorhynchus clarkii henshawi). We found that density-dependent and -independent factors additively contributed to population dynamics. This finding suggests that the observed within-species variability in density dependence might be attributable to local differences in the strength of density-independent factors.     

Female choice and male reproductive success in the angel blenny,Coralliozetus angelica (Teleostei: Chaenopsidae)

Correlates of reproductive success of male Coralliozetus angelica were studied in field populations in the central Gulf of California by collecting males with their shelters (vacant barnacle tests) and counting the number of eggs defended. Male body size was positively correlated with the number of eggs defended. This apparently resulted from female preference for large males since females preferred to mate with the larger of two males in laboratory choice experiments. Condition of males, as assessed by their relative head width, was also positively correlated with the number of eggs defended. Fit of males in their shelter entrance, male colour intensity, and male size relative to immediately surrounding males were not correlated with reproductive success. The quality of shelters was an important determinant of male reproductive success since females apparently avoided mating with males in heavily fouled shelters. Large shelters tended to be more heavily fouled than small shelters and male size and shelter size were positively correlated. The resulting negative correlation of male size and shelter quality apparently constrained female preference for large males and may be an important factor limiting the degree of polygyny in this species.     

The influence of shelter availability on recruitment and early juvenile survivorship of Lythrypnus dalli Gilbert (Pisces: Gobiidae)

The bluebanded goby, Lythrypnus dalli Gilbert, is a common inhabitant of rocky subtidal reefs along the coast of southern California. Experiments utilizing artificial habitats indicate that recruitment and early juvenile survivorship are limited by the presence of adult conspecifics and predators. However, the level of effect is strongly influenced by shelter availability. Both adult and predator effects on limiting juvenile density are significantly reduced as the availability of shelters is increased. Susceptibility to predation is greatest among smaller size classes of fish and is likely to result from displacement from shelters by larger fish. Existing population structures probably reflect the availability of adequate shelter sizes and relative competitive abilities among fish.     

What makes a species common? No evidence of density-dependent recruitment or mortality of the sea urchin Diadema antillarum after the 1983–1984 mass mortality

A potential consequence of individuals compensating for density-dependent processes is that rare or infrequent events can produce profound and long-term shifts in species abundance. In 1983–1984 a mass mortality event reduced the numbers of the abundant sea urchin Diadema antillarum by 95–99 % throughout the Caribbean and western Atlantic. Following this event, the abundance of macroalgae increased and the few surviving D. antillarum responded by increasing in body size and fecundity. These initial observations suggested that populations of D. antillarum could recover rapidly following release from food limitation. In contrast, published studies of field manipulations indicate that this species had traits making it resistant to density-dependent effects on offspring production and adult mortality; this evidence raises the possibility that density-independent processes might keep populations at a diminished level. Decadal-scale (1983–2011) monitoring of recruitment, mortality, population density and size structure of D. antillarum from St John, US Virgin Islands, indicates that population density has remained relatively stable and more than an order of magnitude lower than that before the mortality event of 1983–1984. We detected no evidence of density-dependent mortality or recruitment since this mortality event. In this location, model estimates of equilibrium population density, assuming density-independent processes and based on parameters generated over the first decade following the mortality event, accurately predict the low population density 20 years later (2011). We find no evidence to support the notion that this historically dominant species will rebound from this temporally brief, but spatially widespread, perturbation.     

Geomorphological features drive spatiotemporal dynamics of young-of-the-year brown trout populations in a large New Zealand river catchment

1. Determining the factors driving population dynamics of stream salmonids across broad landscapes is important for understanding stream ecosystem functioning and for the management and conservation of the populations of such fish. A few studies have described the determinants of the early life history of salmonids across catchments, but none of them have examined how this spatial variation develops temporally.
2. We hypothesised that: (1) spatiotemporal dynamics of young-of-the-year (YoY) trout populations vary within the catchment, with loss rates (emigration plus mortality) being higher in lower elevation, downstream reaches than in higher elevation, upstream ones; and (2) loss rate would be density dependent, leading to reduced intra- and inter-cohort competition and supporting establishment of resident populations.
3. We tested the first hypothesis by examining relationships between geomorphic characteristics of spawning streams and temporal dynamics of YoY brown trout density, loss rate and biological traits through an austral summer across the catchment of New Zealand's fourth-longest river, the Taieri River. To test the second hypothesis, we examined whether initial YoY density in spring and other biotic determinants affected loss rate and cohort structure dynamics, resulting in density-dependent self-thinning of YoY fish to avoid resource limitation.
4. Spring YoY density and subsequent loss rate was higher in low elevation sites, corresponding to stream accessibility to migratory spawners. These significant correlations of YoY density and related biotic variables with elevation disappeared later in the season, indicating that regardless of their origin, brown trout YoY tend to form stream resident populations with multicohort structure and densities low enough to limit intraspecific competition.
5. Our findings provide evidence that geomorphological features of the landscape affect temporal dynamics of YoY trout populations driven by density-dependent self-regulatory mechanisms that can control the abundance of growing fish.

