Competition between Eurasian otter Lutra lutra and American mink Mustela vison probed by niche shift

Interspecific competition is one of several constraints that might prevent an individual from maximising its energy intake. When an interspecific competitor is introduced, an individual is often forced to shift its diet according to the intensity of the competitive pressure. In this paper, we explore whether the introduced American mink ( Mustela vison Schreber) shifts its diet when the density of its potential competitor, the Eurasian otter ( Lutra lutra L.), is increased. We compared the diets of otter and mink at the same location but at two moments in time when the relative densities of these two species were different while controlling for the abundance of aquatic prey. Mink and otters are semi-aquatic mammals belonging to the same guild of mustelids and otters are expected to be the dominant competitor because they are larger and better at hunting underwater. The diets of otters and mink overlap to a great extent but while otters specialise mainly on aquatic prey, mink are able to exploit both aquatic and terrestrial prey. These observations prompted the hypothesis investigated in this work that at higher otter densities the diet of mink should change to include a higher proportion of terrestrial items. This hypothesis was supported by the data and at higher otter densities mink diet was observed to consist of a higher proportion of mammals and birds while fewer fish were present, although this pattern was present only in winter while no changes were observed in spring. Meanwhile the diet of otters remained basically unchanged. In the second part of the study, we investigated whether niche breadth and niche overlap between otter and mink changed at different otter densities. We found that niche overlap declined as the density of otters increased, in agreement with the prediction of habitat selection theory.     

Understanding the inter-specific dynamics of two co-existing predators in the Tierra del Fuego Archipelago: the native southern river otter and the exotic American mink

Knowledge about interactions between endangered native southern river otters (Lontra provocax) and introduced American mink (Neovison vison) is essential for effective management of both species. We evaluated competition for spatial and trophic niches between otter and mink in overlapping and non-overlapping areas, comparing distribution, habitat preference, diet and mink marking behavior. We surveyed otter and mink signs along 250 km of Beagle Channel coastline. Habitat suitability models were constructed based on species presence/absence and habitat characteristics, using generalized linear models. Feces were collected for diet analyses. Otters used forested coasts with 12°–32° shoreline slope and without human influence, and our evidence suggests they were not affected by mink presence. Mink preferred forested and shrubland coasts with 10°–28° shoreline slope. Neither human influence nor otter presence affected mink habitat occupation, but in the presence of otters, mink left fewer signs. Otters consumed more aquatic prey than mink, and mink modified their diet in the presence of otters, consuming more exotic small terrestrial mammals and less fish as well as shifting to smaller and shallower fish species that are less consumed by otters. Mink showed more plastic, generalist behavior than otters, being more tolerant of human presence, using more habitat types and having greater diet breadth. At the same time, otters apparently affect mink adversely and could help limit their invasion in sympatric areas. Conservation and recovery of otters, therefore, may produce a secondary benefit of simultaneously reducing the effect of mink, thereby providing an additional way to control this exotic predator’s population.     

Evidence for food competition between mink (Mustela vison) and otter (Lutra lutra) on Scottish islands

Food competition between American mink and otters was measured by comparing the diets of sympatric mink and otter populations with those of allopatric populations. Niche breadth was narrower for otters than mink. Niche breadth was wider for both mink and otters on islands where they co-existed in comparison to that of the allopatric populations. Niche overlap was lower in sympatric populations on islands with mammalian prey, however, niche overlap was not reduced on small islands without mammalian prey. Obtained data suggest that mink and otter compete for food resources and, when alternative prey sources are available, mink become more generalist predators to avoid competition with otters. However, when alternative prey sources are not available, both species become more generalist.     

Interactions of exotic and native carnivores in an ecotone, the coast of the Beagle Channel, Argentina

In coasts bordering highly productive seas, there can be a flux of resources to the terrestrial ecosystem, and terrestrial carnivores can use marine prey extensively. Two native, endangered species (otter Lontra provocax and culpeo fox Pseudalopex culpaeus lycoides) and two exotic species (mink Neovison vison and grey fox Pseudalopex griseus) inhabit the Beagle coast. The objectives of this paper are: (1) to describe the diet, habitat use and distribution of otters, mink and foxes on the coast of the Beagle channel and (2) to discuss the role of marine resources in the ecological interactions among these species. Diet was determined from the analysis of 245 faeces, and distribution was established from sign surveys. Marine prey occurred in the scats of 98.3, 70.4, 35.5 and 18.2% of otters, mink, culpeo and grey foxes, respectively. Other terrestrial species also use marine resources in Southern Patagonia. All this evidence suggests that the Fueguian coastal channels provide an illuminating example of allochthonous food subsidies from the sea. In the community of four sympatric predators, two native (and endangered) and two exotic, coexistence appears to be facilitated by a renewing marine food subsidy.     

