Degree Of Dietary Specialization Research Articles

Colour-specific diet specialization is associated with differences in owlet weight in a polymorphic owl: influence of the trophic quality variation.

The niche divergence hypothesis proposes that the evolution and maintenance of colour polymorphism is based on a mechanism of disruptive selection. In a trophic context, the hypothesis predicts that individuals differing in colour vary in their trophic niche, either because they differ in foraging efficiency or feed in different habitats. A major evolutionary conundrum is how these expectations are affected by variation in trophic quality. Using an owl species with colour plumage polymorphism, the Eurasian scops owl Otus scops, we examined diet and habitat segregation during reproduction in relation to plumage colouration and trophic quality. Intensive sampling revealed that trophic quality for scops owls (i.e. abundance of grasshoppers and locusts) varied more among territories than between years, but scops owls did not segregate among territories of different quality by their colouration. However, we found that sex, plumage colouration and territory differences in trophic quality explained differences in the degree of dietary specialization. Brownish males delivered a higher diversity of prey to the nest than greyish ones in high trophic quality territories. We also found that the more diverse the diet provided by males, the heavier the owlets at fledging. Our study provides evidence for a different sensitivity to trophic quality of the colour morphs with potential fitness consequences in scops owls. We highlight the importance of studying the mechanisms leading to the persistence of colour polymorphism in patchy environments, since segregation may pass otherwise unnoticed if only habitats or years with similar conditions are considered.

Nutrient balancing in a fruit‐specialist primate, the black‐and‐white ruffed lemur (Varecia variegata)

Animals' foraging behavior and dietary choices are, in part, driven by their ultimate function: to meet nutritional demands. However, depending on their degree of dietary specialization and the availability and distribution of food resources in their environment, species may utilize different nutritional strategies. With shifting plant phenology, increasing unpredictability of fruiting, and declining food quality in response to anthropogenic climate change, existing nutritional constraints may become exacerbated. Such changes are especially concerning for Madagascar's endemic fruit specialists given the nutrient-limitation of the island's landscapes. In this study, we examined the nutritional strategy of one such fruit-specialist primate, the black-and-white ruffed lemur (Varecia variegata), over a 12-month period (January to December 2018) in Ranomafana National Park, Madagascar. We hypothesized that Varecia would balance nonprotein energy (NPE) to protein (AP) at a high ratio similar toother frugivorous primates, and that they would prioritize protein intake given their high degree of frugivory. We found that Varecia balance NPE:AP at a ratio of 11:1, higher than in any other primate studied to date; however, diets shifted such that nutrient balancing varied seasonally (12.6:1 abundant-9.6:1 lean). Varecia meet NRC suggested recommendations of 5-8% of calories from protein, despite having a diet mostly comprising fruits. However, seasonal shifts in NPE intakes result in significant energy shortfalls during fruit-lean seasons. Flowers provide an important source of NPE during these periods, with flower consumption best predicting lipid intake, suggesting this species' ability to shift resource use. Nevertheless, achieving adequate and balanced nutrient intakes may become precarious in the face of increasing unpredictability in plant phenology and other environmental stochasticities resulting from climate change.

Degree of Dietary Specialization on Furanocoumarin-Containing Hostplants in a Newly Invasive Web Building Caterpillar.

The genus Depressaria (Lepidoptera: Depressariidae) mostly comprises specialist herbivores with varying capacity for detoxification of defensive phytochemistry. Depressaria depressana, a Eurasian moth recently introduced into North America, is a family-level specialist of the Apiaceae, whose hosts include more than a dozen species in multiple tribes; Depressaria radiella is a super-specialist of Eurasian origin that feeds exclusively on species in the genera Pastinaca and Heracleum throughout its native and introduced range. In eastern North America, it feeds upon Pastinaca sativa, an invasive European species, and Heracleum maximum, a native species. We determined whether differences in furanocoumarin metabolism exist between D. depressana and two isolated populations of D. radiella, feeding exclusively on either P. sativa or H. maximum. We also compared gravimetric estimates of feeding efficiency to assess D. depressana larval performance on different diets. Both populations of D. radiella metabolized furanocoumarins at a greater rate than D. depressana. Although there was no difference in rates of metabolism of linear furanocoumarins in the two populations of D. radiella, individuals collected from H. maximum metabolized angular furanocoumarins more rapidly. The gravimetric assessments of feeding efficiencies revealed that D. depressana exhibited highest efficiencies consuming Daucus carota; moreover, this species survived to pupation consuming fruits of Zizia aurea, an apiaceous species native to North America. Our preliminary phylogenetic analysis, building on an earlier morphological analysis, incorporates mitochondrial cytochrome oxidase subunit 1 data from the BOLD database and revealed that the presence or absence of furanocoumarins is not a strong predictor of species-level evolution in Depressaria.