     

Summer mortality in the hazel dormouse (Muscardinus avellanarius) and its effect on population dynamics

In the period 2000–2012, 38.3 % of 950 marked overwintered hazel dormice (Muscardinus avellanarius) were not recaptured at a study site in Lithuania in autumn. As adult dormice are sedentary, it is presumed that those dormice not recaptured died between late April and August. The highest total number of dormice captured for the last time was recorded in May and the lowest in August. The total summer mortality was significantly higher in females (42.5 %) than in males (34.6 %), but it did not depend on dormouse age or body weight. Tawny owl (Strix aluco) is the main known dormouse predator in Lithuania, and likely, it has the highest impact on summer mortality of M. avellanarius. Over the years, the total summer mortality of adult dormice ranged from 27 % to 52 %. The increased summer mortality resulted in decreased total dormouse population density and particularly decreased density of adult females in summer. Decreased densities led to more intensive breeding in the dormouse population, specifically breeding by young-of-the-year females, a pattern that is not common for this species. The number of breeding cases by young-of-the-year females was inversely related to the density of adult overwintered females in summer and to the number of breeding cases of these females. Breeding by young-of-the-year females was the main factor in the restoration of decreased population density in summer. Lithuanian populations of M. avellanarius are unique in their high proportion of breeding cases by young-of-the-year females amongst all populations investigated in the entire species distributional range.     

Distribution of life cycle stages in a lithophytic and epiphytic orchid

Density-dependent processes may have multiple effects on populations, which among other things include the regulation of population abundance and of the relative distribution of life-cycle stages within populations. The epiphytic habitat is often characterized as highly ephemeral and therefore epiphytic orchid populations may never achieve density-dependent regulation. In this study, we investigated the potential for density-dependent regulation in epiphytic and lithophytic orchids by examining the association between seedlings, juvenile and adult life-history stages in the Caribbean endemic orchid,Lepanthes rupestris in a cross-sectional study of 179 populations surveyed in the Luquillo National Forest along a riparian area where it is locally abundant. Under density-dependent regulation we expected a negative association between the ratio of seedling/adults and juveniles/adults and total population density. Population density was in the range of 140 individuals per m², however patch sizes were small and mostly limited to less than 0.5 m² with a maximum of 3 m². We found no evidence of reduction of the ratio of seedlings or juveniles to adults as population size increased in either tree or boulder populations suggesting negative density dependence for population regulation inL. rupestris is either rare or occurs at even higher densities than those measured here. Moreover, we found positive (although weak) relationship between the ratio of seedlings and juveniles to adults and population size, suggesting that facilitation may be occurring.     

Contestable shelters provoke aggression among 0+ perch, <Emphasis Type="BoldItalic">Perca fluviatilis</Emphasis>

Synopsis We examined behavioural interactions and feeding within triads of young-of-the-year (YOY) Eurasian perch, Perca fluviatilis, in aquaria with and without a shelter. For the first time we showed that competition for shelters, but not for food, provokes aggressiveness and triggers establishment of social hierarchy among young perch. No aggressiveness occurred during feeding bouts, when food was a limited resource. We expect the observed interactions to occur in natural heterogeneous habitats providing limited number of refuges and high local density of fish. They may ultimately lead to previously anticipated interference competition among juvenile perch.     

Recruitment limitation: A theoretical perspective

Abstract A theoretical analysis of the concept of recruitment limitation leads to the conclusion that most populations should he regarded as jointly limited by recruitment and interactions between individuals after recruitment. The open nature of local marine systems does not permit avoidance of density-dependent interactions; it simply may make them more difficult to detect. Local populations consisting of settled organisms may not experience density-dependent interactions under some circumstances, but the entire species population consisting of the collection of local populations and their planktonic larvae must have density-dependent dynamics. Any local population of settled individuals can escape density dependence if sufficient density dependence occurs among planktonic larvae or within other local populations. Common conceptions of density dependence are too narrow, leading too often to the conclusion that it is absent from a system. It is equally wrong to expect that density-dependent interactions after settlement determine local population densities independently of recruitment. Special circumstances allowing density dependence to act strongly and quickly are needed before density dependence can neutralize the effects of recruitment. Recruitment limitation and density-dependent interactions therefore should not be regarded as alternatives but as jointly acting to determine the densities of marine benthic populations. Moreover, the interaction between fluctuating recruitment and density dependence is potentially the most interesting feature of recruitment limitation. For example, this interaction may be an important diversity-maintaining mechanism for marine systems.     

