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At least 217 records · Page 12Linked to original sources

Learning rate and temperament in a high predation risk environment

Living in challenging environments can influence the behavior of animals in a number of ways. For instance, populations of prey fish that experience frequent, nonlethal interactions with predators have a high proportion of individuals that express greater reaction to risk and increased activity and exploration—collectively known as temperament traits. Temperament traits are often correlated, such that individuals that are risk-prone also tend to be active and explore more. Spatial learning, which requires the integration of many sensory cues, has also been shown to vary in fish exposed to different levels of predation threat. Fish from areas of low predation risk learn to solve spatial tasks faster than fish from high predation areas. However, it is not yet known whether simpler forms of learning, such as learning associations between two events, are similarly influenced. Simple forms of associative learning are likely to be affected by temperament because a willingness to approach and explore novel situations could provide animals with a learning advantage. However, it is possible that routine-forming and inflexible traits associated with risk-prone and increased exploratory behavior may act in the opposite way and make risk-prone individuals poorer at learning associations. To investigate this, we measured temperament in Panamanian bishop fish ( Brachyrhaphis episcopi ) sampled from a site known to contain many predators. The B. episcopi were then tested with an associative learning task. Within this population, fish that explored more were faster at learning a cue that predicted access to food, indicating a link between temperament and basic learning abilities.

Oecologia

Methylmercury-Total mercury ratios in predator and primary consumer insects from Adirondack streams (New York, USA)

Mercury (Hg) is a global pollutant that affects biota in remote settings due to atmospheric deposition of inorganic Hg, and its conversion to methylmercury (MeHg), the bioaccumulating and toxic form. Characterizing biotic MeHg is important for evaluating aquatic ecosystem responses to changes in Hg inputs. Aquatic insects possess many qualities desired for MeHg biomonitoring, but are not widely used, largely because of limited information regarding percentages of total mercury (THg) composed of MeHg (i.e., MeHg%) in various taxa. Here, we examine taxonomic, spatial, and seasonal variation in MeHg% of stream-dwelling predator and primary-consumer insects from nine streams in the Adirondack region (NY, USA). Predator MeHg% was high (median 94%) and did not differ significantly among five taxa. MeHg% in selected dragonflies (the most abundant predators, Odonata: Aeshnidae and Libellulidae) exhibited little seasonal and spatial variation, and THg concentration was strongly correlated with aqueous (filtered) MeHg (FMeHg; r s = 0.76). In contrast, MeHg% in primary consumers—shredders (northern caddisflies [Trichoptera: Limnephilidae]) and scrapers (flathead mayflies [Ephemeroptera: Heptageniidae]), were lower (medians 52% and 35%, respectively), and differed significantly between taxa, among sites, and seasonally. Correlations of THg with FMeHg were weak (shredders, r s = 0.45, p = 0.09) or not significant (scrapers, p = 0.89). The higher MeHg% of predators corresponded with their higher trophic positions (indicated by nitrogen stable isotopes). Results suggest obligate predators hold the most promise for the use of THg as a surrogate for MeHg biomonitoring with aquatic insects within the Adirondack region.

Adirondack Mountains

Predators shape distribution and promote diversification of morphological defenses in Leucorrhinia , Odonata

Predators strongly influence species assemblages and shape morphological defenses of prey. Interestingly, adaptations that constitute effective defenses against one type of predator may render the prey susceptible to other types of predators. Hence, prey may evolve different strategies to escape predation, which may facilitate adaptive radiation of prey organisms. Larvae of different species in the dragonfly genus Leucorrhinia have various morphological defenses. We studied the distribution of these larvae in relation to the presence of predatory fish. In addition, we examined the variation in morphological defenses within species with respect to the occurrence of fish. We found that well-defended species, those with more and longer spines, were more closely associated with habitats inhabited by predatory fish and that species with weakly developed morphological defenses were more abundant in habitats without fish. The species predominantly connected to lakes with or without fish, respectively, were not restricted to a single clade in the phylogeny of the genus. Our data is suggestive of phenotypic plasticity in morphological defense in three of the studied species since these species showed longer spines in lakes with fish. We suggest that adaptive phenotypic plasticity may have broadened the range of habitats accessible to Leucorrhinia . It may have facilitated colonization of new habitats with different types of predators, and ultimately, speciation through adaptive radiation.