Impact of released Eurasian otters on a population of American mink: a test using an experimental approach

Competition is considered to be a major evolutionary driving force within assemblages of related and morphologically similar species, but it is notoriously difficult to test and quantify especially amongst species that cannot be easily manipulated in a laboratory or in the field. By exploiting a re-introduction of Eurasian otters ( Lutra lutra ) in the Upper Thames catchment (UK) in 1999 we performed an experiment to test whether there was evidence of competition between otters and American mink ( Mustela vison ). Mink and otters are semi-aquatic mustelids belonging to the same guild. Otters are expected to be the dominant competitor because they are larger and better adapted at exploiting aquatic resources. Our hypothesis was that mink declines when the density of otters increases. We measured the effect of competition at the population level, by observing whether mink distribution and densities changed in an area of 1353 km² in association with the arrival of its putative competitor. We estimated distribution and densities by means of sign surveys and trapping. The results showed that otters were associated with a significant and rapid reduction in the densities of mink, while mink occupancy remained approximately the same in an area of 2464 km² that was used as a control. We observed that the spatial distribution of the mink population throughout its decline was influenced by the yearly cycle of mink activities, with areas being temporarily re-colonized during the dispersal season. Mink is an invasive species in the UK threatening the survival of some native species. Our findings suggest that the re-establishment of otter populations is likely to lead to a decline of mink that may, in turn, be beneficial to native species threatened by this invasive.     

Decline of invasive alien mink (Mustela vison) is concurrent with recovery of native otters (Lutra lutra)

Invasive species often cause the decline of native prey or competitors. We highlight a contrasting example of the large-scale recovery of a native species and the concurrent decline and likely displacement of an established invasive competitor. Invasive American mink Mustela vison became widespread in the British Isles at the same time as native Eurasian otters Lutra lutra were declining as a result of water pollution. In common with other invasive predators, mink cause conservation problems for a range of native prey species, most notably water voles Arvicola terrestris . Recent trends in the distribution of native otters and invasive mink in north-east England were examined using a novel regression modelling approach to analyse presence/absence data from field surveys, corroborated by contemporary predator culling records. Between 1991 and 2002, the percentage of sites where mink signs were found decreased from 80% to 20%, while otter signs increased from 18% to 80%. Annual indices of mink captures on shooting estates increased between 1980 and 1996, but were followed by a decline thereafter. Indices of the incidence of native otters were significantly related to those indicating the decline or displacement of invasive mink. This large-scale field study is supportive of localized experimental evidence for the return of dominant, native otters being concurrent with the decline of the invasive alien mink. The recovery of a dominant native species may represent a reversal of the mesopredator release that allowed invasive mink to establish and may eventually serve to mitigate their impact on native prey species.     

Diet of coastal foraging Eurasian otters (<Emphasis Type="Italic">Lutra lutra</Emphasis> L.) in Pembrokeshire south-west Wales

The importance of the marine environment to Eurasian otters is currently poorly understood. Wales is one of the few countries where coastal activity has been recorded and an increase in marine otter sightings could indicate remarkable developments within Welsh populations. The trophic niche of coastal otter populations around Pembrokeshire was investigated over a 12-month period. Marine activity was more widespread than previously thought and marine prey formed the largest component of otter diet, although, otters also consumed freshwater and terrestrial prey throughout the year. Otter diet was very diverse compared to other European coastal populations and a spring contraction in trophic niche width coincided with the estimated timing of breeding activity. Seasonal variation in prey composition was predominantly due to differences in the consumption of alternate prey types. In areas where wetlands are fragmented and populations of freshwater fish are declining, the marine environment may become an increasingly important habitat for otters. It is necessary to define the historical importance of coastal populations to otter conservation. Coastal areas are often subject to pressure from human activities, so the impact of disturbance needs to be assessed. Importantly, there is no verified otter survey method for coastal areas, so the use of marine habitat is likely to be underestimated.     