Effects of strength and timing of harvest on seasonal population models: stability switches and catastrophic shifts

Population abundance of many species is controlled by a combination of density-dependent processes during different periods of the annual cycle. In the context of population exploitation or conservation programs, sequential density dependence has the potential to dramatically change population responses to harvesting. Looking for a better understanding of the potential effects of harvesting on the dynamics of seasonal populations, we carry out a theoretical analysis of a discrete model for a semelparous population with an annual cycle involving three discrete density-dependent events: breeding, natural mortality, and harvesting. Our study reveals how the interplay between the model parameters determines the importance of harvest timing on stability and population abundance, especially when two nontrivial stable equilibria coexist. We address the possibility for compensatory mortality and report different forms of the hydra effect, including non-smooth ones due to catastrophic shifts. These drastic switches may include hysteresis, which has important implications for conservation goals. Regarding variability, we show that increasing the harvesting effort may either stabilize or destabilize the population, and these effects strongly depend on harvest timing and natural mortality rates. Our results also emphasize the importance of sampling populations after every discrete event occurs during one cycle. Indeed, though the dynamics are not affected by census timing, the model shows that changes in population abundance in response to changes in harvesting pressure are substantially different depending on when population is sampled. Thus, a manager would receive different (and sometimes contradictory) messages depending on census time, which could lead to managing mistakes.     

Predicting the Timing and Magnitude of Tropical Mosquito Population Peaks for Maximizing Control Efficiency

The transmission of mosquito-borne diseases is strongly linked to the abundance of the host vector. Identifying the environmental and biological precursors which herald the onset of peaks in mosquito abundance would give health and land-use managers the capacity to predict the timing and distribution of the most efficient and cost-effective mosquito control. We analysed a 15-year time series of monthly abundance of Aedes vigilax, a tropical mosquito species from northern Australia, to determine periodicity and drivers of population peaks (high-density outbreaks). Two sets of density-dependent models were used to examine the correlation between mosquito abundance peaks and the environmental drivers of peaks or troughs (low-density periods). The seasonal peaks of reproduction (r) and abundance () occur at the beginning of September and early November, respectively. The combination of low mosquito abundance and a low frequency of a high tide exceeding 7 m in the previous low-abundance (trough) period were the most parsimonious predictors of a peak''s magnitude, with this model explaining over 50% of the deviance in . Model weights, estimated using AIC_c, were also relatively high for those including monthly maximum tide height, monthly accumulated tide height or total rainfall per month in the trough, with high values in the trough correlating negatively with the onset of a high-abundance peak. These findings illustrate that basic environmental monitoring data can be coupled with relatively simple density feedback models to predict the timing and magnitude of mosquito abundance peaks. Decision-makers can use these methods to determine optimal levels of control (i.e., least-cost measures yielding the largest decline in mosquito abundance) and so reduce the risk of disease outbreaks in human populations.     

Adaptive plasticity in ontogenetic niche shifts stabilizes consumer-resource dynamics

Ontogenetic niche shifts, changes in the diet or habitats of organisms during their ontogeny, are widespread among various animal taxa. Ontogenetic niche shifts introduce stage structure in a population with different stages interacting with different communities and can substantially affect their dynamics. In this article, I use mathematical models to test the hypothesis that adaptive plasticity in the timing of ontogenetic niche shifts has a stabilizing effect on consumer-resource dynamics. Adaptive plasticity allows consumers in one ontogenetic niche to perform an early shift to the next ontogenetic niche if the resource density of the first niche is low. The early shift will reduce predation by the consumer on the scarce resource. On the other hand, adaptive plasticity allows consumers to delay their shift to the next niche if the resource density of the first niche is high. The delayed shift will increase the predation on the abundant resource. As a result, the scarce resource will tend to increase, and the abundant resource will tend to decrease. This causes density-dependent negative feedback in the resource dynamics, which stabilizes the consumer-resource dynamics. To test this hypothesis, I compare three consumer-resource models differing in terms of mechanisms controlling the timing of the ontogenetic niche shift: the fixed-age model assumes that the age at which the ontogenetic niche shift occurs is fixed; the fixed-size model assumes that the size at the shift is fixed; and the adaptive plasticity model assumes that the timing of the shift is such that the individual fitness of the consumer is maximized. I show that only the adaptive plasticity model has a locally stable equilibrium and that the stabilizing effect is due to the density-dependent negative feedback in the resource dynamics. I discuss the ontogenetic niche shifts of lake fish in light of the obtained result.     

