Is coarse woody debris in lakes a refuge or a trap for benthic invertebrates exposed to fish predation?

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Small but Powerful: Top Predator Local Extinction Affects Ecosystem Structure and Function in an Intermittent Stream

Top predator loss is a major global problem, with a current trend in biodiversity loss towards high trophic levels that modifies most ecosystems worldwide. Most research in this area is focused on large-bodied predators, despite the high extinction risk of small-bodied freshwater fish that often act as apex consumers. Consequently, it remains unknown if intermittent streams are affected by the consequences of top-predators’ extirpations. The aim of our research was to determine how this global problem affects intermittent streams and, in particular, if the loss of a small-bodied top predator (1) leads to a ‘mesopredator release’, affects primary consumers and changes whole community structures, and (2) triggers a cascade effect modifying the ecosystem function. To address these questions, we studied the top-down effects of a small endangered fish species, Barbus meridionalis (the Mediterranean barbel), conducting an enclosure/exclosure mesocosm experiment in an intermittent stream where B. meridionalis became locally extinct following a wildfire. We found that top predator absence led to ‘mesopredator release’, and also to ‘prey release’ despite intraguild predation, which contrasts with traditional food web theory. In addition, B. meridionalis extirpation changed whole macroinvertebrate community composition and increased total macroinvertebrate density. Regarding ecosystem function, periphyton primary production decreased in apex consumer absence. In this study, the apex consumer was functionally irreplaceable; its local extinction led to the loss of an important functional role that resulted in major changes to the ecosystem’s structure and function. This study evidences that intermittent streams can be affected by the consequences of apex consumers’ extinctions, and that the loss of small-bodied top predators can lead to large ecosystem changes. We recommend the reintroduction of small-bodied apex consumers to systems where they have been extirpated, to restore ecosystem structure and function.     

Fish predation affects the structure of a benthic community

1. We conducted an experimental study of predation by benthivorous fish on a natural community of stream invertebrates using a reach‐scale approach. Over a 2‐year period (experimental phase), the benthic invertebrate community of a stretch containing two species of benthivorous fish was compared with a fishless stretch. Thereafter, all fish were removed and benthic community structure was analysed again to account for natural differences between the two stretches (reference phase). 2. Benthivorous fish at the moderate densities investigated did not affect total benthic biomass or density, but did alter species composition. In addition, the fish effect differed between pool and riffle habitats, with larger effects in the pools indicating a habitat‐specific predation effect. In the reference phase, when all fish were removed from the stream, the difference between the two stretches was reduced. 3. The benthivorous fish reduced the densities of four taxa (Pisidium sp., Dugesia gonocephala, Gammarus pulex, Limoniidae), representing 29% of total biomass. It is possible that density reductions of other species were masked by prey migration despite the relatively large spatial scale. Indeed, higher drift activity in the upstream fishless stretch could have increased the density of Baetis rhodani in the fish stretch, as indicated by the results of a drift model. 4. Our results provide insights into stream food web ecology because fish predation showed effects even in a natural system where habitat complexity was high, environmental factors were highly variable and many predator and prey species interacted and because benthivorous fish were the focus, whereas the majority of previous predation experiments in streams have used drift‐feeding trout.     

Habitat overlap between predatory benthic fish and their invertebrate prey in streams: the relative influence of spatial and temporal factors on predation risk

1. The spatial heterogeneity of ecosystems as well as temporal activity patterns of organisms can have far‐reaching effects on predator–prey relationships. We hypothesised that spatiotemporal constraints in mesohabitat use by benthic fish predators would reduce habitat overlap with benthic invertebrates and lead to mesohabitat‐specific predation risks. 2. We analysed the spatiotemporal activity patterns of two small‐bodied benthivorous fishes, gudgeon (Gobio gobio) and stone loach (Barbatula barbatula), and of benthic invertebrates in a small temperate stream during three 24‐h field experiments. By applying a novel method of field video observation, we monitored the spatiotemporal foraging behaviour of the fish in their natural environment. A parallel analysis of invertebrate mesohabitat use by means of small area Hess sampling allowed a direct estimation of habitat overlap at a pool–riffle scale. 3. Gudgeon showed a dominant spatial activity pattern preferring pools at all times of day, whereas stone loach used both mesohabitats but with a distinct temporal (nocturnal) activity pattern. The patterns of residence were not identical with those of active foraging. Invertebrate community composition differed significantly between mesohabitats but not between times of day. More than half of the total dissimilarity between pools and riffles was accounted for by six invertebrate taxa. Five of these were subject to higher fish predation in pools than in riffles. The total prey consumption of the two fish species together in pools was about three times as high as in riffles. Trophic niche breadth of stone loach and thus its predation range was broader than that of gudgeon. 4. These results indicate that the potential predation risk for stream invertebrates depends on the combination of spatial and temporal patterns of both predator and prey. Given the distinct differences in predation risk found between pools and riffles, we conclude that spatial heterogeneity at the mesohabitat scale can influence mechanisms and consequences of selective predation. We also suggest that the analysis of spatiotemporal predator–prey relationships should not be based on the premise that the main residence habitat and active foraging habitat of a predator are identical.     