Evolutionary Ecology

Behavioural response of adult sea lamprey (Petromyzon marinus) to predator and conspecific alarm cues: evidence of additive effects

Sea lampreys Petromyzon marinus, an invasive pest in the Upper Great Lakes, avoid odours that represent danger in their habitat. These odours include conspecific alarm cues and predator cues, like 2-phenylethylamine hydrochloride (PEA HCl), which is found in the urine of mammalian predators. Whether conspecific alarm cues and predator cues function additively or synergistically when mixed together is unknown. The objectives of this experimental study were to determine if the avoidance response of sea lamprey to PEA HCl is proportional to the concentration delivered, and if the avoidance response to the combination of a predator cue (PEA HCl) and sea lamprey alarm cue is additive. To accomplish the first objective, groups of ten sea lampreys were placed in an artificial stream channel and presented with stepwise concentrations of PEA HCl ranging from 5 × 10−8 to 5 × 10−10 M and a deionized water control. Sea lampreys exhibited an increase in their avoidance behaviour in response to increasing concentrations of PEA HCl. To accomplish the second objective, sea lampreys were exposed to PEA HCl, conspecific alarm cue and a combination of the two. Sea lampreys responded to the combination of predator cue and conspecific alarm cue in an additive manner.

St.Mary's River

Effects of rodent species, seed species, and predator cues on seed fate

Seed selection, removal and subsequent management by granivorous animals is thought to be a complex interaction of factors including qualities of the seeds themselves (e.g., seed size, nutritional quality) and features of the local habitat (e.g. perceived predator risk). At the same time, differential seed selection and dispersal is thought to have profound effects on seed fate and potentially vegetation dynamics. In a feeding arena, we tested whether rodent species, seed species, and indirect and direct predation cues influence seed selection and handling behaviors (e.g., scatter hoarding versus larder hoarding) of two heteromyid rodents, Ord's kangaroo rat (Dipodomys ordii) and the Great Basin pocket mouse (Perognathus parvus). The indirect cue was shrub cover, a feature of the environment. Direct cues, presented individually, were (1) control, (2) coyote (Canis latrans) vocalization, (3) coyote scent, (4) red fox (Vulpes vulpes) scent, or (5) short-eared owl (Asio flammeus) vocalization. We offered seeds of three sizes: two native grasses, Indian ricegrass (Achnatherum hymenoides) and bluebunch wheatgrass (Pseudoroegneria spicata), and the non-native cereal rye (Secale cereale), each in separate trays. Kangaroo rats preferentially harvested Indian ricegrass while pocket mice predominately harvested Indian ricegrass and cereal rye. Pocket mice were more likely to scatter hoard preferred seeds, whereas kangaroo rats mostly consumed and/or larder hoarded preferred seeds. No predator cue significantly affected seed preferences. However, both species altered seed handling behavior in response to direct predation cues by leaving more seeds available in the seed pool, though they responded to different predator cues. If these results translate to natural dynamics on the landscape, the two rodents are expected to have different impacts on seed survival and plant recruitment via their different seed selection and seed handling behaviors.

Acta Oecologica

Dynamics of a plant-herbivore-predator system with plant-toxicity

A system of ordinary differential equations is considered that models the interactions of two plant species populations, an herbivore population, and a predator population. We use a toxin-determined functional response to describe the interactions between plant species and herbivores and use a Holling Type II functional response to model the interactions between herbivores and predators. In order to study how the predators impact the succession of vegetation, we derive invasion conditions under which a plant species can invade into an environment in which another plant species is co-existing with a herbivore population with or without a predator population. These conditions provide threshold quantities for several parameters that may play a key role in the dynamics of the system. Numerical simulations are conducted to reinforce the analytical results. This model can be applied to a boreal ecosystem trophic chain to examine the possible cascading effects of predator-control actions when plant species differ in their levels of toxic defense.