Comparative population dynamics of <Emphasis Type="Italic">Peromyscus leucopus</Emphasis> in North America: influences of climate,food, and density dependence

Temporal variation in population size is regulated by both exogenous forces and density-dependent feedbacks. Furthermore, accumulating evidence indicates that temporal and spatial variation in climate and resources can modify the strength of density dependence in animal populations. We analyzed six long-term time series estimates of Peromyscus leucopus (white-footed mouse) abundance from Kansas, Ohio, Pennsylvania, Virginia, Vermont, and Maine, USA, using the Kalman filter. Model-averaged estimates of the strength of delayed density dependence increased from west to east and from south to north. The strength of direct and delayed density dependence was positively related to the annual number of days with minimum temperature below −17.8°C. Annual population growth rates of P. leucopus at the Maine site were positively related to acorn abundance and P. leucopus populations tracked the changes in red-oak acorn abundance. The populations of P. leucopus living in northern latitudes might be more dependent on northern red oak (Quercus rubra) acorns for winter food than P. leucopus in southern latitudes. Furthermore, northern red oak trees mast every 4–5 years. Thus, longer, colder winters in northerly latitudes might result in stronger delayed density dependence in mouse populations with a shortage of winter food. Mice might simply track the acorn fluctuations in a delayed autocorrelated manner; however, delayed density dependence remained in our models for the Maine mouse populations after accounting for acorns, suggesting additional sources for delayed density dependence. Our results suggest that, in seed-eating Peromyscus, cyclicity may be regulated, in part, from low to high trophic levels. Deceased: Jerry O. Wolff     

Temporal and spatial variation of larval parasitism in non-outbreaking populations of a folivorous moth

In order to assess the role of parasitoids in the regulation of non-outbreaking populations of Epirrita autumnata, a geometrid lepidopteran with outbreaking populations in northern Europe, we examined the temporal and spatial variation of larval parasitism in southwestern Finland during 6 successive years. The study was carried out on two spatial scales, among trees within sites of about 1 ha and among sites separated by distances of 2–10 km, using experimental and observational approaches respectively. The overall percent parasitism was independent of host density on both spatial scales, while temporally it fluctuated only little. Of the two main parasitoids, the commoner one, Protapanteles immunis, showed a variable response to host density on the larger spatial scale and negative density dependence on the smaller scale. Temporally, parasitism caused by this species was independent of host density. Another parasitoid, Phobocampe bicingulata, showed positive density dependence on the smaller spatial scale and had a variable response on the larger scale, but exhibited negative density dependence over time. The results of this study caution against drawing conclusions concerning population regulation on the grounds of spatial density dependence alone. Larval parasitoids apparently do not maintain low densities in the E. autumnata populations studied. However, they may suppress E. autumnata densities to a level low enough for density-dependent mortality factor(s) to become regulating. Among other mortality factors of E. autumnata, pupal predation has been found to be temporally positively density-dependent. Received: 19 October 1999 / Accepted: 10 January 2000     

Thermal Carrying Capacity for a Thermally-Sensitive Species at the Warmest Edge of Its Range

Anthropogenic environmental change is causing unprecedented rates of population extirpation and altering the setting of range limits for many species. Significant population declines may occur however before any reduction in range is observed. Determining and modelling the factors driving population size and trends is consequently critical to predict trajectories of change and future extinction risk. We tracked during 12 years 51 populations of a cold-water fish species (brown trout Salmo trutta) living along a temperature gradient at the warmest thermal edge of its range. We developed a carrying capacity model in which maximum population size is limited by physical habitat conditions and regulated through territoriality. We first tested whether population numbers were driven by carrying capacity dynamics and then targeted on establishing (1) the temperature thresholds beyond which population numbers switch from being physical habitat- to temperature-limited; and (2) the rate at which carrying capacity declines with temperature within limiting thermal ranges. Carrying capacity along with emergent density-dependent responses explained up to 76% of spatio-temporal density variability of juveniles and adults but only 50% of young-of-the-year''s. By contrast, young-of-the-year trout were highly sensitive to thermal conditions, their performance declining with temperature at a higher rate than older life stages, and disruptions being triggered at lower temperature thresholds. Results suggest that limiting temperature effects were progressively stronger with increasing anthropogenic disturbance. There was however a critical threshold, matching the incipient thermal limit for survival, beyond which realized density was always below potential numbers irrespective of disturbance intensity. We additionally found a lower threshold, matching the thermal limit for feeding, beyond which even unaltered populations declined. We predict that most of our study populations may become extinct by 2100, depicting the gloomy fate of thermally-sensitive species occurring at thermal range margins under limited potential for adaptation and dispersal.     