Stream inflow and predation risk affect littoral habitat selection by benthic fish

1. We examined small, fishless headwater streams to determine whether transport of macroinvertebrates into the littoral zone of an oligotrophic lake augmented food availability for Cottus asper, an abundant predatory fish in our study system. We sampled fish and macroinvertebrates during the recruitment and growth season of 2 years, either monthly (2004) or bi‐monthly (2005), to observe whether stream inputs increased prey availability and whether variation in total macroinvertebrate biomass was tracked by fish. 2. Observations from eight headwater streams indicated that streams did not increase the total macroinvertebrate biomass in the shallow littoral zone at stream inflows, relative to adjacent plots without stream inputs (controls). The taxonomic composition of stream macroinvertebrates drifting toward the lake differed from that in the littoral lake benthos itself, although there was no evidence of any species change in the composition of the littoral benthos brought about by stream inputs. 3. Although streams made no measurable contribution to the biomass or taxonomic composition of the littoral macroinvertebrate benthos, there was substantial temporal variation in biomass among the eight sites for each of the (n = 7) sample periods during which observations were made. Variation in total biomass was primarily a function of bottom slope and benthic substrata in the lake habitats. Dominant taxonomic groups were Baetidae, Ephemerellidae (two genera), Leptophlebiidae, Chironomidae (three subfamilies) and Perlodidae, although we did not determine the specific substratum affinities of each taxon. 4. Mixed effects linear models identified a significant interaction between macroinvertebrate biomass and plot type (stream inflow vs. control) associated with fish abundance. Across the observed range of macroinvertebrate biomass, fish showed a significant preference for stream inflows, but more closely tracked food availability in the controls. For young‐of‐the‐year (YOY), a negative effect of temperature was also included in the model, and we observed lower temperatures at stream inflows. However, abundance of predatory adults affected habitat selection for YOY. Lake‐bottom slope also accounted for variation in abundance in both fish models. 5. Our results suggest that the effect of fishless headwater streams on downstream fish may not always be through direct delivery of food. In this study system, fish preferred stream inflow plots, but this preference interacted with macroinvertebrate biomass in a manner that was difficult to explain. For YOY, predation risk was related to the preference for stream inflows, although the specific factor that mitigates predation risk remains poorly understood.     

Turbidity amplifies the non‐lethal effects of predation and affects the foraging success of characid fish shoals

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Assessing changes in arthropod predator–prey interactions through DNA‐based gut content analysis—variable environment,stable diet

Analysing the structure and dynamics of biotic interaction networks and the processes shaping them is currently one of the key fields in ecology. In this paper, we develop a novel approach to gut content analysis, thereby deriving a new perspective on community interactions and their responses to environment. For this, we use an elevational gradient in the High Arctic, asking how the environment and species traits interact in shaping predator–prey interactions involving the wolf spider Pardosa glacialis. To characterize the community of potential prey available to this predator, we used pitfall trapping and vacuum sampling. To characterize the prey actually consumed, we applied molecular gut content analysis. Using joint species distribution models, we found elevation and vegetation mass to explain the most variance in the composition of the prey community locally available. However, such environmental variables had only a small effect on the prey community found in the spider's gut. These observations indicate that Pardosa exerts selective feeding on particular taxa irrespective of environmental constraints. By directly modelling the probability of predation based on gut content data, we found that neither trait matching in terms of predator and prey body size nor phylogenetic or environmental constraints modified interaction probability. Our results indicate that taxonomic identity may be more important for predator–prey interactions than environmental constraints or prey traits. The impact of environmental change on predator–prey interactions thus appears to be indirect and mediated by its imprint on the community of available prey.     