Mathematical Biosciences

Sizes of prey consumed by two pelagic predators in US reservoirs: Implications for quantifying biomass of available prey

Striped bass Morone saxatilis and hybrid bass M. saxatilis x M. chrysops have been stocked to establish fisheries in many US reservoirs, but success has been limited by a poor understanding of relations between prey biomass and predator growth and survival. To define sizes of prey that are morphologically available, we developed predictive relationships between predator length, mouth dimensions, and expected maximum prey size; predictions were then validated using published data on sizes of clupeid prey (Dorosoma spp.) in five US reservoirs. Further, we compared the biomass of prey considered available to predators using two forms of a length-based consumption model - a previously published AP/P ratio and a revised model based on our results. Predictions of maximum prey size using predator GW were consistent with observed prey sizes in US reservoirs. Length of consumed Dorosoma was significantly, but weakly, correlated with predator length in four of the five reservoirs (r2 = 0.006-0.336, P < 0.05). Model predictions of available prey biomass differed by as much as 800% between the original AP/P model and a revision based on our estimates of maximum available prey size. The revised model predicted less available prey biomass in cases where large Dorosoma (>150 mm TL) were abundant. (C) 2000 Elsevier Science B.V.

Fisheries Research

Quantifying the importance of ontogeny and prey type in modeling top-down and bottom-up effects of an ectothermic predator

Dietary decisions by predators can affect prey abundance and overall food web dynamics. Many predators do not forage on the same prey at the same frequency throughout their lives. Ontogenetic shifts in prey preference are not, however, often accounted for when modeling food web relationships, despite growing literature that suggests that stage specific dietary relationships may be an important consideration when modeling trophic interactions. We investigated the importance of considering size-structure of a predator population with ontogenetic diet shifts in evaluating relationships with prey response using a manipulative experiment with the brown treesnake ( Boiga irregularis ) in Guam. After removing ~ 40% of the snake population via toxic mammal carrion, we measured the strength of the relationship between snake density and the response of two types of prey (lizards and mammals). We evaluated these relationships based on total population size or division of the population into stage specific size categories based on diet preference predictions. We hypothesized that the density of juvenile snakes would correlate more strongly with lizard detections, while adult snakes would better correlate to rodent detections. We also measured reproductive output following changes in rates of prey detection. As expected by known ontogenetic shifts in dietary preference, explicit stage-based models better predicted shifts in rates of observed prey items than did total predator density for both lizards and mammals. Additionally, rodent detections were predictive of one reproductive pulse from snakes, while lizard detections were not predictive or correlated. Our findings support that consideration of predatory species stage-based dietary preference can be meaningful for understanding food web dynamics, particularly when a predator has a broad diet or one that changes through time.

Guam

Trophic ontogeny of fluvial Bull Trout and seasonal predation on Pacific Salmon in a riverine food web

Bull Trout Salvelinus confluentus are typically top predators in their host ecosystems. The Skagit River in northwestern Washington State contains Bull Trout and Chinook Salmon Oncorhynchus tshawytscha populations that are among the largest in the Puget Sound region and also contains a regionally large population of steelhead O. mykiss (anadromous Rainbow Trout). All three species are listed as threatened under the Endangered Species Act (ESA). Our objective was to determine the trophic ecology of Bull Trout, especially their role as predators and consumers in the riverine food web. We seasonally sampled distribution, diets, and growth of Bull Trout in main-stem and tributary habitats during 2007 and winter–spring 2008. Consumption rates were estimated with a bioenergetics model to (1) determine the annual and seasonal contributions of different prey types to Bull Trout energy budgets and (2) estimate the potential impacts of Bull Trout predation on juvenile Pacific salmon populations. Salmon carcasses and eggs contributed approximately 50% of the annual energy budget for large Bull Trout in main-stem habitats, whereas those prey types were largely inaccessible to smaller Bull Trout in tributary habitats. The remaining 50% of the energy budget was acquired by eating juvenile salmon, resident fishes, and immature aquatic insects. Predation on listed Chinook Salmon and steelhead/Rainbow Trout was highest during winter and spring (January–June). Predation on juvenile salmon differed between the two study years, likely due to the dominant odd-year spawning cycle for Pink Salmon O. gorbuscha . The population impact on ocean- and stream-type Chinook Salmon was negligible, whereas the impact on steelhead/Rainbow Trout was potentially very high. Due to the ESA-listed status of Bull Trout, steelhead, and Chinook Salmon, the complex trophic interactions in this drainage provide both challenges and opportunities for creative adaptive management strategies.

