Habitat selection responses of parents to offspring predation risk: an experimental test

The ability of nest predation to influence habitat settlement decisions in birds is widely debated, despite its importance in limiting fitness. Here, we experimentally manipulated nest predation risk across a landscape and asked the question, do migratory birds assess and respond to variation in nest predation risk when choosing breeding habitats? We examined habitat preference by quantifying the density and settlement date of eight species of migratory passerines breeding in areas with and without intact nest predator communities. We found consistently more individuals nesting in areas with reduced nest predation than in areas with intact predator assemblages, although predation risk had no influence on settlement or breeding phenology. Additionally, those individuals occupying safer nesting habitats exhibited increased singing activity. These findings support a causal relationship between habitat choice and nest predation risk and suggest the importance of nest predation risk in shaping avian community structure and breeding activity.     

Spatial heterogeneity in predator activity, nest survivorship, and nest-site selection in two forest thrushes

The ability of prey to find and use predator-free space has far-reaching consequences for their persistence and interactions with their predators. We tested whether nest survivorship of the ground-nesting veery (Catharus fuscescens) and shrub-nesting wood thrush (Hylocichla mustelina) was related to the local absence of a major nest predator, the white-footed mouse (Peromyscus leucopus). Mouse-free space was defined by trap stations that failed to trap a mouse during the avian breeding season (~May through July). In addition, mouse activity was quantified at individual trap stations based on the number of captures during the same period (six 2.25-ha trapping grids, each containing 121 trap stations trapped repeatedly throughout the summer between 1998 and 2002.) Annual mouse-free space was correlated with other measures of mouse activity based on trapping data. Both mouse-free space and activity metrics were significantly related to annual rates of nest predation (i.e., nest daily mortality rate) in veery but not wood thrush. Likewise, mouse-free space and mouse activity within the nest neighborhood (~30×30 m² surrounding each nest) was significantly related to nest survivorship in veery but not wood thrush. More trap stations had consistently greater (hotspots) and lesser (coldspots) mouse activity than expected by chance, and veeries were significantly more likely to nest near stations that had below the grid-average trapping success. Our study thus documented significant spatial variability in predator activity and its relationship to nest predation and nest-site selection in a ground-nesting songbird.     

Nest predation patterns in ground-nesting passerines on the Iberian Peninsula

The idea that ground-nesting birds have a high risk of predation is widely accepted This paper analyzes the nest predation patterns of ground-nesting passerines on the Iberian Peninsula The nest predation rates are higher m open land species x= 71%) than m countryside farmland (10%) or forest birds (29%) The latter species have no differences in daily survival between ground and above-ground nests The higher nest predation in ground-nesting open land species was similar m peninsular Spain and North America We conclude that it is incorrect to generalize that all ground nesters have high nest predation rates, and we discuss the possible link of the high nest predation rates of open land birds to the decline observed in their population trends in Spain     

Fear factor: prey habitat selection and its consequences in a predation risk landscape

Predation risk influences prey use of space. However, little is known about how predation risk influences breeding habitat selection and the fitness consequences of these decisions. The nest sites of central-place foraging predators may spatially anchor predation risk in the landscape. We explored how the spatial dispersion of avian predator nests influenced prey territory location and fitness related measures. We placed 249 nest boxes for migrant pied flycatchers Ficedula hypoleuca , at distances between 10 and 630 m, around seven different sparrowhawk nests Accipiter nisus . After closely monitoring flycatcher nests we found that flycatcher arrival dates, nest box occupation rates and clutch size showed a unimodal relationship with distance from sparrowhawk nests. This relationship suggested an optimal territory location at intermediate distances between 330 and 430 m from sparrowhawk nests. Furthermore, pied flycatcher nestling quantity and quality increased linearly with distance from sparrowhawk nests. These fitness related measures were between 4 and 26% larger in flycatcher nestlings raised far from, relative to those raised nearby, sparrowhawk nests. Our results suggest that breeding sparrowhawk affected both flycatcher habitat selection and reproductive success. We propose that nesting predators create predictable spatial variation in predation risk for both adult prey and possibly their nests, to which prey individuals are able to adaptively respond. Recognising predictable spatial variation in perceived predation risk may be fundamental for a proper understanding of predator-prey interactions and indeed prey species interactions.     