Differences in predator composition alter the direction of structure‐mediated predation risk in macrophyte communities

Structural complexity strongly influences the outcome of predator–prey interactions in benthic marine communities affecting both prey concealment and predator hunting efficacy. How habitat structure interacts with species‐specific differences in predatory style and antipredatory strategies may therefore be critical in determining higher trophic functions. We examined the role of structural complexity in mediating predator–prey interactions across several macrophyte habitats along a gradient of structural complexity in three different bioregions: western Mediterranean Sea (WMS), eastern Indian Ocean (EIO) and northern Gulf of Mexico (NGM). Using sea urchins as model prey, we measured survival rates of small (juveniles) and medium (young adults) size classes in different habitat zones: within the macrophyte habitat, along the edge and in bare sandy spaces. At each site we also measured structural variables and predator abundance. Generalised linear models identified biomass and predatory fish abundance as the main determinants of predation intensity but the efficiency of predation was also influenced by urchin size class. Interestingly though, the direction of structure‐mediated effects on predation risk was markedly different between habitats and bioregions. In WMS and NGM, where predation by roving fish was relatively high, structure served as a critical prey refuge, particularly for juvenile urchins. In contrast, in EIO, where roving fish predation was low, predation was generally higher inside structurally complex environments where sea stars were responsible for much of the predation. Larger prey were generally less affected by predation in all habitats, probably due to the absence of large predators. Overall, our results indicate that, while the structural complexity of habitats is critical in mediating predator–prey interactions, the direction of this mediation is strongly influenced by differences in predator composition. Whether the regional pool of predators is dominated by visual roving species or chemotactic benthic predators may determine if structure dampens or enhances the influence of top–down control in marine macrophyte communities.     

Shift in predation regime mediates diversification of foraging behaviour in a dragonfly genus

1. Behavioural adaptations to avoid and evade predators are common. Many studies have investigated population divergence in response to changes in predation regime within species, but studies exploring interspecific patterns are scant. Studies on interspecific divergence can infer common outcomes from evolutionary processes and highlight the role of environmental constraints in shaping species traits. 2. Species of the dragonfly genus Leucorrhinia underwent well‐studied shifts from habitats being dominated by predatory fish (fish lakes) to habitat being dominated by predatory invertebrates (dragonfly lakes). This change in top predators resulted in a set of adaptive trait modifications in response to the different hunting styles of both predator types: whereas predatory fish actively search and pursue prey, invertebrate predator follow a sit‐and‐wait strategy, not pursuing prey. 3. Here it is shown that the habitat shift‐related change in selection regime on larval Leucorrhinia caused species in dragonfly lakes to evolve increased larval foraging and activity, and results suggest that they lost the ability to recognise predatory fish. 4. The results of the present study highlight the impact of predators on behavioural trait diversification with habitat‐specific predation regimes selecting for distinct behavioural expression.     

Virtual plankton: A novel approach to the investigation of aquatic predator‐prey interactions

Small‐scale zooplankton swimming behaviors can affect aquatic predator‐prey interactions. Difficulties in controlling prey swimming behavior however, have restricted the ability to test hypotheses relating differences in small‐scale swimming behavior to frequency of predation by fish. We report here a Virtual Plankton (VP) system that circumvents this problem by allowing the observation of fish “preying"on computer‐generated prey images whose size, shape, color and swimming behavior can be precisely controlled. Two experiments were performed in which bluegill sunfish (Lepomis macrochirus) were given a choice of either two VP images, one of which moved twice as fast as the other, or six VP, one of which moved either faster (1.25 x, 1.5 x or 2 x ) or slower (0.5 x) than the other five. Current predator‐prey models based on encounter probabilities and prey visibility predict that moving faster increases predation risk and conversely, moving slower decreases predation risk. In agreement with existing predator‐prey models, in both experiments, fish chose faster moving VP significantly more often than their slower moving neighbors. Contrary to the predictions of existing models, in the second experiment with six VP, the rate at which fish chose a prey image moving half as fast as the five surrounding images did not differ significantly from the rate predicted by chance(l/6). These results suggest that current fish‐zooplankton predation models would benefit by the incorporation of small‐scale swimming behavior and assessments of its influence on overall prey visibility.     

