Ultimate and proximate mechanisms underlying the occurrence of bears close to human settlements: review and management implications

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Attitudes of local people towards the mountain nyala (Tragelaphus buxtoni) in Munessa,Ethiopia

Due to their direct dependence on wildlife products for subsistent use, people living in poverty are often viewed as being in conflict with wildlife conservation. We studied the attitudes of local people towards mountain nyala (Tragelaphus buxtoni) in Munessa, Ethiopia. A household survey (n = 214) was used to examine the socio‐economic characteristics and beliefs of local people from three peasant associations and one village. Generally, the study revealed that local people had positive attitudes towards mountain nyala, its conservation and population increase, which is consistent with our predictions. For example, a greater percentage of the respondents had positive (62.5%) rather than negative (37.5%) attitudes towards mountain nyala and its conservation. Moreover, the larger proportion of the respondents had positive (66.67%) rather than negative (33.3%) attitudes towards an increase in mountain nyala population. However, compared to socio‐economic characteristics, beliefs towards mountain nyala were more powerful and consistent predictor of attitudes and explained much of the variances of the two groups of the dependent variables. Thus, informing local communities about the value of mountain nyala (e.g. recreational and economic) through conservation education, advocating the need for sustainable utilization and introducing an economic benefit sharing may improve positive attitude and increase participation of local people in conservation and management of the mountain nyala population in Munessa.     

Predators or prey? Spatio-temporal discrimination of human-derived risk by brown bears

Prey usually adjust anti-predator behavior to subtle variations in perceived risk. However, it is not clear whether adult large carnivores that are virtually free of natural predation adjust their behavior to subtle variations in human-derived risk, even when living in human-dominated landscapes. As a model, we studied resting-site selection by a large carnivore, the brown bear (Ursus arctos), under different spatial and temporal levels of human activity. We quantified horizontal and canopy cover at 440 bear beds and 439 random sites at different distances from human settlements, seasons, and times of the day. We hypothesized that beds would be more concealed than random sites and that beds would be more concealed in relation to human-derived risk. Although human densities in Scandinavia are the lowest within bear ranges in Western Europe, we found an effect of human activity; bears chose beds with higher horizontal and canopy cover during the day (0700?C1900?hours), especially when resting closer to human settlements, than at night (2200?C0600?hours). In summer/fall (the berry season), with more intensive and dispersed human activity, including hunting, bears rested further from human settlements during the day than in spring (pre-berry season). Additionally, day beds in the summer/fall were the most concealed. Large carnivores often avoid humans at a landscape scale, but total avoidance in human-dominated areas is not possible. Apparently, bears adjust their behavior to avoid human encounters, which resembles the way prey avoid their predators. Bears responded to fine-scale variations in human-derived risk, both on a seasonal and a daily basis.     

Livestock‐wildlife conflicts in the Ethiopian highlands: assessing the dietary and spatial overlap between mountain nyala and cattle

Livestock grazing is an increasing conservation challenge throughout protected wildlife areas in Africa. We report here on a negative correlation in abundance between cattle and the mountain nyala across seven study areas in the Bale Mountains of Ethiopia, and investigate if this can be explained by similarity in feeding ecology and spatial overlap. Mountain nyala foraging behaviour was studied using scan sampling of 17 GPS/VHF‐collared animals, while faecal sampling of both species was used to quantify abundance and spatial overlap. We found that female and male mountain nyala grazed during 87% and 50% of the feeding observations respectively. Although mountain nyala has earlier been classified as a browser, our findings indicate a feeding ecology closer to a grazer. The overall spatial overlap between mountain nyala and cattle was 22%, with a range of 2.4–100% among areas, and illegal cattle grazing was common. Based on the negative correlation in abundance, the similarity in feeding ecology and the relatively high spatial overlap we speculate that a competitive interaction between mountain nyala and cattle might occur. Our findings points to the potential for existence of a wildlife‐cattle conflict also in the Ethiopian highlands, as frequently reported in savannah ecosystems.     

