Predators of bird nests in the Neotropics: a review

Predation is the main cause of nest failure among birds and, therefore, a strong selective agent. To fully understand patterns of nest predation, determining the identities of nest predators is crucial. Information about nest predators in the Neotropics, however, is largely anecdotal and not easily accessible in the literature. Our objective was to search the literature and compile a list of the known predators of nests in the Neotropics. We identified 256 species belonging to 67 families of birds, reptiles, mammals, and arthropods as nest predators. Families with at least 10 species of identified nest predators included Colubridae, Accipitridae, Corvidae, Ramphastidae, Falconidae, Furnariidae, Icteridae, and Didelphidae. Species in the first five of these families, plus the family Cebidae, predated nests of at least 30 species of birds. Many species not included on our list are also likely nest predators, e.g., 79 species identified as nest predators in the Nearctic that also occur in the Neotropics, but have not yet been confirmed as predators there. Increased use of video technology in the future should lead to an increase in the numbers of nest predators identified, particularly those that are nocturnal. By determining which species on our list occur in a given study area, researchers can now consider the likely nest predators in their study areas when designing hypotheses and conservation plans.     

Nest excavation does not reduce harmful effects of ectoparasitism: an experiment with a woodpecker, the northern flicker Colaptes auratus

It has been hypothesized that it is adaptive for birds to build new nests annually in order to avoid the accumulation of ectoparasites. Previous studies on costs of ectoparasitism have focused on cavity nesters in nestboxes while largely ignoring reproductive consequences in natural tree cavity nests, the context where nest selection strategies presumably evolved. To see whether ectoparasitism could be a driving selective force in the evolution of nest excavation in a woodpecker, I experimentally fumigated a subset of freshly excavated cavities and a subset of reused cavities of the northern flicker Colaptes auratus and compared reproductive performance with a set of control nests. The main ectoparasite of nestlings, a blood-sucking fly Carnus hemapterus , may have appeared one or two days earlier in reused nests but there was no difference between fresh and reused nests in intensities of flies one week post-hatching. Prevalence of parasitism reached 100% in both reused and freshly-excavated control nests in the second week. Nestlings from control nests had lower body mass residuals than those from fumigated nests after 15 d and fledged at a lower weight, suggesting that ectoparasitism by C. hemapterus was costly. However, fresh nest construction was no benefit, likely because the high dispersal ability of the ectoparasite meant all nests were colonized rapidly. Parents did not adjust provisioning effort according to parasitism as delivery rates did not differ between control and fumigated nests but delivery rates increased with brood size.     

Nest height, nest concealment, and predator type predict nest predation in superb fairy-wrens (Malurus?cyaneus)

Three factors and their interaction effects are increasingly recognized as important determinants of nest predation: nest concealment, nest height, and predator type. The risk of nest predation is predicted to vary across these variables because of nest detectability and accessibility. In general, however, few studies examine how these three variables interact in relation to nest predation, focusing instead on either nest concealment or nest height (whereby predator identity is usually not known). In this study, we examine the role of nest concealment and nest height for nest survival using both artificial and natural nests in the superb fairy-wren (Malurus cyaneus). We indirectly identified potential predators through marks left on artificial eggs and footprints left on tracking tunnels. Predation level at artificial nests was lower than at natural nests, and this could be due to a failure of some nest predators to locate cryptic nests in the absence of cues provided by parental activity. Our results supported the prediction that exposed and concealed nests have different levels of nest predation, which can be explained by variation in predator type. Visual predators were only detected at exposed nests, and survival from visual predators was lower for high nests that were also exposed. However, olfactory predators were detected irrespective of nest height or nest concealment. Because rodents use olfaction to locate nests, this could explain the lack of association between nest concealment and predation outcome at low nests. In addition, rodent footmarks near nests were significantly associated with rodent tooth marks on eggs.     