Consequences of temperature and temperature variability on swimming activity,group structure,and predation of endangered delta smelt

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Implications of shared predation for space use in two sympatric leporids

Spatial variation in habitat riskiness has a major influence on the predator–prey space race. However, the outcome of this race can be modulated if prey shares enemies with fellow prey (i.e., another prey species). Sharing of natural enemies may result in apparent competition, and its implications for prey space use remain poorly studied. Our objective was to test how prey species spend time among habitats that differ in riskiness, and how shared predation modulates the space use by prey species. We studied a one‐predator, two‐prey system in a coastal dune landscape in the Netherlands with the European hare (Lepus europaeus) and European rabbit (Oryctolagus cuniculus) as sympatric prey species and red fox (Vulpes vulpes) as their main predator. The fine‐scale space use by each species was quantified using camera traps. We quantified residence time as an index of space use. Hares and rabbits spent time differently among habitats that differ in riskiness. Space use by predators and habitat riskiness affected space use by hares more strongly than space use by rabbits. Residence time of hare was shorter in habitats in which the predator was efficient in searching or capturing prey species. However, hares spent more time in edge habitat when foxes were present, even though foxes are considered ambush predators. Shared predation affected the predator–prey space race for hares positively, and more strongly than the predator–prey space race for rabbits, which were not affected. Shared predation reversed the predator–prey space race between foxes and hares, whereas shared predation possibly also released a negative association and promoted a positive association between our two sympatric prey species. Habitat riskiness, species presence, and prey species’ escape mode and foraging mode (i.e., central‐place vs. noncentral‐place forager) affected the prey space race under shared predation.     

Sacrificial males: the potential role of copulation and predation in contributing to copepod sex‐skewed ratios

Predation is thought to play a selective role in the emergence of behavioural traits in prey. Differences in behaviour between prey demographics may, therefore, be driven by predation with select components of the population being less vulnerable to predators. While under controlled conditions prey demography has been shown to have consequences for predation success, investigations linking these implications to natural prey population demographics are scarce. Here we assess predator–prey dynamics between notonectid predators (backswimmers) and Lovenula raynerae (Copepoda), key faunal groups in temperate ephemeral pond ecosystems. Using a combination of field and experimental approaches we test for the development and mechanism of predation‐induced sex‐skewed ratios. A natural population of L. raynerae was tracked over time in relation to their predator (notonectid) and prey (Cladocera) numbers. In the laboratory, L. raynerae sex ratios were also assessed over time but in the absence of predation pressure. Predation success and prey performance experiments evaluating differences between L. raynerae male, female, gravid female and copulating pairs exposed to notonectid predation were then examined. Under natural conditions, a female dominated copepod population developed over time and was correlated to predation pressure, while under predator‐free conditions non sex‐skewed prey population demographics persisted. Predator–prey laboratory trials showed no difference in vulnerability and escape performance for male, female and gravid female copepods, but pairs in copula were significantly more vulnerable to predation. This vulnerability was not shared by both sexes, with only female copepods ultimately escaping from successful predation on a mating pair. These results suggest that contact periods during copula may contribute to the development of sex‐skewed copepod ratios over time in ecosystems dominated by hexapod predators. This is discussed within the context of vertebrate and invertebrate predation and how these dissimilar types of predation are likely to have acted as selective pressures for copepod mating systems.     

Picking personalities apart: estimating the influence of predation,sex and body size on boldness in the guppy Poecilia reticulata

Predation is a strong selective force in most natural systems, potentially fueling evolutionary changes in prey morphology, life history and behaviour. Recent work has suggested that contrasting predation pressures may lead to population differentiation in personality traits. However, there are indications that these personality traits also differ between sexes and not necessarily in a consistent way between populations. We used an integrative approach to quantify boldness (latency to emerge from a shelter) in wild‐caught guppies in relation to predation pressure, population origin, sex and size. In addition we quantified the repeatability of these personality traits. We show that predation regime had significant effects on emergence time. In general, fish from high predation localities emerged sooner from the shelter compared to those from low predation localities. We found strong sex differences; males were significantly bolder than females. The relationship between emergence time and body size was non‐significant in all populations. We discuss what responses to expect from predator‐naïve versus predator‐experienced individuals and how this can be linked to the shyness–boldness continuum.     