Consequences of information use in breeding habitat selection on the evolution of settlement time

The role of temporal changes and spatial variability in predation risk and prey's means of mitigating such risks is poorly understood in the context of potential threats of global climate change for migratory birds. Yet nest predation, for example, represents a primary source of reproductive mortality in birds. To assess risk birds must spend time prospecting potential breeding sites for cues or signals of predator presence. However, competition for breeding sites with advantage to prior residency poses an evolutionary dilemma as individuals also benefit from early settling. We develop a model to examine adaptive prospecting time for predator cues on breeding grounds characterized by spatial heterogeneity in nest predation risk. We study how populations respond to environmental change represented by variation in habitat specific levels of nest predation, habitat composition, population vital rates, and availability of information (via prospecting) in the form of acoustic predator cues. We identify two mechanisms that regulate and buffer impacts of environmental change on populations. First, the adaptive response to lower population abundance under deteriorating environmental conditions is to increase prospecting time, which in turn increases individuals nest success to counteract population declines. This occurs because reduced competition for sites decreases the benefit of early settlement. Second, per capita success in site choice increases during population declines owing to reduced competition that increases the availability of good sites. We also show that the increased benefit to settling early when competition increases can lead to the paradoxical result that with greater spatial heterogeneity, less effort is placed on discerning good and bad sites. Our analysis thus contributes several novel results by which nest predation, settlement phenology, prospecting time and information gathering can influence species capacity to adapt to changing environments.     

Adaptive plasticity in nest-site selection in response to changing predation risk

Birds selecting a nest site have to find the best compromise between the risk of encountering predation, the availability of food near to the nest and microclimatic requirements. As the optimal solution of this problem will vary with changes in predator abundance, we ask whether birds are capable of assessing such changes and of adjusting their nest-site choice accordingly. The reproductive success of dusky warblers Phylloscopus fuscatus , breeding in a mosaic of bushland and tundra habitat in the Russian Far East, varied greatly depending on the abundance of a nest predator, the Siberian chipmunk Tamias sibiricus . Using artificial nests we analysed which strategies dusky warblers should follow to avoid nest predation by chipmunks. We then compared the nest sites which dusky warblers actually had chosen in years with very high chipmunk densities (1998 and 1999) with those chosen in 1997, when chipmunks were almost absent from the study area. We found that safe nest-sites were preferred over those offering other advantages (microclimate, proximity to food) when the risk of predation was high, and we could not detect any confounding factor that might alternatively have caused these striking between-year differences. Our study suggests that even a short-lived passerine may be capable of choosing its nest site according to the actual predation risk. We show that such behavioural plasticity can lead to a paradoxical situation where better-protected nest-sites (selected in years and areas with high risk), on average, suffer greater predation than sites offering low safety. Thus, behavioural plasticity, if undetected, may result in serious misinterpretation of nest-predation patterns. A review of the literature suggests that adaptive plasticity in nest placement may be more widespread than is currently recognized.     

Nesting Success in Crimson Finches: Chance or Choice?

In avian systems, nest predation is one of the most significant influences on reproductive success. Selection for mechanisms and behaviours to minimise predation rates should be favoured. To avoid predation, breeding birds can often deter predators through active nest defence or by modifying behaviours around the nest (e.g. reducing feeding rates and vocalisations). Birds might also benefit from concealing nests or placing them in inaccessible locations. The relative importance of these strategies (behaviour vs. site selection) can be difficult to disentangle and may differ according to life history. Tropical birds are thought to experience higher rates of predation than temperate birds and invest less energy in nest defence. We monitored a population of crimson finches (Neochmia phaeton), in the Australian tropics, over two breeding seasons. We found no relationship between adult nest defence behaviour (towards a model reptile predator) and the likelihood of nest success. However, nest success was strongly related to the visibility of the nest and the structure of the vegetation. We found no evidence that adult nest building decisions were influenced by predation risk; individuals that re‐nested after a predation event did not build their nest in a more concealed location. Therefore, predator avoidance, and hence nest success, appears to be largely due to chance rather than due to the behaviour of the birds or their choice of nesting sites. To escape high predation pressures, multiple nesting attempts both within and between seasons may be necessary to increase reproductive success. Alternatively, birds may be limited in their nest‐site options; that is, high‐quality individuals dominate quality nest sites.     