Landscape-scale habitat response of African elephants shows strong selection for foraging opportunities in a human dominated ecosystem

Balancing trade-offs between foraging and risk factors is a fundamental behavior that structures the spatial distribution of species. For African elephants Loxodonta africana, human pressures from poaching and conflict are primary drivers of species decline, but little is known about how elephants structure their spatial behavior in the face of human occupancy and predation. We seek to understand how elephants balance trade-offs between resource access, human presence and human predatory risk factors (poaching and conflict killing) in an unfenced, dynamic ecosystem where elephants persist primarily outside protected areas in community rangelands. We used tracking data from 101 elephants collected between 2001 and 2016. We investigated elephant behavior in response to landcover, topography, productivity, water, human features and human predation risk using third-order resource selection functions. We extended this analysis by employing a mixed-effects multinomial regression to identify temporal shifts in habitat use, and evaluated temporal shifts in movement patterns by estimating mean squared displacement across different productivity periods. Across periods, elephants displayed strong selection for productive areas and areas near water. Temporal shifts in habitat use showed that, during the dry period, elephants were clustered around permanent water sources where humans also congregated. At the onset of the wet period, a shift occurred where elephants moved away from permanent water and from permanent settlements towards seasonal water sources and seasonal settlements. Our findings indicate that foraging and water access are important limiting factors affecting elephants that potentially restrain their spatial responses to humans at the scale of our analysis. Given that pastoralists and elephants rely on the same resources, increasing human and livestock populations enhance pressure on shared resources and space in Africa's drylands. The long-term conservation of elephants will require approaches that reduce poaching as well as landscape level planning to prevent negative impacts from increasing competition for preferred resources.     

Behavioural Responses of European Roe Deer to Temporal Variation in Predation Risk

In natural environments, predation risk varies over time. The risk allocation hypothesis predicts that prey is expected to adjust key anti‐predator behaviours such as vigilance to temporal variation in risk. We tested the predictions of the risk allocation hypothesis in a natural environment where both a species‐rich natural predator community and human hunters are abundant and where the differences in seasonal and circadian activity between natural and anthropogenic predators provided a unique opportunity to quantify the contributions of different predator classes to anti‐predator behaviour. Whereas natural predators were expected to show similar levels of activity throughout the seasons, hunter activity was high during the daytime during a clearly defined hunting season. According to the risk allocation hypothesis, vigilance should then be higher during the hunting season and during daytime hours than during the non‐hunting season and night‐time hours. Roe deer (Capreolus capreolus) on the edge of Bia?owie?a Primeval Forest in Eastern Poland displayed vigilance behaviour consistent with these predictions. The behavioural response of roe deer to temporarily varying predation risks emphasises the behavioural plasticity of this species and suggests that future studies of anti‐predator behaviour need to incorporate circadian variation in predation pressure as well as risk gradients of both natural and anthropogenic predators.     

Habitat overlap of enemies: temporal patterns and the role of spatial complexity

Environmental heterogeneity can promote coexistence of conflicting species by providing spatial or temporal refuges from strong interactions (e.g., intraguild predation, competition). However, in many systems, refuge availability and effectiveness may change through time and space because of variability in habitat use by either species. Here I consider how the intensity of intraguild predation risk varies from day to night for aquatic insects that use both vegetated and open water habitats. Large (1,265 l) and small (42 l) mesocosms were used to test the hypothesis that Buenoa would choose an open-water habitat that minimized predation by the ambush predator Notonecta during the day, but that at night Buenoa would safely use both vegetated and open water. Regardless of container size, Notonecta remained in vegetated water during the day and exploited both habitats at night, despite exhibiting greatest instantaneous predation rates in open water during the day. In contrast, Buenoa maintained an even distribution throughout the mesocosms during day and night, even though habitat-specific predation risks were fivefold lower in open waters than in vegetation during the day and habitat-specific predation risk would have been reduced threefold by fully exploiting open waters. Thus, temporal heterogeneity was both beneficial and detrimental to Buenoa; darkness of night reduced predation, but spatial refuges also disappeared. Together, these patterns suggest that while environmental heterogeneity can dampen intense biotic interactions, enemies do not select habitats solely on the basis of conflict avoidance. Instead, it appears that habitat-specific variation in other biotic (e.g., visual predators) or physical factors (e.g., UV radiation) may also mediate species interactions by influencing habitat selection.     