18S rRNA metabarcoding diet analysis of a predatory fish community across seasonal changes in prey availability

Predator–prey relationships are important ecological interactions, affecting biotic community composition and energy flow through a system, and are of interest to ecologists and managers. Morphological diet analysis has been the primary method used to quantify the diets of predators, but emerging molecular techniques using genetic data can provide more accurate estimates of relative diet composition. This study used sequences from the 18S V9 rRNA barcoding region to identify prey items in the gastrointestinal (GI) tracts of predatory fishes. Predator GI samples were taken from the Black River, Cheboygan Co., MI, USA (n = 367 samples, 12 predator species) during periods of high prey availability, including the larval stage of regionally threatened lake sturgeon (Acipenser fulvescens Rafinesque 1817) in late May/early June of 2015 and of relatively lower prey availability in early July of 2015. DNA was extracted and sequenced from 355 samples (96.7%), and prey DNA was identified in 286 of the 355 samples (80.6%). Prey were grouped into 33 ecologically significant taxonomic groups based on the lowest taxonomic level sequences that could be identified using sequences available on GenBank. Changes in the makeup of diet composition, dietary overlap, and predator preference were analyzed comparing the periods of high and low prey abundance. Some predator species exhibited significant seasonal changes in diet composition. Dietary overlap was slightly but significantly higher during the period of high prey abundance; however, there was little change in predator preference. This suggests that change in prey availability was the driving factor in changing predator diet composition and dietary overlap. This study demonstrates the utility of molecular diet analysis and how temporal variability in community composition adds complexity to predator–prey interactions.     

Facilitation and interference among three predators affect their consumption of a stream-dwelling mayfly

1. We experimentally tested if a multiplicative risk model accurately predicted the consumption of a common mayfly at risk of predation from three predator species in New Zealand streams. Deviations between model predictions and experimental observations were interpreted as indicators of ecologically important interactions between predators. 2. The predators included a drift‐feeding fish [brown trout (T), Salmo trutta], a benthivorous fish [galaxiid (G), koaro, Galaxias brevipennis] and a benthic predatory stonefly (S; Stenoperla sp.) with Deleatidium sp. mayflies as prey. Eight treatments with all predator species combinations and a predator‐free control were used. Experiments were performed in aquaria with cobbles as predator refuges for mayflies and we measured the proportion of prey consumed after 6 h for both day and night trials. 3. Trout consumed a higher proportion of prey than other predators. For the two predator treatments we found less than expected prey consumption in the galaxiid + trout treatment (G + T) for both day and night trials, whereas a higher than expected proportion of prey was consumed during night time in the stonefly + trout (S + T) treatment. 4. The results indicate interference (G + T) and facilitation (S + T) between predators depending on predator identity and time of day. Thus, to make accurate predictions of interspecific interactions, it is necessary to consider the ecology of individual species and how differences influence the direction and magnitude of interactions.     

Two common invertebrate predators show varying predation responses to different types of sentinel prey

Sentinel prey (an artificially manipulated patch of prey) are widely used to assess the level of predation provided by natural enemies in agricultural systems. Whilst a number of different methodologies are currently in use, little is known about how arthropod predators respond to artificially manipulated sentinel prey in comparison with predation on free‐living prey populations. We assessed how attack rates on immobilized (aphids stuck to cards) and artificial (plasticine lepidopteran larvae mimics) sentinel prey differed to predation on free‐moving live prey (aphids). Predation was assessed in response to density of the common invertebrate predators, a foliar‐active ladybird Harmonia axyridis (Coleoptera: Coccinellidae), and a ground‐active beetle Pterostichus madidus (Coleoptera: Carabidae). Significant increases in attack rates were found for the immobilized and artificial prey between the low and high predator density treatments. However, an increased predator density did not significantly reduce numbers of free‐living live aphids included in the mesocosms in addition to the alternate prey. We also found no signs of predation on the artificial prey by the predator H. axyridis. These findings suggest that if our assessment of predation had been based solely on the foliar artificial prey, then no increase in predation would have been found in response to increased predator density. Our results demonstrate that predators differentially respond to sentinel prey items which could affect the level of predation recorded where target pest species are not being used.     