Coexistence of Habitat Specialists and Generalists in Metapopulation Models of Multiple-Habitat Landscapes

In coarse-grained environments specialists are generally predicted to dominate. Empirically, however, coexistence with generalists is often observed. We present a simple, but previously unrecognized, mechanism for coexistence of a habitat generalist and a number of habitat specialist species. In our model all species have a metapopulation structure in a landscape consisting of patches of different habitat types, governed by local extinction and colonization. Each specialist is limited to its specific type of habitat. The generalist can use more types of habitat, has a lower local competitive ability but can exploit patches left open by the specialists. Our modeling shows that coexistence is easily possible. The mechanism amounts to a colonization/competition trade-off at the landscape level, where the colonization advantage of the inferior competitor does not arise from a higher colonization rate but from its ability to use more types of habitat. Habitat availability has to be intermediate: when there are few patches of each habitat, only the generalist is able to maintain itself and when there are many patches, high propagule pressure of the specialists excludes the generalist. Habitat selection or temporal variations in relative habitat quality are not necessary for coexistence. Increased niche-width, colonization rate or local competitive ability of the generalist enhances its performance compared to the specialists. Various types of habitat degradation favour generalism. When able to use a broad range of habitats, generalists can generate so much propagule pressure that only a low level of local competitive ability is needed to globally exclude the specialists. Hence, in a reversal of the original problem, the question is why there are so many specialist metapopulations?     

Do alien North American mink compete for resources with native South American river otter in Argentinean Patagonia?

American mink Mustela vison , originally bred in fur farms, have become established in areas occupied by native endangered Southern river otter Lontra provocax , in Patagonia. In accordance with European experience, this biological invasion in South America raises questions about the interaction between invasive mink and native otter, from the viewpoints of both community assembly and conservation. We set out (1) to find which aspects of habitat structure were related to the distribution of signs of both this invasive species and Southern river otter Lontra provocax , in Argentinean Patagonia and their most common prey and (2) to test general predictions of niche partitioning between these two species. Based on surveys of 447 of 600 m transects for otter and mink scats/footprints along the waterside of lakes and rivers in the Andean Patagonian region, we compared diet composition (from scat analysis) and micro-habitat preferences (from field signs) of the two species. Otters were more specialist than mink in habitat use and diet. Mink used different habitats in other river basins where otters were absent. Where they occurred together in the basin of the Limay River, the distributions of their signs were similar, and mink diet was more similar to that of otters. There was no detectable difference in otter diet before and after mink arrival in the Limay basin. Contrary to the prediction of niche partitioning, and to the findings of European studies, resource use by mink was more similar to that of otters where the species occurred sympatrically than where they were allopatric.     

The Smell of New Competitors: The Response of American Mink, Mustela vison, to the Odours of Otter, Lutra lutra and Polecat, M. putorius

We tested the response of wild American mink (an established alien species in the UK), to the odours of unfamiliar mink, European polecat and Eurasian otter. Polecats are similar in size and habits to mink, otters are larger than mink and a dominant competitor; both are native to the UK and both were absent during the original colonization by mink but are now undergoing natural population recoveries. The response of mink to experimental odours was assessed by counting the numbers of tracks (footprints) on rafts treated with anal gland secretions, and compared with response to a control raft, on two rivers in the Upper Thames valley, UK. Remote video showed that the number of tracks was positively correlated with the time that mink spent investigating an odour. We found that mink were attracted to the odours of both unfamiliar mink and polecats. There was little evidence that mink avoided the odour of otters. We suggest that, during an encounter with a polecat, mink may behave much as they would to a conspecific. We infer from the response of mink to the odour of otters, that, if mink do avoid otters, the mechanism of avoidance is likely to be complex, situation-dependent and perhaps affected by prior experience.     