Late Pleistocene hyena skeleton remains of a communal/prey depot cave den in the Bohemian Mountains (Czech Republic) – its osteology,taphonomy and palaeoecology

A total of 366 bones are arranged to six incomplete individual skeletons (=MNI) of different age classes and taphonomic preservation from the Late Pleistocene Crocuta crocuta spelaea (Goldfuss 1823) hyena den Srbsko Chlum–Komín Cave, Czech Republic, Bohemian Mountains (Central Europe). The hyena assemblage counts no siblings, 37% cub, 33% early adult/subadult and 30% adult/senile individuals. A senile scavenged individual demonstrates the cannibalism within the cave. About 10% of the 3569 megafauna bones are from hyenas which larger amounts are typical at communal/prey depot den sites. This site is different to three other larger natal/birth and communal or prey depot hyena den cave sites in Czech Republic. The Chlum–Komín Cave hyena prey bone assemblage (=NISP) consists, similar as at two other Czech Republic sites, mainly of Equus ferus przewalskii (50%), including pregnant females being hunted in spring/early summer. The fewer amounts of Coelodonta antiquitatis (4%), Bison priscus (2%), Rangifer tarandus (15%), and alpine Rupicapra rupicapra (2%) or Capra ibex (1%) and the absence of mammoth correlate to hyena den bone assemblages in middle high elevated mountain regions of northern/central Europe. The rareness of cave bears in the Bohemian Mountains, on which hyenas specialized to feed additionally in European mountain regions, explains predominant predation on horses.     

Temporal variation in habitat selection breaks the catch‐22 of spatially contrasting predation risk from multiple predators

Predator avoidance depends on prey being able to discern how risk varies in space and time, but this is made considerably more complicated if risk is simultaneously present from multiple predators. This is the situation for an increasing number of mammalian prey species, as large carnivores recover or are reintroduced in ecosystems on several continents. Roe deer Capreolus capreolus in southern Norway illustrate a case in which prey face two predators with contrasting patterns of predation risk. They face a catch‐22 situation: spatially avoiding the risk from one predator (lynx Lynx lynx in dense habitat) implies exposure to the other (hunters in open habitat). Using GPS‐data from 29 roe deer, we tested for daily and seasonal variation in roe deer selection for habitat with respect to the habitats’ year‐round average risk level. Generally, roe deer altered their habitat selection between night and day in a pattern consistent with being able to avoid predicted risk from the nocturnal lynx during night and predicted risk from human hunters during day. However, seasonal variation in habitat selection only partially corresponded with the predicted seasonal variation in risk. Whereas roe deer avoided areas with high risk from hunters more strongly during the hunting season than in other seasons, there was a lack of selection towards areas and time periods lowering the risk of lynx predation during winter. It seems likely that the risk of starvation and thermal stress constrain roe deer habitat selection during this energetically challenging season with cold temperatures, snow cover and limited natural forage. The habitat selection pattern of roe deer fits thus only partly with the two contrasting risk gradients they face. Adjusting risk‐avoidance behavior temporally can be an adaptive response in the case of several predators whose predation patterns differ in space and time.     

Wait Until Dark? Daily Activity Patterns and Nest Predation by Snakes

Predation involves costs and benefits, so predators should employ tactics that reduce their risk of injury or death and that increase their success at capturing prey. One potential way that predators could decrease risk and increase benefits is by attacking prey at night when risks may be reduced and prey more vulnerable. Because some snakes are facultatively nocturnal and prey on bird nests during the day and night, they are ideal for assessing the costs and benefits of diurnal vs. nocturnal predation. We used automated radiotelemetry and cameras to investigate predation on nesting birds by two species of snakes, one diurnal and the other facultatively nocturnal. We predicted that snakes preying on nests at night should experience less parental nest defence and capture more adults and nestlings. Rat snakes (Pantherophis obsoletus) were relatively inactive at night (23–36% activity) but nearly always preyed on nests after dark (80% of nest predations). Conversely, racers (Coluber constrictor) were exclusively diurnal and preyed on nests during the times of day they were most active. These results are consistent with rat snakes strategically using their capacity for facultative nocturnal activity to prey on nests at night. The likely benefit is reduced nest defence because birds defended their nests less vigourously at night. Consistent with nocturnal predation being safer, rat snake predation events lasted three times longer at night than during the day (26 vs. 8 min). Nocturnal nest predation did not make nests more profitable by increasing the likelihood of capturing adults or removing premature fledging of nestlings. The disconnect between rat snake activity and timing of nest predation seems most consistent with rat snakes locating prey during the day using visual cues but waiting until dark to prey on nests when predation is safer, although designing a direct test of this hypothesis will be challenging.     