Alternative nesting behaviours following colonisation of a novel environment by a passerine bird

When birds settle in new environments, they may encounter different suites of nest-predators, resulting in strong selection to nest in novel locations. An important axis of variation is in height of nest placement, because off-ground nests are likely to be subject to different predation pressures than on-ground nests. In their native habitats, i.e. temperate montane forest, dark-eyed juncos Junco hyemalis predominantly nest on the ground. A population of juncos became established on the urban campus of the Univ. of California at San Diego in the early 1980s, and now has a relatively high frequency (∼20%) of off-ground nests. Off-ground nesting is associated with strong fitness benefits: based on returns from 579 nests, an egg laid off the ground has ∼80% higher chance of producing a recruit to the next generation than one laid on the ground. In addition, some nests were occasionally re-used by ∼10% of females and off-ground nests were more likely to be re-used. Many females that bred off-ground also built an on-ground nests and we did not detect an association in nest site positions of mothers and daughters, implying low heritability of this novel nest placement trait. By contrast, certain territories consistently had on-ground nests irrespective of the identity of the breeding female. We suggest that a limitation of suitable off-ground nest sites for a species predominantly adapted to on-ground nesting greatly slows the rate of evolution of this trait.     

Nest orientation of the Spotted Barbtail, <Emphasis Type="Italic">Premnoplex brunnescens</Emphasis>, is strongly correlated with stream flow

The placement and orientation of bird nests may strongly influence reproductive success. For many species, nest orientation is related to the cardinal directions and has to do with prevailing winds or solar exposure. Nest orientation differs among species in different environments, variously cooling or warming nests to maintain a nest microclimate optimal for successful development of eggs and young. The Spotted Barbtail (Premnoplex brunnescens) builds mossy, enclosed nests along shaded streams in Neotropical cloud forests. It shows a unique pattern of nest orientation, whereby nests are oriented in relation to the direction of water flowing below the nest entrance, rather than compass bearing. Nests face in one of three directions in relation to the stream: downstream, upstream, or perpendicular to the flow of the water. I discuss the ability of various hypotheses to explain this pattern in the context of factors affecting nest orientation in other species. While the reasons for this pattern of orientation remain unclear, orientation is likely driven by the need to create and maintain an optimal internal nest microclimate.     

Common waxbills use carnivore scat to reduce the risk of nest predation

For many animals, nest construction is a prerequisite for successfulbreeding. The choice of nesting materials is an important componentof nest construction, because material properties can influencenest design and, potentially, reproductive success. Common waxbillsare small African finches that select carnivore scat as a materialto include in, on, and around their nests. I investigated thehypothesis that scat functions to reduce predation risk by documentingits use in a wild population of common waxbills and by conductingan artificial nest experiment. Among natural nests, scat waspresent in every nest that hatched young, and parents continuedto add scat to nests throughout the nestling period. Among artificialnests, those that received experimental additions of carnivorescat survived at a significantly higher rate than did untreatednests, suggesting that scat functions to reduce predation risk.The mechanism by which nests are protected remains unclear,although it is likely that scat acts as an olfactory deterrentand/or camouflage. Researchers have long focused on the implicationsof nest site characteristics for avian life-history evolution.Results of the present study suggest that nest materials, similarto nest sites, may influence life histories of nest-buildinganimals by affecting predation risk.     

Threat-sensitive Nest Preferences in Male Upland Bullies,Gobiomorphus breviceps

A male's ability to acquire and defend a nest site against competitors is an important requirement for successful breeding in many fish species. In the upland bully Gobiomorphus breviceps (Eleotridae), not all males are successful in gaining ownership of nesting rocks; therefore, male-male competition may be important in nest acquisition. Although larger nests have the potential to hold more eggs, in nature there is a preferred nest size well below the maximum available. I found that in over 88% of dyadic contests, the larger male was successful in acquiring the single nesting site provided. When males were individually given a choice of two artificial nest rocks, they consistently preferred the larger nest. These results suggest that larger males have the ability to occupy the largest nests. In the presence of a trout predator, males preferred nests with three closed sides 93% of the time. This preference disappeared when the threat was a conspecific. This shows that while males were choosing the larger nests their decisions were also threat-sensitive.     