Feeding habits of the otter and the American mink in Bialowieza Primeval Forest (Poland) compared to other Eurasian populations

Diets of the otter Lutra lutra and the American mink Mustela vison were studied by scat analysis on five woodland rivers and streams in eastern Poland. Fish constituted 51% of food biomass consumed by otters in spring‐summer and 40% in autumn‐winter, with common fish (perch Perca fluviatilis, pike Esox lucius, and roach Rutilus rutilus) being captured most frequently by the otters. Amphibians (mainly Rana temporaria, which also dominated in the living community) made up 34% of otters’ food biomass in spring‐summer and 58% in autumn‐winter. American mink relied on three prey groups: fish (40% in spring‐summer, and 10% in autumn‐winter), frogs (32% and 51%, respectively), and small mammals (21% and 36%). Out of available Micromammalia, mink strongly selected the root vole Microtus oeconomus. The cold season diet of both otter and mink depended on river size. On small rivers with forested valleys, otters and mink fed nearly exclusively on amphibians (72–90% of food biomass). With size of a river increasing and riverside habitat becoming more open (sedge and reed marshes instead of forests), otters shifted to catching predominantly fish (up to 76% in diet) and mink to preying on small mammals (up to 65% in diet).
Review of literature on otter and mink in Eurasia showed that their diets did not change with latitude (as indicators of climate severity and duration of water freezing) but they depended on habitats. In otter diet, the mean share of fish declined from 94% (SE 1.7) on sea shores, to 71% (SE 2.9) on lakes and fish ponds, to 64% (SE 2.8) on rivers and streams. The roles of amphibians and crustaceans increased in the same gradient (from 0 to 15%, and from 3 to 7%, respectively). On inland waters, the abundance of crayfish was the essential factor differentiating otters’ diet composition. In Eurasia, the staple food types of American mink on rivers and streams were fish (on average, 27% in diet, SE 3.9), mammals (30%, SE 5.0), and amphibians (17%, SE 4.8), whereas on lakes and ponds mink fed predominantly on birds (on average, 33% in diet, SE 10.1) and fish (28%, SE 9.5). In the Palaearctic region, over a wide gradient of habitats, otters appeared strongly specialised on prey taken from water, whereas American mink was a typical generalist capable of utilising several prey groups originating from both water and land.     

Neo-tropical felid activity patterns in relation to potential prey and intraguild competitors in the Calakmul Biosphere Reserve,Mexico

Predator behaviors influence, and are influenced by, prey and competitor behaviors. Jaguars (Panthera onca), pumas (Puma concolor), and ocelots (Leopardus pardalis) coexist throughout their geographic range as the three largest predators in a multi-predator community across diverse environments. This study tested for non-random segregation and overlap in the activity patterns of these felids and their shared prey in the southern buffer zone of the Calakmul Biosphere Reserve, in southern Mexico, using camera traps during February to August 2019. We detected little temporal segregation between the nocturnal activities of jaguars, pumas, and ocelots, although pumas were more active closer to dawn. Jaguars had low activity overlap with species likely to be common prey, whereas ocelots had high overlap with their potential prey. Pumas displayed finer-scale similarities in activity with species likely to be common prey. In an understudied area of conservation importance, this study shows that temporal segregation is an unlikely mechanism of coexistence. Further research should incorporate spatio-temporal avoidance and dietary differences to improve our understanding of the mechanisms that drive coexistence between generalist species in a diverse assemblage of threatened felids. Abstract in Spanish is available with online material.     

The dynamics of predation and competition in patchy environments

The model of N. D. Atkinson and B. Shorrocks (J. Anim. Ecol. 50, 461–471 (1981)) as two competing species distributing their progeny amongst patches according to independent negative binomial distributions. The resulting separation of the species increases the likelihood of coexistence. We have assumed a much simpler distribution of the competitors which has enabled us to explore analytically the dynamics of interactions with two competing species and a shared natural enemy in a patchy environment. Two types of natural enemy have been considered: a generalist predator whose dynamics are uncoupled from those of the two prey species, and a specialist (e.g., a parasitoid) whose dynamics are entirely coupled to those of its two prey. The following conclusions emerge. Non-aggregating generalist predators causing random predation across patches are generally destabilizing (although asymmetrical predation may in some case enhance coexistence as a result of preferential predation on the superior competitor). Predator aggregation in patches of high prey density, however, produces a switching effect which tends to promote stability. Coexistence is now even possible with high degrees of correlation in the distribution of the two prey and in situations of extreme competition where the competition coefficients exceed one. The main difference in the models with a specialist parasitoid as the natural enemy is a reduction in stability compared with the equivalent generalist-prey interaction. But stable coexistence can still readily occur if the natural enemies aggregate markedly in patches of high prey density.     