Temporal changes in the strength of density-dependent mortality and growth in intertidal barnacles

1. In demographically open marine systems, the extent to which density-dependent processes in the benthic adult phase are required for population persistence is unclear. At one extreme, represented by the recruitment limitation hypothesis, larval supply may be insufficient for the total population size to reach a carrying capacity and density-independent mortality predominates. At the opposite extreme, populations are saturated and density-dependent mortality is sufficiently strong to reshape patterns established at settlement. 2. We examined temporal variation in the way density-independent and density-dependent mortality interact in a typical sessile marine benthic invertebrate, the acorn barnacle Semibalanus balanoides (L.), over a 2-year period. 3. Recruitment was manipulated at two high recruitment sites in north Wales, UK to produce recruit densities covering the range naturally found in this species. Following manipulation, fixed quadrats were monitored using digital photography and temporal changes in mortality and growth rate were examined. 4. Over a 2-year period there was a clear, spatially consistent, over-compensatory relationship between the density of recruits and adult abundance indicating strong density-dependent mortality. The strength of density dependence intensified with increasing recruitment. 5. Density-dependent mortality did not operate consistently over the study period. It only operated in the early part of the benthic phase, but the pattern of adult abundance generated was maintained throughout the whole 2-year period. Thus, early life-history processes dictated adult population abundance and dynamics. 6. Examination of the natural recruitment regime in the area of study indicated that both positive and negative effects of recruitment will occur over scales varying from kilometres to metres.     

Evidence for density-dependent habitat occupancy at varying scales in an expanding bird population

Understanding factors shaping the spatial distribution of animals is crucial for the conservation and management of wildlife species. However, few studies have investigated density-dependent habitat selection in wild populations in non-equilibrium conditions and at varying spatial scales. Here, we investigated density-dependent habitat selection at varying spatial scales in an increasing white stork Ciconia ciconia population using a long term data set in western France. During a 16-year study period, the breeding population density increased from 0.66 nests per 100 km² to 6.6 nests per 100 km². At the beginning of the colonisation of the area settlement probability of storks was mainly positively affected by grasslands located near wetlands. Areas with intensive or moderately intensive agriculture were extremely unlikely to be occupied by breeding birds. However, selection for the initially preferred habitats faded as stork density increased although the proportions of habitat types remained unchanged. At the same time selection for initially less favoured habitats had increased. Moreover, we found that the spatial scale of selection for each foraging habitat variable was consistent over time. Our results suggest that snapshot analyses of resource selection in populations at high density may be misleading for population conservation or management. In contrast, a longitudinal approach to resource selection can be a valuable tool for understanding resource limitation.     

Factors influencing the abundance of Trioxys pallidus,a successful introduced biological control agent of walnut aphid in California

This field study reports on factors influencing the abundance of Trioxys pallidus 35 years after its successful introduction in California, and in the presence of continued hyperparasitism and intraguild predation. The seasonal activity of Chromaphis juglandicola and T. pallidus were monitored in 2005–06 in relation to hyperparasitism and mummy predation in three orchards. Walnut aphids remained under the economic threshold in all three orchards with insecticide treatments for other pests in some cases reducing aphid populations as well. No significant density-independent or density-dependent aggregation in the risk of hyperparasitism was found at the individual tree scale, suggesting the absence of a refuge from hyperparasitism. In contrast, significant negative density-dependent aggregation in the risk of intraguild predation was found, but no density-independent aggregation, suggesting a refuge at higher mummy densities. Using a linear mixed effects model, the following factors were found to influence mummy density: aphid density from the previous sample date and intraguild predation, with the latter linked to hyperparasitism from the previous sample date through a significant interaction that appears to be mediated by current aphid density.