Increasing temperatures increase the risk of reproductive failure in a near threatened alpine ground-nesting bird,the Cape Rockjumper Chaetops frenatus

A major cause of reproductive failure in birds is nest predation. Predation risk depends on predator type, as predators vary in their ecology and sensory modalities (e.g. visual vs. olfactory). Snakes (generally olfactory predators) are a major nest predator for small birds, with predation strongly associated with higher temperatures. We investigated nest survival in a ground-nesting alpine species, the Cape Rockjumper Chaetops frenatus, endemic to alpine fynbos in southwestern South Africa. We collected 3 years of nest data, testing whether nest survival was related to (1) habitat stage (early post-fire vs. late post-fire habitat, ≤ 3 and > 3 years since fire respectively), (2) nest concealment and (3) temperature. We found that nests had better survival at lower temperatures, with snake predation (our main source of predation) increasing in higher temperatures.     

EVOLUTION OF AVIAN COOPERATIVE BREEDING: COMPARATIVE TESTS OF THE NEST PREDATION HYPOTHESIS

Comparative analyses carried out on two different phylogenies of cooperatively and noncooperatively breeding Australian passerine birds (parvorder Corvida) were unable to detect a significant difference in nest predation rates after controlling for body mass and risk of predation due to location of the nest (nest safety). Nest predation rates, however, decrease as nest safety and body mass increase. We suggest that cooperative breeding does not bring about a current net change in rates of nest predation among Australian passerines. Species breeding cooperatively may have developed antipredator strategies that produce results similar to those adopted by noncooperatively breeding species. The function of cooperative breeding may lie outside of antipredator strategies.     

Predators and the breeding bird: behavioral and reproductive flexibility under the risk of predation

A growing body of work suggests that breeding birds have a significant capacity to assess and respond, over ecological time, to changes in the risk of predation to both themselves and their eggs or nestlings. This review investigates the nature of this flexibility in the face of predation from both behavioural and reproductive perspectives, and also explores several directions for future research. Most available work addresses different aspects of nest predation. A substantial change in breeding location is perhaps the best documented response to nest predation, but such changes are not always observed and not necessarily the best strategy. Changes in nesting microhabitat (to more concealed locations) following predation are known to occur. Surprisingly little work addresses the proactive avoidance of areas with many nest predators, but such avoidance is probably widespread. Individual birds could conceivably adopt anti‐predator strategies based on the nest predators actually present in an area, but such effects have yet to be demonstrated. In fact, the ways in which birds assess the risk of nest predation is unclear. Nest defence in birds has historically received much attention, but little is known about how it interacts with other aspects of decision‐making by parents. Other studies concentrate on predation risk to adults. Some findings suggest that risk to adults themselves influences territory location, especially relative to raptor nests. An almost completely unexplored area concerns the sorts of social protection from predators that might exist during the breeding season. Flocking typical of the non‐breeding season appears unusual while breeding, but a mated pair may sometimes act as a “flock of two”. Opportunistic heterospecific sociality may exist, with heterospecific protector species associations more prevalent than currently appreciated. The dynamics of singing during the breeding season may also respond to variation in predation risk, but empirical research on this subject is limited. Furthermore, a few theoretical and empirical studies suggest that changes in predation risk also influence the behaviour of lekking males. The major influence of predators on avian life histories is undoubtedly expressed at a broad phylogenetic scale, but several studies hint at much flexibility on an ecological time scale. Some species may forgo breeding completely if the risk of nest predation is too high, and a few studies document smaller clutch sizes in response to an increase in nest predation. Recent evidence suggests that a female may produce smaller eggs rather than smaller clutches following an increase in nest predation risk. Such an increase may also influence decisions about intraspecific brood parasitism. There are no clear examples of changes in clutch/egg size with changes in risk experienced by adults, but parental responses to predators have clear consequences for offspring fitness. Changes in risk to adults may also influence body mass changes across the breeding season, although research here is sparse. The topics highlighted herein are all in need more empirical attention, and more experimental field work whenever feasible.     