Nest sites as limiting resources for cavity‐nesting birds in mature forest ecosystems: a review of the evidence

ABSTRACT Assumptions that populations of cavity‐nesting birds are limited by access to nest sites have largely been based on anecdotal reports or correlative data. Nest‐box‐addition experiments or tree‐cavity‐blocking experiments are potentially rigorous ways to investigate how densities of breeding birds are affected by access to nest cavities. Experimental evidence indicates that natural tree holes are limited in human‐altered landscapes, but the possibility that cavity nests are limited in old growth (unmanaged) forests is less clear. I reviewed 31 nest‐cavity‐removal or addition experiments conducted with 20 species of cavity‐nesting birds in mature forests. Of these 31 experiments conducted with a variety of different species of birds, only 19% reported statistically significant changes in breeding densities. However, none of these studies included data about the reproductive history of individuals colonizing the boxes (i.e., whether birds using the boxes would have otherwise been floaters or that birds excluded from blocked cavities on the plots did not simply move elsewhere), so they provided no strong evidence that the number of breeding pairs was limited by availability of nest sites at the population scale. Although some studies indicate that nest sites are limited at local (plot) scales in old growth forests, there is still little empirical evidence for nest‐site limitation at the population‐ and landscape‐level in mature, unmanaged forests. I review the challenges in designing and interpreting box‐addition experiments and highlight the main gaps in knowledge that should be targeted in the future.     

A predator's perspective of nest predation: predation by red squirrels is learned,not incidental

Nest predation has been used to explain aspects of avian ecology ranging from nest site selection to population declines. Many arguments rely on specific assumptions regarding how predators find nests, yet these predatory mechanisms remain largely untested. Here we combine artificial nest experiments with behavioural observations of individual red squirrels Tamiasciurus hudsonicus to differentiate between two common hypotheses: predation is incidental versus learned. Specifically, we tested: 1) whether nest survival could be explained solely by a squirrel's activity patterns or habitat use, as predicted if predation was incidental; or 2) if predation increased as a squirrel gained experience preying on a nest, as predicted if predation was learned. We also monitored squirrel activity after predation to test for evidence of two search mechanisms: area‐restricted searching and use of microhabitat search images. Contrary to incidental predation and in support of learning, squirrels did not find nests faster in areas with high use (e.g. forest edges). Instead, survival of artificial nests was strongly related to a squirrel's prior experience preying on artificial nests. Experience reduced nest survival times by over half and increased predation rates by 150–200%. Squirrels returned to and doubled their activity at the site of a previously preyed on nest. However, neither area‐restricted searching nor microhabitat search images can explain how squirrels located artificial nests more readily with experience. Instead, squirrels likely used cues associated with the nests or eggs themselves. Learning implies that squirrels could be increasingly effective predators as the density or profitability of nests increases. Our results add support to the view that nest predation is complex and broadly influenced (e.g. by predator experience, motivation), and is unlikely to be predicted consistently by simple relationships with predator activity, abundance or habitat.     

Beach erosion and nest site selection by the leatherback sea turtle Dermochelys coriacea (Testudines: Dermochelyidae) and implications for management practices at Playa Gandoca, Costa Rica

Leatherback sea turtles (Dermochelys coriacea) nest on dynamic, erosion-prone beaches. Erosive processes and resulting nest loss have long been presumed to be a hindrance to clutch survival. In order to better understand how leatherbacks cope with unstable nesting beaches, I investigated the role of beach erosion in leatherback nest site selection at Playa Gandoca, Costa Rica. I also examined the potential effect of nest relocation, a conservation strategy in place at Playa Gandoca to prevent nest loss to erosion, on the temperature of incubating clutches. I monitored changes in beach structure as a result of erosion at natural nest sites during the time the nest was laid, as well as in subsequent weeks. To investigate slope as a cue for nest site selection, I measured the slope of the beach where turtles ascended from the sea to nest, as well as the slopes at other random locations on the beach for comparison. I examined temperature differences between natural and relocated nest sites with thermocouples placed in the sand at depths typical of leatherback nests. Nests were distributed non-randomly in a clumped distribution along the length of the beach and laid at locations that were not undergoing erosion. The slope at nest sites was significantly different than at randomly chosen locations on the beach. The sand temperature at nest depths was significantly warmer at natural nest sites than at locations of relocated nests. The findings of this study suggest leatherbacks actively select nest sites that are not undergoing erosive processes, with slope potentially being used as a cue for site selection. The relocation of nests appears to be inadvertently cooling the nest environment. Due to the fact that leatherback clutches undergo temperature-dependent sex determination, the relocation of nests may be producing an unnatural male biasing of hatchlings. The results of this study suggest that the necessity of relocation practices, largely in place to protect nests from erosion, should be reevaluated to ensure the proper conservation of this critically endangered species.     