The influence of wave exposure on the foraging activity of marine otter, <Emphasis Type="Italic">Lontra felina</Emphasis> (Molina, 1782) (Carnivora: Mustelidae) in northern Chile

The marine otter Lontra felina has been said to prefer wave-exposed habitats over more protected sites in response to a greater prey abundance in exposed habitat. We examined how the foraging activity of L. felina is affected by the regime of wave exposure and prey availability at Isla Choros, northern Chile. Through focal sampling we recorded time spent by otters in foraging, the duration of dives, and the hunting success on a wave-exposed and a wave-protected site on the island. In addition, we quantified the abundance of prey in both habitats. Marine otters spent more time foraging in the wave-protected site compared with the wave-exposed habitat. Successful dives reached 26.9% in the wave-exposed habitats, and 38.2% in the wave-protected habitat. Foraging dives were 18% shorter in wave-exposed as compared with wave-protected habitat. Numerically, available prey did not differ significantly with habitat. Our results are more consistent with the hypothesis that wave-exposed habitats represent a sub-optimal habitat to foraging marine otters. Marine otters’ use of wave-exposed patches through northern and central Chile coastal areas probably reflects a low availability of suitable protected areas and greater human disturbance of more protected habitat.     

A Comparison of the diet and distribution of southern river otter (Lutra provocax) and mink (Mustela vison) in Southern Chile

In Chile, between latitudes 39S and 4330'S, southern river otter ( Lutra provocax ) and introduced mink ( Mustela vison ) scats were collected. Mink sign was recorded in 29% of the aquatic habitat where otter sign was found. Sixty-eight per cent of mink scats were collected at otter rest sites. A significant difference between the diets and low percentage of habitat use overlap of the two species (5-22%) suggest that, in Chile, river otter and mink coexist with little competition for space or food. There are no data supporting a relationship between the introduction of mink and the decline of southern river otters.     

Variation in the degree of specialization can maintain local diversity in model communities

We hypothesize that the continuum between generalist and specialist adaptations is an important general trade-off axis in the maintenance of local diversity, and we explore this idea with a simple model in which there are patch types to which species arrive as propagules and compete. Each patch type is defined by a competitive ranking of all species. A highly specialist species is the top competitor in one patch type but has a relatively low average ranking across different patch types, while a generalist species has a high average rank across patch types but is not the top competitor in any patch type. We use random dispersal and vary the fecundity of all species together to vary total propagule density and therefore recruitment limitation and density-dependent mortality. When fecundity is very high, each patch becomes occupied by its specialist species and generalists go extinct, so the number of species at equilibrium is equal to the number of patch types. If fecundity is very low, generalists dominate and specialists go extinct. There is a range of fecundity levels in which specialists, generalists, and intermediates coexist, and the number of species is substantially greater than the number of patch types. While coexistence of specialists and generalists has been considered a problem in evolutionary ecology, our results suggest to the contrary that this trade-off contributes to the maintenance of local diversity.     

Jack of all trades, master of all: a positive association between habitat niche breadth and foraging performance in pit-building antlion larvae

Species utilizing a wide range of resources are intuitively expected to be less efficient in exploiting each resource type compared to species which have developed an optimal phenotype for utilizing only one or a few resources. We report here the results of an empirical study whose aim was to test for a negative association between habitat niche breadth and foraging performance. As a model system to address this question, we used two highly abundant species of pit-building antlions varying in their habitat niche breadth: the habitat generalist Myrmeleon hyalinus, which inhabits a variety of soil types but occurs mainly in sandy soils, and the habitat specialist Cueta lineosa, which is restricted to light soils such as loess. Both species were able to discriminate between the two soils, with each showing a distinct and higher preference to the soil type providing higher prey capture success and characterizing its primary habitat-of-origin. As expected, only small differences in the foraging performances of the habitat generalist were evident between the two soils, while the performance of the habitat specialist was markedly reduced in the alternative sandy soil. Remarkably, in both soil types, the habitat generalist constructed pits and responded to prey faster than the habitat specialist, at least under the temperature range of this study. Furthermore, prey capture success of the habitat generalist was higher than that of the habitat specialist irrespective of the soil type or prey ant species encountered, implying a positive association between habitat niche-breadth and foraging performance. Alternatively, C. lineosa specialization to light soils does not necessarily confer upon its superiority in utilizing such habitats. We thus suggest that habitat specialization in C. lineosa is either an evolutionary dead-end, or, more likely, that this species' superiority in light soils can only be evident when considering additional niche axes.     