Washington

Effects of hatchery fish density on emigration, growth, survival, and predation risk of natural steelhead parr in an experimental stream channel

Hatchery supplementation of steelhead Oncorhynchus mykiss raises concerns about the impacts on natural populations, including reduced growth and survival, displacement, and increased predation. The potential risks may be density dependent.We examined how hatchery stocking density and the opportunity to emigrate affect the responses of natural steelhead parr in an experimental stream channel and after 15 d found no density-dependent effects on growth, emigration, or survival at densities ranging from 1-6 hatchery parr/m 2 . The opportunity for steelhead parr to emigrate reduced predation by coastal cutthroat trout O. clarkii clarkii on both hatchery and natural steelhead parr. The cutthroat trout exhibited a type-I functional response (constant predation rate with increased prey density) for the hatchery and composite populations. In contrast, the predation rate on natural parr decreased as hatchery stocking density increased. Supplementation with hatchery parr at any experimental stocking density reduced the final natural parr density. This decline was explained by increased emigration fromthe supplemented groups. Natural parr had higher mean instantaneous growth rates than hatchery parr. The proportion of parr emigrating decreased as parr size increased over successive experimental trials. Smaller parr had lower survival and suffered higher predation. The final density of the composite population, a measure of supplementation effectiveness, increased with the hatchery steelhead stocking rate. Our results indicate that stocking larger hatchery parr (over 50 d postemergence) at densities within the carrying capacity would have low short-term impact on the growth, survival, and emigration of natural parr while increasing the density of the composite population; in addition, a stocking density greater than 3 fish/m 2 might be a good starting point for the evaluation of parr stocking in natural streams.

North American Journal of Fisheries Management

Prey ration, temperature, and predator species influence digestion rates of prey DNA inferred from qPCR and metabarcoding

Diet analysis is a vital tool for understanding trophic interactions and is frequently used to inform conservation and management. Molecular approaches can identify diet items that are impossible to distinguish using more traditional visual-based methods. Yet, our understanding of how different variables, such as predator species or prey ration size, influence molecular diet analysis is still incomplete. Here, we conducted a large feeding trial to assess the impact that ration size, predator species, and temperature had on digestion rates estimated with visual identification, qPCR, and metabarcoding. Our trial was conducted by feeding two rations of Chinook salmon ( Oncorhynchus tshawytscha ) to two piscivorous fish species (largemouth bass [ Micropterus salmoides ] and channel catfish [ Ictalurus punctatus ]) held at two different temperatures (15.5 and 18.5°C) and sacrificed at regular intervals up to 120 h from the time of ingestion to quantify the prey contents remaining in the digestive tract. We found that ration size, temperature, and predator species all influenced digestion rate, with some indication that ration size had the largest influence. DNA-based analyses were able to identify salmon smolt prey in predator gut samples for much longer than visual analysis (~12 h for visual analysis vs. ~72 h for molecular analyses). Our study provides evidence that modelling the persistence of prey DNA in predator guts for molecular diet analyses may be feasible using a small set of controlling variables for many fish systems.

Molecular Ecology Resources

The rise of an apex predator following deglaciation

Aim Sea otters ( Enhydra lutris ) are an apex predator of the nearshore marine community and nearly went extinct at the turn of the 20th century. Reintroductions and legal protection allowed sea otters to re‐colonize much of their former range. Our objective was to chronicle the colonization of this apex predator in Glacier Bay, Alaska, to help understand the mechanisms that governed their successful colonization. Location Glacier Bay is a tidewater glacier fjord in southeastern Alaska that was entirely covered by glaciers in the mid‐18th century. Since then, it has endured the fastest tidewater glacier retreat in recorded history. Methods We collected and analysed several data sets, spanning 20 years, to document the spatio‐temporal dynamics of an apex predator expanding into an area where they were formerly absent. We used novel quantitative tools to model the occupancy, abundance and colonization dynamics of sea otters, while accounting for uncertainty in the data collection process, the ecological process and model parameters. Results Twenty years after sea otters were first observed colonizing Glacier Bay, they became one of the most abundant and widely distributed marine mammal. The population grew exponentially at a rate of 20% per year. They colonized Glacier Bay at a maximum rate of 6 km per year, with faster colonization rates occurring early in the colonization process. During colonization, sea otters selected shallow areas, close to shore, with a steep bottom slope, and a relatively simple shoreline complexity index. Main conclusions The growth and expansion of sea otters in Glacier Bay demonstrate how legal protection and translocation of apex predators can facilitate their successful establishment into a community in which they were formerly absent. The success of sea otters was, in part, a consequence of habitat that was left largely unperturbed by humans for the past 250 years. Further, sea otters and other marine predators, whose distribution is limited by ice, have the potential to expand in distribution and abundance, reshaping future marine communities in the wake of deglaciation and global loss of sea ice.