Influence of olfactory and visual cover on nest site selection and nest success for grassland‐nesting birds

Habitat selection by animals is influenced by and mitigates the effects of predation and environmental extremes. For birds, nest site selection is crucial to offspring production because nests are exposed to extreme weather and predation pressure. Predators that forage using olfaction often dominate nest predator communities; therefore, factors that influence olfactory detection (e.g., airflow and weather variables, including turbulence and moisture) should influence nest site selection and survival. However, few studies have assessed the importance of olfactory cover for habitat selection and survival. We assessed whether ground‐nesting birds select nest sites based on visual and/or olfactory cover. Additionally, we assessed the importance of visual cover and airflow and weather variables associated with olfactory cover in influencing nest survival. In managed grasslands in Oklahoma, USA, we monitored nests of Northern Bobwhite (Colinus virginianus), Eastern Meadowlark (Sturnella magna), and Grasshopper Sparrow (Ammodramus savannarum) during 2015 and 2016. To assess nest site selection, we compared cover variables between nests and random points. To assess factors influencing nest survival, we used visual cover and olfactory‐related measurements (i.e., airflow and weather variables) to model daily nest survival. For nest site selection, nest sites had greater overhead visual cover than random points, but no other significant differences were found. Weather variables hypothesized to influence olfactory detection, specifically precipitation and relative humidity, were the best predictors of and were positively related to daily nest survival. Selection for overhead cover likely contributed to mitigation of thermal extremes and possibly reduced detectability of nests. For daily nest survival, we hypothesize that major nest predators focused on prey other than the monitored species’ nests during high moisture conditions, thus increasing nest survival on these days. Our study highlights how mechanistic approaches to studying cover informs which dimensions are perceived and selected by animals and which dimensions confer fitness‐related benefits.     

Competition, predation and nest niche shifts among tropical cavity nesters: ecological evidence

I studied cavity-nesting birds in an undisturbed site in lowland Peru to determine the relative roles of competition and predation in favoring termitarium nesting over tree cavity nesting. Occupancy rates of both nest boxes and natural tree cavities near 2% suggest that competition for tree cavities is not favoring the use of termitaria. Artificial nests and bird nests in termitaria suffer significantly lower predation rates than similar nests in old tree cavities showing that predation is favoring the use of termitaria over old tree cavities. Bird nests in newly excavated tree cavities also show lower predation rates than older cavities suggesting that cavity age is more important than substrate (tree or termitaria) per se . This study suggests that nest predation has a greater influence than nest competition on life history evolution for many cavity-nesting birds.     

Density-dependent nest predation in waterfowl: the relative importance of nest density versus nest dispersion

When nest predation levels are very high or very low, the absolute range of observable nest success is constrained (a floor/ceiling effect), and it may be more difficult to detect density-dependent nest predation. Density-dependent nest predation may be more detectable in years with moderate predation rates, simply because there can be a greater absolute difference in nest success between sites. To test this, we replicated a predation experiment 10 years after the original study, using both natural and artificial nests, comparing a year when overall rates of nest predation were high (2000) to a year with moderate nest predation (2010). We found no evidence for density-dependent predation on artificial nests in either year, indicating that nest predation is not density-dependent at the spatial scale of our experimental replicates (1-ha patches). Using nearest-neighbor distances as a measure of nest dispersion, we also found little evidence for "dispersion-dependent" predation on artificial nests. However, when we tested for dispersion-dependent predation using natural nests, we found that nest survival increased with shorter nearest-neighbor distances, and that neighboring nests were more likely to share the same nest fate than non-adjacent nests. Thus, at small spatial scales, density-dependence appears to operate in the opposite direction as predicted: closer nearest neighbors are more likely to be successful. We suggest that local nest dispersion, rather than larger-scale measures of nest density per se, may play a more important role in density-dependent nest predation.     

Interspecific nest destruction in the Tropical Kingbird Tyrannus melancholicus: kleptoparasitism or competition?

Bird nest destruction and nest material kleptoparasitism (i.e., the theft of nest material from other bird’s nests) are poorly documented behaviors, and little is known about the parasite species and their hosts. Here, I present the first account of nest material kleptoparasitism in the Tropical Kingbird (Tyrannus melancholicus) followed by nest destruction, which occurred on a Variegated Flycatcher nest (Empidonomus varius). I explore the implications of these behaviors for both the kleptoparasite and the victim species and, due to the lack of basic information on the general behavior of both species, I point out some directions to guide future researches on the subject.