Resource Availability, Sex, and Individual Differences in Exploration Drive Individual Diet Specialization.

AbstractIndividual diet specialization (IDS) is widespread and can affect the ecological and evolutionary dynamics of populations in significant ways. Extrinsic factors (e.g., food abundance) and individual variation in energetic needs, morphology, or physiology have been suggested as drivers of IDS. Behavioral traits like exploration and boldness can also impact foraging decisions, although their effects on IDS have not yet been investigated. Specifically, variation among individuals in exploratory behavior and their position along the exploration/exploitation trade-off may affect their foraging behavior, acquisition of food items, and home range size, which may in turn influence the diversity of their diet. Here, we analyzed stable carbon and nitrogen isotopes in hair of wild eastern chipmunks, Tamias striatus, to investigate the influence of individual differences in exploration on IDS. We found that exploration profile, sex, and yearly fluctuations in food availability explained differences in the degree of dietary specialization and in plasticity in stable carbon and stable nitrogen over time. Thus, consistent individual differences in exploration can be an important driver of within-population niche specialization and could therefore affect within-species competition. Our results highlight the need for a more thorough investigation of the mechanisms underlying the link between individual behavioral differences and diet specialization in wild animal populations.

Seasonality affects specialisation of a temperate forest herbivore community

Understanding spatiotemporal trends on insect‐plant interaction networks is essential to unveil the ecological and evolutionary processes driving herbivore specialisation. However, community studies accounting for temporal dynamics in host‐plant specialisation of herbivorous insects are surprisingly scarce. Here, we investigated how seasonality affects specialisation of a temperate forest herbivore community. A substantial body of literature suggests that young plant foliage tends to be more protected by defence mechanisms effective against generalist insect herbivores than mature leaves. We thus hypothesised that herbivore specialisation would be highest in the early season, when young leaves are available, and that the degree of dietary specialisation would decline with leaf maturation. To test this hypothesis, we sampled more than 4700 folivorous caterpillars associated with 16 tree species within a 0.2 ha deciduous forest stand in eastern North America. We further examined the dietary specialisation of exposed feeders versus shelter builders (e.g. leaf rollers, leaf tiers, webbers). Contrary to our prediction, we observed a significantly less specialised herbivore fauna in the early season than in the two subsequent summer seasons. We further found that the seasonal increase in specialisation was driven by a remarkable turnover in species composition rather than by shifts in guild structure or intraspecific changes in diet breadth of the herbivores. These findings run counter to the widespread belief that young leaves are less susceptible to generalist insect herbivores. Our study underscores the need to account for temporal dynamics when contrasting herbivore specialisation among sites with pronounced seasonality.

Can mandible morphology help predict feeding habits in Antarctic amphipods?

AbstractIn Antarctica, amphipods form a highly diverse group, occupy many different ecological niches and hold an important place in food webs. Here, we aimed to test whether differences in Antarctic amphipod feeding habits were reflected in their mandible morphology, and if mouthpart specialization could be used to describe amphipod trophic ecology. To do so, we compared mandible morphology in nine species spanning seven families and five functional groups (grazers, suspension feeders, generalist predators, specialist predators and scavengers). Mandible morphology adequately depicted some aspects of amphipod trophic ecology, such as the trophic level at which animals feed or their degree of dietary specialization. On the other hand, links between mandible morphology and amphipod diet were seldom unambiguous or straightforward. Similar adaptations were found in distinct functional groups. Conversely, mandible morphology could vary within a single functional group, and phylogenetic effects sometimes complicated the interpretation of form-function relationships. Overall, mandible morphology on its own was generally not sufficient to precisely predict amphipod feeding strategies. However, when combined with other methods (e.g. gut contents, trophic markers), it constitutes a valuable source of information for integrative studies of amphipod ecological diversity in the Southern Ocean.