Factors affecting nest predation on forest songbirds in North America

Nest predation is an important factor in the ecology of passerines and can be a large source of mortality for birds. I provide an overview of factors affecting nest predation of passerines in North America with the goal that it may provide some insight into the ecology and management of woodland birds in the United Kingdom. Although several factors influence productivity, nest success is perhaps the most widely measured demographic characteristic of open-cup-nesting birds, and nest predation is usually the largest cause of nest failure. The identity of predator species, and how their importance varies with habitat and landscape factors, must be known for managers and scientists to design effective conservation plans and place research on nest predation in the appropriate context. Recent studies using video surveillance have made significant contributions to our understanding of the relative importance of different predator taxa in North America. Spatial and temporal variation in nest predation can be better understood when landscapes are placed in a biogeographical context and local habitat and nest-site effects are placed in a landscape context. Low productivity resulting from high nest predation is one of several potential causes of bird population declines in North America and the UK. Although the 'forest fragmentation paradigm' from the eastern US may not apply directly to the UK, thinking about avian demographics from a multiscale perspective, and consideration of factors affecting nest predation with knowledge of the dominant predator species, may provide insight into population declines.     

Management of reedbeds: mosaic reed cutting does not affect prey abundance and nest predation rate of reed passerine birds

Reed passerine birds are strict habitat specialists inhabiting reedbed habitats. In Europe, many of these species are threatened due to loss and degradation of natural reedbeds. Another important factor that can negatively affect the abundance of reed passerines is commercial reed harvesting. Previous studies have shown negative impacts of large-scale winter reed cutting on passerine breeding assemblages and arthropod communities. The effect of reed cutting on a small scale, however, has not been studied experimentally to date. The aim of this study was to investigate whether and how small-scale, mosaic reed cutting influences prey abundance and nest predation rate of reed passerines. In June, after the reed had reached maturity, we conducted nest predation experiments with artificial nests and arthropod sampling using pan traps in cut reed patches, adjacent uncut reed patches and unmanaged reedbed. We found no differences in the risk of egg predation between three types of reedbeds. In contrast, the abundance of arthropods in cut and adjacent uncut reed patches was significantly higher than that in unmanaged reedbed. We assume this was caused by habitat heterogeneity, small size of cut patches and their rapid recolonization by arthropods from adjacent uncut patches. Our results suggest that in contrast to large-scale reed cutting, small-scale, mosaic reed cutting has no negative effect on nest survival and food abundance of reed passerine birds. However, given that we performed all experiments in June, i.e., when the reed was mature, our findings cannot be generalized to whole breeding period of all reed passerine birds. Therefore, temporal variation in nest predation rate and arthropod abundance in managed and unmanaged reedbeds during the entire breeding season should be examined in future studies.     

Temporal variation in nest predation risk along habitat edges between grassland and secondary forest in Central Europe

Habitat fragmentation alters many ecological processes, including trophic cascades. For example, increased predation pressure along habitat edges has often been observed in fragmented landscapes. Here, we studied how nest predation risk varies along the transition zone between grassland and mixed forest in Central Europe. Using artificial nests, we tested the two mechanisms that are expected to underlie higher predation rates along edges: (1) the matrix effect model that supposes predator penetration from a habitat type with higher predator density to one with lower predator density and (2) the ecotonal effect model that assumes specific predator preferences for habitat edges. Although our results do not fully support either of these scenarios, our data show high temporal instability in nest predation along forest–grassland edges. Predation was higher in habitat interiors compared to edges during the first year, whereas the opposite pattern was observed during the subsequent year. In addition, dramatic between-year differences in the species composition of nest predators were observed. Therefore, we hypothesise that the effect of edges on nest predation is difficult to predict in landscapes with high predator diversity. In addition, our data indicate that a high abundance of wild boar considerably increases the risk of predation for ground-nesting birds.     

Kea Nestor notabilis mothers produce nest-specific calls with low amplitude and high entropy

Vocal behaviour of nesting altricial birds is subject to selection pressure from several sources. Offspring beg to attract parents’ attention, thus increasing the chances of being fed, but also increasing the chances of being detected by predators. Research on passerines has shown that parents may reduce the risk of nest predation by alarm calling to warn nestlings to be quiet, and by producing food calls which solicit begging when parents are present to defend the nestlings. Both nestlings and parents may reduce the risk of predator detection by producing calls of low amplitude and high entropy which are acoustically difficult to locate. Although extensive research has been undertaken on nesting passerine vocalizations, little is known about parrots in this regard, and studies are needed to determine whether parrots show similar adaptations. We investigated the calling behaviour of Kea Nestor notabilis mothers during the nesting period to determine whether maternal vocalizations were adapted in a way that could increase the chance of brood success. A microphone was installed inside the nest to record calls produced both inside the nest and in the direct vicinity. Our prediction was that calls outside the nest would be easy to locate and could function as alarm calls to alert conspecifics or distract the predator, whereas calls inside the nest would be difficult to locate and could serve to communicate with nestlings without alerting predators. Our results accorded with these predictions. Calls produced outside the nest were loud and tonal, and corresponded to previously described Kea alarm calls. Calls produced inside the nest, however, were high-entropy and low-amplitude calls, and formed a distinct structural category. We thus provide the first evidence that a parrot species has a vocal category for communication inside the nest, and that calls within this category are structured in a way that could reduce the risk of nest predation.     