     

Blue tits (Cyanistes caeruleus) respond to an experimental change in the aromatic plant odour composition of their nest

Although the use of olfaction by birds is now widely recognised, the olfactory abilities of passerine birds remain poorly explored, for historical reasons. Several studies however suggest that passerines can perceive volatile compounds in several biologically relevant contexts. In Corsica, recent findings suggest that cavity-nesting blue tits may use volatile compounds in the context of nest building and maintenance. Although they build their nests mainly from moss, female blue tits also frequently incorporate fragments of several species of aromatic plants in the nest cup. In field experiments, breeding female blue tits altered their nest maintenance behaviour in response to experimental addition of aromatic plants in their nest. In aviary experiments, captive male blue tits could be trained to detect lavender odour from a distance. Here I report results from a field study aimed to test whether adult blue tits altered their chick-feeding behaviour after an experimental change in nest odour composition. I experimentally added fragments of aromatic plant species that differed from those brought in the nests before the start of the experiment in a set of experimental nests and added moss, the basic nest material, in a set of control nests. Both male and female blue tits hesitated significantly longer entering the nest cavity after addition of new aromatic plant fragments, as compared to moss addition. This response was especially observed during the first visit following the experimental change in nest plant composition. Nest composition treatment had no effect on the time spent in the nest. This study demonstrates that free-ranging blue tits detect changes in nest odour from outside the nest cavity.     

Understanding avian nest predation: why ornithologists should study snakes

Despite the overriding importance of nest predation for most birds, our understanding of the relationship between birds and their nest predators has been developed largely without reliable information on the identity of the predators. Miniature video cameras placed at nests are changing that situation and in six of eight recent studies of New World passerine birds, snakes were the most important nest predators. Several areas of research stand to gain important insights from understanding more about the snakes that prey on birds' nests. Birds nesting in fragmented habitats often experience increased nest predation. Snakes could be attracted to habitat edges because they are thermally superior habitats, coincidentally increasing predation, or snakes could be attracted directly by greater prey abundance in edges. Birds might reduce predation risk from snakes by nesting in locations inaccessible to snakes or in locations that are thermally inhospitable to snakes, although potentially at some cost to themselves or their young. Nesting birds should also modify their behavior to reduce exposure to visually orienting snakes. Ornithologists incorporating snakes into their ecological or conservation research need to be aware of practical considerations, including sampling difficulties and logistical challenges associated with quantifying snake habitat use.     

Do artificial nests reveal relative nest predation risk for real nests?

Present knowledge of the effects of nest predation on spatial distribution, habitat selection and community structure of birds is to a large extent based on results from experiments with artificial nests. Although nest predation risk is likely to differ between artificial and real nests, most previous studies of nest predation using artificial nests have been lacking a proper control. We investigated whether predation rates on artificial nests predicted those on real nests by simultaneously comparing the fate of real and artificial nests (containing quail Coturnix coturnix and plasticine eggs) in 92 territories of the northern wheatear Oenanthe oenanthe in 1996. We also investigated whether risk for artificial nests was related to relative average risk for real nests in these territories (based on data collected two years before and two years after the experiment). Nest predation on artificial nests did predict relative predation risk for real nests only when quail egg depredation was used as the criterion for artificial nest predation. Despite plasticine egg depredation being the most common type of predation it was not associated with predation risk for real nests. Small mice and vole species dominated among cases with only plasticine egg depredation, while predatory mammals and snakes destroyed most quail eggs in artificial nests and most eggs in real wheatear nests. The results suggest that artificial nests may only predict the risk for real nests when the nest predator species are similar among the two types of nest. Furthermore, our data suggest that small mice and vole species rarely depredate nests of mid-sized passerine birds . Our results cast doubt on many previous conclusions based on experiments with artificial nests, since predation risk for such nests is likely to be uncorrelated with risk for real nests due to nest-type-specific differences in nest preying species.     