Macroevolutionary analyses indicate that repeated adaptive shifts towards predatory diets affect functional diversity in Neotropical cichlids

AbstractDuring adaptive radiation, diversification within clades is limited by adaptation to the available ecological niches, and this may drive patterns of both trait and species diversity. However, adaptation to disparate niches may result in varied impacts on the timing, pattern and rate of morphological evolution. In this study, we examined the relationship between feeding ecology and functional diversification across a diverse clade of freshwater fishes, the Neotropical cichlids. Species dietary niches were ordinated via multivariate analysis of stomach content data. We investigated changes in the rate and pattern of morphological diversification associated with feeding, including dietary niche and degree of dietary specialization. A major division in dietary niche space was observed between predators that consume fish and macroinvertebrates vs. other groups with diets dominated by small invertebrates, detritus or vegetation. These trophic niches were strongly associated with groupings defined by functional morphospace. Clades within the piscivore/macroinvertivore group rarely transitioned to other dietary niches. Comparatively, high dietary specialization enhanced functional diversification, driving the evolution of more extreme morphologies. Divergent patterns of trophic diversification among Neotropical cichlids appear to derive from different performance demands in regional abiotic and biotic environments associated with biogeographical history.

Trophic interactions and isotopic niche of octopuses and sea stars in North Patagonia

Coexistence of species is associated with differentiation of niches and partitioning of resources, which lessen niche overlap and the intensity of competition. Here, we used stable isotope analysis to evaluate and compare the trophic relationships and isotopic niche of two coexisting predators, the small octopus Octopus tehuelchus (d’Orbigny 1834) and the brooding sea star Anasterias antarctica (Lutken 1857), in the rocky intertidal community of northern Patagonia (Argentina). Our results show differences in the proportional contribution of sources between octopuses and sea stars. Octopuses consume mainly sessile filter-feeders, and sea stars show intra-specific diet differences, changing its preference with size and sex. Small and medium sea stars feed on sessile filter-feeders, such as small mussels, while large sea stars consume more frequently grazer gastropods and scavenger crustaceans. In the food web of the rocky north Patagonian ecosystem, differences in the isotopic niche space and the degree of dietary specialization between octopuses and sea stars indicate differential use of resources and trophic niche partitioning, possibly by limiting inter- and intra-specific competition.

Frequency and consequences of individual dietary specialisation in a wide-ranging marine predator, the northern gannet

Individual specialisations in animals are important contributors to a wide range of ecological and evolutionary processes, and have been particularly documented in relation to multiple aspects of foraging behaviours. Central-place foragers, such as seabirds, frequently exhibit pronounced specialisations and individual differences in a variety of foraging traits. In particular, the availability of fisheries discards alongside natural prey resources provides additional potential for differentiation and specialisation for opportunistically scavenging seabird species. However, the consequences of such specialisations for at-sea distributions and intraspecific interactions are not well known. Here, we investigated the links between the degree of dietary specialisation on natural or discarded prey and the foraging movements and spatial occupancy of northern gannets Morus bassanus in relation to differing intraspecific competition at 6 colonies of differing sizes. We found that, at most colonies, individuals with different dietary strategies concentrated foraging at differing levels of intraspecific competition. In addition, individuals pursuing different strategies were frequently, but not consistently, spatially separated, distinctions that were most acutely seen in females. However, this variation in individual strategy had no significant impact on current body condition. These analyses demonstrate how foraging-associated metrics need not covary within an unconstrained system. They also reveal that specialisation can have important consequences for the competitive regimes individuals experience, highlighting the complexity of examining interacting consequences at large spatial scales.

Diet comparison between rainforest and cave populations of Craugastor alfredi (Anura: Craugastoridae): does diet vary in contrasting habitats?

ABSTRACTInformation about interpopulational variation of diet is important for thoroughly understanding the ecological niche of a species, the possible attributes that have allowed it to live in different habitats and ecosystems (if it is the case) and the degree of dietary specialization. This study compared dietary composition and diversity, size of prey consumed and feeding intensity between two geographically proximate populations of Craugastor alfredi, inhabiting contrasting habitats: one located in a remnant tropical rainforest and the other population located within a cave entrance, in eastern Mexico. A total of 118 frogs (50 from rainforest and 68 from cave) were stomach-flushed and, from those, 86 had at least one prey item (42 from rainforest and 44 from cave). Consumption proportions of orthopterans differed between habitats, with frogs from the cave having higher importance values of this prey than frogs from the rainforest. Additionally, both dietary diversity and feeding intensity were higher in the rainforest population. It was concluded that the generalized diet of C. alfredi, besides the high consumption of orthopterans by the cave population, may have contributed to its success in cave entrances. Nonetheless, other non-feeding attributes like small body size, direct development and semiarboreality may also facilitate the use of rock crevices in caves for foraging and reproduction.