The effect of predation on begging-call evolution in nestling wood warblers

I combined a comparative study of begging in ground- and tree-nesting wood warblers (Parulidae) with experimental measures of the predation costs of warbler begging calls. Throughout their development, ground-nesting warbler nestlings had significantly higher-frequency begging calls than did tree-nesting warblers. There was also a trend for ground-nesting birds to have less rapidly modulated calls. There were no consistent associations between nesting site and the amplitude of the calls. Using miniature walkie-talkies hidden inside artificial nests, I reciprocally transplanted the begging calls of 5- and 8-day-old black-throated blue warblers, Dendroica caerulescens (tree-nesting) and ovenbirds, Seiurus aurocapillus (ground-nesting) and measured the corresponding changes in rates of nest predation. For the begging calls of 8-day-old nestlings, but not those of 5-day-olds, the calls of the tree-nesting species coming from ground nests incurred greater costs than did the calls of ground nesters. The reciprocal transplant had little effect on the rate of predation. Tooth imprints on clay eggs placed in artificial nests indicated that eastern chipmunks, Tamias striatus, were responsible for the increased cost of begging for black-throated blue calls coming from the ground. These data suggest that nest predation may be responsible for maintaining some of the interspecific differences in the acoustic structure of begging calls. Copyright 1999 The Association for the Study of Animal Behaviour.     

Experimental study of alarm calls of the oriental tit (Parus minor) toward different predators and reactions they induce in nestlings

Anti-predatory strategies of birds are diverse and may include predator-specific alarm calls. For example, oriental tit (Parus minor) parents can distinguish snakes from other predators and produce snake-specific referential vocalizations ("jar" call) when a snake poses a threat to their nest. The “jar” call has a very specific function to induce fledging of nestlings close to fledging age. This reaction ensures nestlings' survival in natural encounters with snakes that are capable of entering nest cavities and kill entire broods. Sciurid rodents, like chipmunks, may pose a similar threat to cavity-nesting birds. We explored the hypothesis that parents use the fledging-inducing alarm vocalizations in this situation, because chipmunks, like snakes, can kill the brood upon entering the nest cavity. We compared alarm calls of parents toward two predators (chipmunk and snake) who pose a similar threat to the nestlings in a nest cavity, and toward an avian predator (Eurasian jay) who cannot enter nest cavities and poses no threat to the nestlings in a nest. Our results show that the vocal responses of oriental tits were different among the three predators. This suggests that the acoustic properties of vocal responses to predators are different between predators of a similar hunting strategy (nest-cavity entering). The playback of recorded vocal responses of parents to chipmunks did not trigger the fledging of old nestlings, whereas the vocalizations toward a snake did, as shown by earlier studies. Our study suggests that the vocal response of parents does not carry information about the ability of predators to enter the nest cavity and confirms the special status of alarm calls triggered by snakes.     

Non-additivity among multiple cues of predation risk: a behaviorally-driven trophic cascade between owls and songbirds