Alaska

Habitat overlap of juvenile and adult lake trout of Great Bear Lake: Evidence for lack of a predation gradient?

A range of organisms, from plankton to fish, commonly shift their habitat distributions horizontally or vertically due to predation risk. Juvenile lake trout, Salvelinus namaycush , are generally viewed as occupying deep areas of lakes to decrease predation pressure from adults. In contrast, we found that juvenile lake trout from Great Bear Lake, NT, Canada, occupied a variety of habitats and from shallow to deep depths (0–150 m), overlapping with adult lake trout. No evidence occurred for a length depth‐based segregation (e.g., ontogenetic shift). Genetic variation was also similar among juveniles in the different depth zones. However, isotopic niches and C:N ratios among juveniles showed some variability in niche widths and positions for individuals caught from the 51–150 m zone compared to juvenile individuals caught from 0–20 m and 21–50 m zones. The uniformly distributed adult lake trout in Great Bear Lake may evenly distribute predation pressure (including cannibalism) across shallow‐ and deep‐water habitats more than in other lakes. As a result, juveniles may respond to differences in foraging opportunities rather than predation risks. Juvenile lake trout did not appear to conform to the general pattern of juveniles seeking a deep‐water refuge to reduce predation risks. In contrast, juvenile lake trout of Great Bear Lake displayed broad resource use across all depths and habitats.

Northwest Territories

Is predation on waterfowl nests density dependent? Tests at three spatial scales

We tested whether predation on duck nests ( Anas spp.) was density dependent at three spatial scales using artificial and natural nests in the Suisun Marsh, California, USA. At the largest spatial scale, we used 5 years (1998–2002) of data to examine the natural variation in duck nest success and nest densities among 8–16 fields per year, each 5–33 ha in size (n=62 fields). At an intermediate spatial scale, we deployed artificial nests (2000, n=280) within 1-ha plots at three experimental densities (5, 10, and 20 nests ha −1 ) in a complete randomized block design and examined differences in nest predation. At the smallest spatial scale, we examined nest success in relation to nearest-neighbor fates and distances for artificial (2000, n=280) and natural nests (2000, n=507). We detected no relationship between nest success and the density of natural nests among fields in any year, nor when we pooled data for all years after controlling for year effects. The proportion of artificial nests that survived also did not depend on experimental nest densities within 1-ha plots. Overall, 15.0±12.4%, 15.0±11.0%, and 6.2±4.3% of artificial nests survived the 32-day exposure period in the low, intermediate, and high nest densities, respectively. Additionally, we detected no consistent effect of nearest-neighbor fate or distance on the success of artificial or natural nests. Thus, our results provide no evidence of density-dependent predation on duck nests at any scale of analysis, in contrast to a number of previous studies. Variation among geographical locations in the degree to which predation is density-dependent may reflect the composition of the predator community and the availability of alternate prey.

Oikos

Variation in wind and piscivorous predator fields affecting the survival of Atlantic salmon, Salmo salar, in the Gulf of Maine

Observations relevant to the North American stock complex of Atlantic salmon, Salmo salar L., suggest that marine mortality is influenced by variation in predation pressure affecting post‐smolts during the first months at sea. This hypothesis was tested for Gulf of Maine (GOM) stocks by examining wind pseudostress and the distribution of piscivorous predator fields potentially affecting post‐smolts. Marine survival has declined over recent decades with a change in the direction of spring winds, which is likely extending the migration of post‐smolts by favouring routes using the western GOM. In addition to changes in spring wind patterns, higher spring sea surface temperatures have been associated with shifting distributions of a range of fish species. The abundance of several pelagic piscivores, which based on their feeding habits may predate on salmon post‐smolts, has increased in the areas that serve as migration corridors for post‐smolts. In particular, populations of silver hake, Merluccius bilinearis (Mitchell), red hake, Urophycis chuss (Walbaum), and spiny dogfish, Squalus acanthias L., increased in size in the portion of the GOM used by post‐smolts. Climate variation and shifting predator distributions in the GOM are consistent with the predator hypothesis of recruitment control suggested for the stock complex.