Evolution of dentition in salamanders: relative roles of phylogeny and diet

Despite being widely regarded as generalist predators, amphibians exhibit a diversity of tooth shapes and dentition patterns, which may indicate the influence of dietary specialization on the evolution of tooth morphology. Very few studies have analysed the relationship between tooth morphology and diet (i.e., prey items) in amphibians, and those existing studies are highly speculative. We investigated the evolution of salamander teeth and the relationship between tooth morphology and diet in a phylogenetically independent fashion. We used a phylogeny of 23 species of salamander representing three families (Ambystomatidae, Plethodontidae, and Salamandridae) to, first, analyse the divergence of tooth morphology and its relationship to phylogeny and, second, to analyse the relationship between tooth morphology and diet diversity. We used electron scanning microscopy and a statistical comparative approach using Spatial Evolutionary and Ecological Analysis (SEEVA) and phylogenetic generalized least-squares regression in R. Our results indicated significant divergence in tooth morphology at major phylogenetic splits. Moreover, there was a significant, phylogenetically independent relationship between tooth morphology and diet diversity. The relationship between diet and tooth morphology indicates not only a reflection of phylogenetic history, but also a degree of dietary specialization, indicating that evolution in tooth morphology has had an adaptive aspect in relation to salamander diet.

Land cover influences dietary specialization of insectivorous bats globally

Land-use changes are a leading cause of biodiversity loss and ecosystem service degradation worldwide, but these changes do not affect all organisms equally. Understanding factors influencing resistance to environmental change is vital for informed conservation. In particular, dietary generalists may withstand environmental change better than specialists due to their ability to exploit variable resources. Bats are voracious insectivores, but vary widely in their degree of dietary specialization. Here, we analyze the effect of land cover and morphology on dietary diversity and the two most common prey items on which bats specialize (Lepidoptera and Coleoptera), selecting important independent variables using phylogenetic generalized least squares and model selection. Dietary diversity increased with increasing amount of cropland near the study area, consumption of Lepidoptera decreased with increasing habitat diversity, and consumption of Coleoptera decreased with increasing distance from the equator. Biodiversity (and hence, prey diversity) is expected to decrease with agricultural intensity, but the observed pattern suggests dietary specialists may avoid agricultural habitats due to lack of preferred prey. Dietary specialists may thus be increasingly at risk as agricultural intensity increases around the world, and it is essential that we continue to document their ecological roles and the services they provide to society.

Field and laboratory selection of brassicaceous plants that differentially affect infestation levels by Delia radicum

AbstractSeveral plant traits control plant–insect interactions and shape host range of herbivorous insects according to their degree of dietary specialization. Understanding how plant species diversity influences herbivore infestations is of interest for the development of alternative crop protection strategies. In a pest management context, an appropriate selection of plants can modify pest distribution at the field scale. To develop a ‘push–pull’ strategy against the cabbage root fly, Delia radicum, we conducted a field study to both determine which plants exhibit contrasted pest infestation levels and to evaluate their influence on egg predation activity. Our field experiment reveals that infestation levels of brassicaceous plants by the cabbage root fly in the field can vary considerably according to plant genotype and species, while the number of predated eggs is only slightly affected by plant species. Olfactometry studies carried out under laboratory conditions revealed that plants harbouring the highest number of eggs in the field were also highly attractive, suggesting that olfactory stimuli are responsible, at least partially, for the differential infestation levels observed in the field. In a ‘push–pull’ context, this study demonstrates that different brassicaceous plants could be used to redistribute cabbage root flies in broccoli crops without compromising herbivore control by natural enemies. In addition, the importance of plant volatiles for infestation levels suggests a potential for developing a semiochemically assisted ‘push–pull’ system in which trap plants would be enhanced by synthetic release of attractive compounds.