Non-lethal effects of predators on prey are initiated in the form of responses to direct and indirect cues of predation risk. Like their lethal equivalents, non-lethal effects may affect species further down the food web initiating a behaviorally-driven trophic cascade. I presented a direct cue of predation risk, owl vocalizations, to white-footed mice ( Peromyscus leucopus ) during either a new or full moon (indirect cue). Mice reduced their activity in space by nearly two-thirds in response to playbacks of owl vocalizations during a full moon. However, neither moonlight (full vs new) nor the presence/absence of owl calls had an effect on space use when each cue varied singly. Previous studies have demonstrated a tight correlation between spatial activity in mice as used in the current experiment and nest predation rates on ground-nesting birds. Because moonlight is a ubiquitous deterrent of activity in nocturnal rodents I used of long-term nesting records the veery ( Catharus fuscescens ) to test whether nest predation rates were correlated negatively with moonlight. For half the lunar cycle (∼full moon to new moon) predation rates decreased with moonlight as predicted. During the second half of the lunar cycle predation and moonlight did not correlate as expected, but this was likely due to the depletion of vulnerable nests after a period of in which predation rates were at their maximum near the full moon. These studies suggest that the non-lethal effects of predatory risk on mice (i.e. changes in space use) cascade to affect their prey. Through the mechanism of reduced space use by rodents, perceived predation risk has the potential to significantly and indirectly affect songbird nest predation rates.     

Increased nest defence of upland‐nesting ducks in response to experimentally reduced risk of nest predation

Parent birds should take greater risks defending nests that have a higher probability of success. Given high rates of mammalian nest predation, therefore, parents should risk more for nests in areas with a lower risk of mammalian predation. We tested this hypothesis using nest defence data from over 1300 nests of six species of dabbling ducks studied in an area where predation risk had been reduced through removal of mammalian predators. When predator removal reduced nest predation, the ducks increased risk taking as predicted. Also as predicted, risk taking varied inversely with body size, an index of annual survival, among species. For ducks to vary nest defence in response to variation in predation risk they must be able to assess the risk of nest predation. Because ducks modified nest defence in the breeding season immediately following predator removal, ducks may be able to assess predator abundance indirectly (e.g. by UV reflection from urine) rather than by seeing or interacting directly with the predators.     

Nest predators of Lance-tailed Manakins on Isla Boca Brava, Panamá

ABSTRACT.   Nest predation is often the primary cause of nest failure for passerines. Despite this, little is known about predation rates and the nest predators of birds in the tropics. I used video cameras to monitor seven Lance-tailed Manakin ( Chiroxiphia lanceolata ) nests on Isla Boca Brava, Panamá. One nest fledged young and six nests failed due to predation. I recorded five predation events involving four avian predators and one mammalian predator. Crested Oropendolas ( Psarocolius decumanus ) predated two nests and a Roadside Hawk ( Buteo magnirostris ) and a Black-chested Jay ( Cyanocorax affinis ) each predated one. The mammalian predator was a common opossum ( Didelphis marsupialis ). All avian predation was diurnal; the mammalian predation was nocturnal. My results suggest that tropical birds are subject to a diverse suite of nest predators, and that avian predators may be an important cause of nest failure at my study site.     

Predation as a landscape effect: the trading off by prey species between predation risks and protection benefits

1. Predators impose costs on their prey but may also provide benefits such as protection against other (e.g. nest) predators. The optimal breeding location in relation to the distance from a nesting raptor varies so as to minimize the sum of costs of adult and nest predation. We provide a conceptual model to account for variation in the relative predation risks and derive qualitative predictions for how different prey species should respond to the distance from goshawk Accipiter gentilis nests. 2. We test the model predictions using a comprehensive collection of data from northern Finland and central Norway. First, we carried out a series of experiments with artificial bird nests to test if goshawks may provide protection against nest predation. Second, we conducted standard bird censuses and nest-box experiments to detect how the density or territory occupancy of several prey species varies with distance from the nearest goshawk nest. 3. Nest predation rate increased with distance from goshawk nest indicating that goshawks may provide protection for birds' nests against nest predation. Abundance (or probability of presence) of the main prey species of goshawks peaked at intermediate distances from goshawk nests, reflecting the trade-off. The abundance of small songbird species decreased with distance from goshawk nests. The goshawk poses little risk to small songbirds and they may benefit from goshawk proximity in protection against nest predation. Finally, no pattern with distance in pied flycatcher territory (nest box) occupation rate or the onset of egg-laying was detected. This is expected, as flycatchers neither suffer from marked nest predation risk nor are favoured goshawk prey. 4. Our results suggest that territory location in relation to the nest of a predator is a trade-off situation where adult birds weigh the risk of themselves being predated against the benefits accrued from increased nest survival. Prey species appear able to detect and measure alternative predation risks, and respond adaptively. From the prey perspective, the landscape is a mosaic of habitat patches the quality of which varies according to structural and floristic features, but also to the spatial distribution of predators.