A description of nesting behaviors,including factors impacting nest site selection,in black‐and‐white ruffed lemurs (Varecia variegata)

Nest site selection is at once fundamental to reproduction and a poorly understood component of many organisms’ reproductive investment. This study investigates the nesting behaviors of black‐and‐white ruffed lemurs, Varecia variegata, a litter‐bearing primate from the southeastern rainforests of Madagascar. Using a combination of behavioral, geospatial, and demographic data, I test the hypotheses that environmental and social cues influence nest site selection and that these decisions ultimately impact maternal reproductive success. Gestating females built multiple large nests throughout their territories. Of these, females used only a fraction of the originally constructed nests, as well as several parking locations as infants aged. Nest construction was best predicted by environmental cues, including the size of the nesting tree and density of feeding trees within a 75 m radius of the nest, whereas nest use depended largely on the size and average distance to feeding trees within that same area. Microhabitat characteristics were unrelated to whether females built or used nests. Although unrelated to nest site selection, social cues, specifically the average distance to conspecifics’ nest and park sites, were related to maternal reproductive success; mothers whose litters were parked in closer proximity to others’ nests experienced higher infant survival than those whose nests were more isolated. This is likely because nesting proximity facilitated communal crèche use by neighboring females. Together, these results suggest a complex pattern of nesting behaviors that involves females strategically building nests in areas with high potential resource abundance, using nests in areas according to their realized productivity, and communally rearing infants within a network of nests distributed throughout the larger communal territory.     

The Function of Hitchhiking Behavior in the Leaf‐cutting Ant Atta cephalotes1

In some leaf‐cutting ant species, minim workers ride on the fragments of leaves as they are carried back to the nest from the cutting site. There is convincing evidence that these “hitchhikers” can protect the leaf carriers from attack by phorid (Diptera: Phoridae) parasitoids, but we consider the possibility of other functions for the hitchhiking behavior. It has been hypothesized that the hitchhikers (1) feed on leaf sap from the edges of the cut leaves; (2) ride back to the nest to save energy; (3) get caught on the fragments as they are cut, and hitchhike because they cannot (or will not) get off; and (4) begin the process of preparing the leaf to enter the fungal gardens in the nest, perhaps by removing microbial contaminants. We observed hitchhikers of Atta cephalotes in 14 nests at the La Selva Biological Station in Costa Rica. There was no difference in the proportion of leaf carriers with hitchhikers between day and night. Because the nests we observed were largely nocturnal, more than 90 percent of the hitchhiking occurred at night. The phorid parasitoids are usually considered to be diurnal, so the preponderance of nocturnal hitchhiking suggests other functions in addition to parasitoid defense. Hitchhikers spent more time in the defensive head‐up posture during the day, but spent more time in the head‐down posture at night. The head‐down posture may indicate cleaning or other leaf preparation. The hitchhikers were never observed feeding on sap. Hitchhikers frequently got onto and off of the fragments, and so they were not “marooned.” Few hitchhikers rode all the way back to the nest and were often moving on the leaf fragment; these observations make the energy conservation hypothesis less likely, although we cannot reject it. We conclude that parasitoid defense is an important function of hitchhiking but also that there are probably other functions when parasitoids are absent. Based on available data, the most likely possibility is preparation of the leaf fragment before it enters the nest.     

Patterns of predator behaviour and Wood Warbler Phylloscopus sibilatrix nest survival in a primaeval forest

Understanding the foraging behaviour of predators is key to interpreting the role of anti‐predator adaptations of birds in reducing nest losses. Conducting research in primaeval habitats, with a low level of direct human interference, is particularly valuable in the understanding of predator–prey interactions. Using nest cameras, we investigated the identity and behaviour of potential and actual predators appearing at Wood Warbler Phylloscopus sibilatrix nests, and the importance of different predator groups for nest survival, in the primaeval part of Bia?owie?a Forest (Poland). Mammals formed the main predator group (30 of 32 nest depredations), particularly medium‐sized carnivores (24 of 32), which attacked nests more frequently than merely passing by. This contrasted with other species, especially small rodents, which were commonly recorded near nests but rarely attacked them. Most nest attacks (22 of 32) took place at night and nest survival did not depend on nest visibility, indicating a reduced utility of nest concealment in defence against predators using mainly sound or olfaction when hunting. Daily nest survival declined strongly with nest progression (from egg‐laying to fledging of chicks), probably due to increased predator detection of nests containing older and louder chicks, rather than to increasing parental activity at nests during the day. The set of actual nest predators differed from some previous studies in human‐transformed habitats, showing that Wood Warblers may face different threats in modified vs. near‐pristine environments.