Strategies of zooplanktivory shape the dynamics and diversity of littoral plankton communities: a mesocosm approach

Planktivorous fish can exert strong top‐down control on zooplankton communities. By incorporating different feeding strategies, from selective particulate feeding to cruising filter feeding, fish species target distinct prey. In this study, we investigated the effects of two species with different feeding strategies, the three‐spined stickleback (Gasterosteus aculeatus (L.)) and roach (Rutilus rutilus (L.)), on a low‐diversity brackish water zooplankton community using a 16‐day mesocosm experiment. The experiment was conducted on a small‐bodied spring zooplankton community in high‐nutrient conditions, as well as a large‐bodied summer community in low‐nutrient conditions. Effects were highly dependent on the initial zooplankton community structure and hence seasonal variation. In a small‐bodied community with high predation pressure and no dispersal or migration, the selective particulate‐feeding stickleback depleted the zooplankton community and decreased its diversity more radically than the cruising filter‐feeding roach. Cladocerans rather than copepods were efficiently removed by predation, and their removal caused altered patterns in rotifer abundance. In a large‐bodied summer community with initial high taxonomic and functional diversity, predation pressure was lower and resource availability was high for omnivorous crustaceans preying on other zooplankton. In this community, predation maintained diversity, regardless of predator species. During both experimental periods, predation influenced the competitive relationship between the dominant calanoid copepods, and altered species composition and size structure of the zooplankton community. Changes also occurred to an extent at the level of nontarget prey, such as microzooplankton and rotifers, emphasizing the importance of subtle predation effects. We discuss our results in the context of the adaptive foraging mechanism and relate them to the natural littoral community.     

Evolutionary trends within bivalve prey of chesapeake group naticid gastropods

Five genera of Miocene bivalves evolved antipredatory adaptations in response to predation by drilling naticid gastropods. I examined the evolution of two traits affecting predator‐prey interaction, prey shell thickness (TH) and internal volume (IV). Thickness controls predation costs by determining drilling time, and internal volume influences the benefit derived by the predator.

Internal volume showed no consistent pattern of temporal change among the taxa studied. IV fluctuated nondirectionally during the history of most genera, though both increasing and decreasing trends occurred within species ofAstarte. In contrast, all five genera exhibited significant thickness increases (from 8–157%) during the three‐million‐year interval. Both gradual intraspecific and interspecific directional changes occurred. Taxa with the greatest predation intensities displayed the most change, suggesting that predation selected for the thickness increases. Increased thickness apparently reduced predation; a significant negative correlation between TH and predation intensity occurred within four of the bivalve genera. Improvement of predator capabilities apparently did not keep pace with increased antipredatory morphologic adaptations over the interval studied.     

The underrated importance of predation in transmission ecology of direct lifecycle parasites

Most parasites with complex life cycles exploit trophic webs to pass from host to host in order to develop and, eventually, reproduce. Thus predation constitutes the necessary route for transmission. Conversely, the transmission of parasites that use a single host to develop and reproduce should be, in principle, not particularly affected by host trophic ecology. Here I challenge this view, showing that predation may be relevant also for direct lifecycle parasites. I used a large dataset of fish trophic interactions to investigate if the degree of monogenean species overlap in predators and prey deviated from randomness. I demonstrated that predators and prey often share more monogenean parasite genera than explained by host habitat ecology, geographical distribution and phylogeny. This suggests that predation may play an important role in promoting monogenean host range expansion. In addition, a non‐negligible proportion of considered prey–predator pairs showed a significantly high overlap in their monogenean parasites at the species level. This may indicate a tendency of some monogenean parasites to evolve transmission strategies targeted towards host interactions. If this hypothesis is true, these monogenean parasites would be much more vulnerable to co‐extinction than previously thought. Synthesis Predation is not expected to play an important role in the ecology and evolution of simple life cycle parasites. Yet, several predator fish tend to share with their prey more monogenean parasites than one would expect predicted from their geographical distribution, habitat preference, and or phylogenetic relationships. This suggests that some monogenean parasites have evolved transmission strategies more targeted towards host interactions than towards species‐specific traits. If this hypothesis is supported, it would have strong implications on host–parasite evolutionary ecology, primarily, suggesting the existence of peculiar situations where some parasites have evolved high specialized host finding behaviors to expand their host range.