Gulf Of Maine

The behavioural response of adult Petromyzon marinus to damage-released alarm and predator cues

Using semi‐natural enclosures, this study investigated (1) whether adult sea lamprey Petromyzon marinus show avoidance of damage‐released conspecific cues, damage‐released heterospecific cues and predator cues and (2) whether this is a general response to injured heterospecific fishes or a specific response to injured P. marinus . Ten replicate groups of 10 adult P. marinus , separated by sex, were exposed to one of the following nine stimuli: deionized water (control), extracts prepared from adult P. marinus , decayed adult P. marinus (conspecific stimuli), sympatric white sucker Catostomus commersonii , Amazon sailfin catfish Pterygoplichthys pardalis (heterospecific stimuli), 2‐phenylethylamine (PEA HCl) solution, northern water snake Nerodia sipedon washing, human saliva (predator cues) and an adult P. marinus extract and human saliva combination (a damage‐released conspecific cue and a predator cue). Adult P. marinus showed a significant avoidance response to the adult P. marinus extract as well as to C. commersonii , human saliva, PEA and the adult P. marinus extract and human saliva combination. For mobile P. marinus , the N. sipedon washing induced behaviour consistent with predator inspection. Exposure to the P. pardalis extract did not induce a significant avoidance response during the stimulus release period. Mobile adult female P. marinus showed a stronger avoidance behaviour than mobile adult male P. marinus in response to the adult P. marinus extract and the adult P. marinus extract and human saliva combination. The findings support the continued investigation of natural damage‐released alarm cue and predator‐based repellents for the behavioural manipulation of P. marinus populations in the Laurentian Great Lakes.

Journal of Fish Biology

Predator avoidance ability of juvenile chinook salmon ( Oncorhynchus tshawytscha ) subjected to sublethal exposures of gas-supersaturated water

To assess the effects of gas bubble trauma (GBT) on the predator avoidance ability of juvenile chinook salmon ( Oncorhynchus tshawytscha ), we created groups of fish that differed in prevalence and severity of gas emboli in their lateral lines, fins, and gills by exposing them to 112% total dissolved gas (TDG) for 13 days, 120% TDG for 8 h, or 130% TDG for 3.5 h. We subjected exposed and unexposed control fish simultaneously to predation by northern squawfish ( Ptychocheilus oregonensis ) in water of normal gas saturation in 6, 18, and 10 tests using prey exposed to 112, 120, and 130% TDG, respectively. Only fish exposed to 130% TDG showed a significant increase in vulnerability to predation. The signs of GBT exhibited by fish sampled just prior to predator exposure were generally more severe in fish exposed to 130% TDG, which had the most extensive occlusion of the lateral line and gill filaments with gas emboli. Fish exposed to 112% TDG had the most severe signs of GBT in the fins. Our results suggest that fish showing GBT signs similar to those of our fish exposed to 130% TDG, regardless of their precise exposure history, may be more vulnerable to predation.

Canadian Journal of Fisheries and Aquatic Sciences

The effects of hydropattern and predator communities on amphibian occupancy

Complex, interactive ecological constraints regulate species distributions, and understanding these factors is crucial for predicting species persistence. We used occupancy analysis, which corrects for imperfect detection, to test the importance of abiotic and biotic habitat and landscape factors on probability of occupancy by Boreal Chorus Frog ( Pseudacris maculata (Agassiz, 1850)) tadpoles. We hypothesized that hydropattern and predators are primarily important because they affect desiccation and predation risk and can interact in ways difficult to predict. We surveyed 62 wetland sites across an elevational gradient in Colorado, USA, and modeled patterns in P. maculata occupancy. Tadpoles were most frequently present in intermediate-length hydropattern systems with lower desiccation risk and no predatory fish because of occasional drying. Pseudacris maculata occupancy had a strong negative relationship with fish presence, while tadpoles, odonate larvae, and Barred Tiger Salamanders ( Ambystoma mavortium mavortium Baird, 1850) frequently co-occurred. Dry seasonal conditions will likely result in fewer intermediate-length hydropattern ponds available for amphibian breeding. We hypothesize that this will force P. maculata to breed in habitats with fish. As habitats shrink, predators that co-occur with P. maculata are expected to concentrate in the remaining habitat and increase predation risk for developing tadpoles (assuming predators are similarly constricted in their habitat use as amphibians are).

Colorado