Dietary breadth of frugivorous birds in relation to their feeding strategies in the lowland forests of central Honshu, Japan

Variation in the dietary breadth of frugivorous birds, i.e. variation in the degree of dietary specialization, is a key factor for understanding the construction of the architecture of diffused fruit–bird interactions. However, little information is available on the dietary breadth of frugivorous birds in a community and the avian attributes influencing it. In this study, we evaluated and compared the dietary breadth of 23 frugivorous birds (including both seed dispersers and seed predators) in relation to their feeding strategies (depending on which the birds were categorized into four types as gulpers, grinders, crushers and peckers) and other avian attributes. In particular, we hypothesized that feeding types with long fruit‐handling time (i.e. crushers and peckers) have a narrow dietary breadth. Our analysis was based on the data obtained through long‐term volunteer monitoring in the lowland forests of Kanagawa prefecture, Japan. We evaluated the dietary breadth of the birds while controlling for differences in sampling efforts and seasonal variation in the available fruit assemblages among the birds. Our analysis revealed that there was a large variation in dietary breadth among the birds, even after controlling for differences in sampling efforts and seasonal variation in the available fruit assemblages. Feeding strategies of the birds defined their dietary breadth to some degree, but we found little support for the hypothesis that fruit handling determines dietary breadth. We also observed a large variation in dietary breadth among the gulpers, which act mainly as seed dispersers for plants, and this variation could not be explained fully by other avian attributes. Our results showed that variation in the dietary breadth of frugivorous birds is partly determined by their feeding strategies and suggested that other unknown factors may play a role in determining dietary breadth variation and in structuring fruit–bird interaction networks.

Testing the Threat-Sensitive Hypothesis with Predator Familiarity and Dietary Specificity

In a system with multiple predators, the threat-sensitive predator avoidance hypothesis predicts that prey respond differently to predators relative to the risks each poses (e.g., degree of dietary specialization). Aquatic animals often rely heavily on detecting predators via chemical cues (kairomones) and respond with a suite of behaviors including detection and avoidance. However, little is known about how animals respond to kairomones of specialist versus generalist predators. In laboratory experiments, we compared behavioral responses of a poorly studied aquatic salamander, the greater siren (Siren lacertina), to cues from specialist and generalist predator snakes to evaluate threat-sensitive responses. Sirens exhibited a novel behavior (gill-flushing) most often in the presence of specialist predator cues. Avoidance behavior (reversing direction following cue detection) was higher in response to specialist predator and novel animal control cues and lowest in response to generalist predator cues. An intermediate response to the animal control, an unfamiliar amphibian predator, indicated that sirens respond cautiously to a novel cue. The gradient of observed responses to different snake cues indicates that sirens may be evaluating predation potential of animals based on their foraging specificity and familiarity.

Generalists, specialists and opportunists: niche metrics of Poicephalus parrots in southern Africa

There are distinct differences in the way in which Poicephalus parrots utilise food resources within their distributional range. Our study evaluated the niche metrics of all Poicephalus parrots studied thus far for evidence of linkages between body mass, distributional range and niche breadth. We calibrated the degree of dietary specialisation of Meyer's Parrots in the Okavango Delta, Botswana, over 12 months using a modified Hurlbert's expanded and standardised niche breadth index, and for comparison using Levins's niche breadth, with the food item calendars of the other Poicephalus parrots in southern Africa. Our study showed that Meyer's Parrot has the widest niche breadth of any Poicephalus parrot studied thus far, which likely plays a role in their wide distributional range. In addition, degree of dietary specialisation was inversely correlated with body mass, whereby the narrower the trophic niche, the larger the body size.

Relationship between demographics and diet specificity of Imperial Eagles Aquila heliaca in Kazakhstan

The demographic consequences of within‐population variability in predator foraging are not well understood. We assessed the relationship between the degree of diet specialization and two demographic parameters, population density and reproductive output, within a single population of Imperial Eagles Aquila heliaca at a nature reserve in north‐central Kazakhstan. Nearest‐neighbour distances between eagle nests throughout the reserve, and thus population density, were correlated with the degree to which diets were specialized. Diet diversity showed an extensive regional variability that was linked to prey distributions, but within‐year analyses of reproductive output did not show similar linkages. However, multi‐year analyses of breeding performance showed inter‐regional differences in nesting success that were paralleled, and probably driven by, similar trends in diet diversity. In contrast, brood size at fledging was not linked to diet diversity and was more probably driven by reserve‐wide influences such as weather. Finally, the decision to initiate breeding was associated neither with diet diversity nor with environmental variability. Our results indicate that the degree of dietary specialization is linked to the demographics of Imperial Eagle populations. For these and other raptor populations, it is possible that management could be used separately to increase or decrease nesting success, brood size at fledging, and the likelihood that a pair will initiate breeding.