Trophic ecology of a top predator colonizing the southern extreme of South America: Feeding habits of invasive American mink (Neovison vison) in Tierra del Fuego

The American mink (Neovison vison) is a semi-aquatic, generalist carnivore released onto Tierra del Fuego (TDF) Island in the 1940s, subsequently spreading to adjacent islands in the archipelago with potential effects on native prey populations. Knowledge of this new predator's trophic ecology is essential to identify threats, plan control strategies and conserve native fauna. We studied seasonal mink diet in TDF in different habitats. We identified undigested remains from 493 scats collected between May 2005 and March 2009 along marine coasts and freshwater shores (rivers and lakes). Small mammals and fish were the main mink prey in TDF (over 65% of diet items). Seasonal variations were not detected, but diet did vary significantly between marine and freshwater habitats, where more terrestrial items were consumed. Among mammals, mink consumed more small native rodents than exotic species. Native fish consumption was also important with greater representation of species from the families Nototheniidae and Galaxiidae in marine and freshwater habitats respectively. Birds were the third item in importance, but did not constitute a particularly large part of the mink's diet on TDF. Overall, differences found in mink diet between habitats reflected their generalist/opportunistic feeding behaviour and did not differ greatly from observations in its native range or in other areas where it has been introduced. Our results establish the interactions between this novel predator and its prey and also illustrate the need to continue research on native prey populations to quantify mink impact on them and understand the ecological context of this biotic assemblage.     

Dynamics of a intraguild predation model with generalist or specialist predator

Intraguild predation (IGP) is a combination of competition and predation which is the most basic system in food webs that contains three species where two species that are involved in a predator/prey relationship are also competing for a shared resource or prey. We formulate two intraguild predation (IGP: resource, IG prey and IG predator) models: one has generalist predator while the other one has specialist predator. Both models have Holling-Type I functional response between resource-IG prey and resource-IG predator; Holling-Type III functional response between IG prey and IG predator. We provide sufficient conditions of the persistence and extinction of all possible scenarios for these two models, which give us a complete picture on their global dynamics. In addition, we show that both IGP models can have multiple interior equilibria under certain parameters range. These analytical results indicate that IGP model with generalist predator has “top down” regulation by comparing to IGP model with specialist predator. Our analysis and numerical simulations suggest that: (1) Both IGP models can have multiple attractors with complicated dynamical patterns; (2) Only IGP model with specialist predator can have both boundary attractor and interior attractor, i.e., whether the system has the extinction of one species or the coexistence of three species depending on initial conditions; (3) IGP model with generalist predator is prone to have coexistence of three species.     

Asymmetric reproductive interference between specialist and generalist predatory ladybirds

1.?Closely related species often differ greatly in the quality and breadth of resources exploited, but the actual mechanisms causing these differences are poorly understood. Because in the laboratory specialized species often survive and perform as well or better on host species that are never utilized in nature, negative ecological interactions restricting host range must exist. Here, we focused on reproductive interference, which has been theoretically predicted to drive niche separation between closely related species with overlapping mating signals. 2.?We examined the interspecific sexual interactions in relation to ecological specialization and generalization in two sibling ladybird species, Harmonia yedoensis and Harmonia axyridis. Harmonia yedoensis is a specialist predator that preys only on pine aphids, which are highly elusive prey for ladybird hatchlings, whereas H.?axyridis is a generalist predator with a broad prey and habitat range. 3.?We experimentally showed that conspecific sperm fertilized the vast majority of eggs regardless of mating order (i.e. conspecific sperm precedence) when a female of H.?yedoensis or H.?axyridis mated with both a conspecific and a heterospecific male. Moreover, we demonstrated that mating opportunities of H.?yedoensis females strongly decreased as heterospecific density increased relative to conspecific density. In contrast, in H.?axyridis, female mating success was high regardless of conspecific or heterospecific density. 4.?Our results suggest that the generalist H.?axyridis should be dominant to the specialist H.?yedoensis in terms of reproductive interference. Our results support the hypothesis that asymmetric reproductive interference from the dominant species may force the non-dominant species to become a specialist predator that exclusively utilizes less preferred prey in nature.