Evolution of Dietary Specialization and Chemical Defense in Poison Frogs (Dendrobatidae): A Comparative Analysis

Defensive mechanisms, including noxious or toxic substances, are favored by predation-driven natural selection. The acquisition of noxious/toxic substances can be either endogenous, in which the substances are produced by the organism, or exogenous, in which the substances are produced by another organism and are sequestered. Evidence indicates that the defensive skin alkaloids of Neotropical poison frogs (Dendrobatidae) have an exogenous source: a diet of ants and other small alkaloid-containing arthropods, which we term the diet-toxicity hypothesis. A critical prediction of the diet-toxicity hypothesis is that independent origins of dietary specialization will be found to be correlated with independent origins of skin alkaloids. We tested this prediction in an integrated framework using comparative methods with new and published data on feeding ecology and chemical defense for 15 species of dendrobatids in five genera. We found a significant correlation between alkaloid profiles and degree of dietary specialization. This reveals a recurring association of dietary specialization and alkaloid sequestration in dendrobatids, which suggests parallel evolutionary trends in the origins of defensive mechanisms.

Phylogenetic Relationships, Species Limits, and Host Specificity of Gall-Forming <I>Fergusonina</I> Flies (Diptera: Fergusoninidae) Feeding on <I>Melaleuca</I> (Myrtaceae)

Phylogenetic analysis of recently described gall-forming Fergusonina Malloch flies was performed using DNA sequence data from the mitochondrial cytochrome oxidase I gene. Fifty-three flies reared from nine species of Melaleuca L. were sequenced. Species boundaries delimited by mitochondrial data confirm recent morphological investigation with one exception. Fergusonina turneri Taylor, believed to feed on both Melaleuca quinquenervia (Cav.) S. T. Blake and Melaleuca fluviatilis Barlow, seems to be comprised of two cryptic species, each specialized on one of the two hosts. Because F. turneri is under consideration as a potential biological control agent for invasive M. quinquenervia in the Florida Everglades, understanding cryptic variation and the degree of dietary specialization of this species is critical.

Host specificity and comparative foraging behaviour of Aenasius vexans and Acerophagus coccois, two endo‐parasitoids of the cassava mealybug

AbstractTwo encyrtid parasitoids, Aenasius vexans Kerrich (Hymenoptera: Encyrtidae) and Acerophagus coccois Smith (Hymenoptera: Encyrtidae), were compared for their degree of dietary specialisation and the impact this has on their foraging strategies. Both parasitoid species are significant for biological control of the cassava mealybug, Phenacoccus herreni, Cox & Williams (Homoptera: Sternorrhyncha) a major Latin American pest of cassava, Manihot esculenta Crantz, an important root crop. Host acceptance and parasitism were analysed in seven mealybug species (with different levels of polyphagy) occurring in and around cassava fields. Results demonstrate that, in this ecosystem, An. vexans is a specialist for P. herreni while Ac. coccois is a generalist on the first and second trophic level. Of the seven mealybug species, P. herreni and P. madeirensis Green were the most acceptable hosts for Ac. coccois, followed by Ferrisia virgata Cockerell. Ac. coccois did not accept the other four mealybug species.The foraging and oviposition behaviour of individual parasitoids was observed in bioassays with cassava leaves infested by P. herreni. The two species used different strategies to locate their host. Aenasius vexans spent significantly more time walking and standing on an infested leaf and examined a host longer than did Ac. coccois. Acerophagus coccois, in contrast, spent more time for oviposition. As a consequence An. vexans parasitised more hosts in a given time than did Ac. coccois. Because the rate of offspring production of the two species did not differ, we conclude that the gregarious Ac. coccois's strategy to deposit several eggs at once might compensate for its relatively low number of ovipostitions, compared with the solitary An. vexans. These findings suggest that, given the advantages and limitations of each species, a multi‐species approach to biological control of P. herreni may yield best results.