GPS tracking of the foraging movements of Manx Shearwaters Puffinus puffinus breeding on Skomer Island, Wales

We report the first successful use of miniature Global Positioning System loggers to track the ocean‐going behaviour of a c. 400 g seabird, the Manx Shearwater Puffinus puffinus. Breeding birds were tracked over three field seasons during the incubation and chick‐rearing periods on their foraging excursions from the large colony on Skomer Island, Pembrokeshire, UK. Foraging effort was concentrated in the Irish Sea. Likely foraging areas were identified to the north, and more diffusely to the west of the colony. No foraging excursions were recorded significantly to the south of the colony, conflicting with the conclusions of earlier studies based on ringing recoveries and observations. We discuss several explanations including the hypothesis that foraging may have shifted substantially northwards in recent decades. We found no obvious relationship between birds’ positions and water depth, although there was a suggestion that observations at night were in shallower water than those during the day. We also found that, despite the fact that Shearwaters can be observed rafting off‐shore from their colonies in the hours prior to making landfall at night, breeding birds are usually located much further from the colony in the last 8 h before arrival, a finding that has significance for the likely effectiveness of marine protection areas if they are only local to the colony. Short sequences of precise second‐by‐second fixes showed that movement speeds were bimodal, corresponding to sitting on the water (most common at night and around midday) and flying (most common in the morning and evening), with flight behaviour separable into erratic (indicative of searching for food) and directional (indicative of travelling). We also provide a first direct measurement of mean flight speed during directional flight (c. 40 km/h), slower than a Shearwater's predicted maximum range velocity, suggesting that birds are exploiting wave or dynamic soaring during long‐distance travel.     

Night — day drift patterns and the size of larvae of two aquatic insects

A test of Allan's (1978) hypothesis about differential drift abundance of mayflies of the genus Baetis between night and day, and the size of larvae was performed at a mountain stream in Idaho. Palisades Creek, Idaho, contains a different species of mayfly, B. tricaudatus, and vertebrate predator, Salmo clarki, than Cement Creek, Colorado (B. bicaudatus, and brook trout, Salvelinus fontinalis). Consequently it was not known if B. tricaudatus would exhibit a similar pattern as its congener in Cement Creek, with large instars tending to avoid daylight drift, as found by Allan (1978). However, similar results were observed in the present study. It appears that the earlier hypothesis may have generality for geographically distinct streams with a different vertebrate predator and mayfly prey. The existence of a similar pattern for chironomid larvae was also tested, however, no such pattern existed. This discrepency between taxa may be due to differential predation, or to inherent differences in drift abilities.     

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

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

The diet of the striped hyena in Nepal's lowland regions

Striped hyenas (Hyaena hyaena) are extremely rare in Nepal, and only a few people have studied them in their natural forest and grassland habitat. Their rarity is due to anthropogenic pressures such as hunting, habitat modification, being killed on roads, and depletion of their natural prey. Here, we studied the feeding ecology of hyenas in lowland, Nepal. We employed an opportunistic sampling to collect hyena scats in a range of habitats and the line transect sampling to identify the prey of the hyena in the study site. We collected 68 hyena scats between 2015 and 2018. Most of the hyena scat (39.7%) was found in the Churia Hill forest followed by riverbed (26.4%), mixed forest (14.7%), Sal (Shorea robusta)‐dominated forest (11.7%), and grassland area (7.3%). We found eleven mammalian prey species, plants, and some unidentified items in the hyena scats. The frequency of occurrence and relative biomass of the medium‐sized wild boar (Sus scrofa) were higher than other smaller prey species such as hare (Lepus nigricollis) and rhesus macaque (Macaca mulatta). Similarly, the proportion of large prey species such as nilgai (Boselaphus tragocamelus) in the hyena diet was lower compared with wild boar, hares, and rhesus macaques indicating medium‐sized wild boar is the most preferred prey species. Livestock contributed 17.3% of the total dietary biomass. Domesticated species such as goats, sheep, cows, and even dogs were found in the diet of hyenas. Predation of livestock by hyenas could cause conflict, especially if this ongoing issue continues in the future. Rather, more conservation effort is required in lowland areas of Nepal to protect the hyenas' natural prey species, particularly in wildlife habitats to reduce the lure of taking domestic livestock. Similarly, conservation education at the local level and active involvement of government authorities in the conservation of this species might be helpful to mitigate human–hyena conflict in the human‐dominated landscape.     

The Effect of Moonlight on Scopoli's Shearwater Calonectris diomedea Colony Attendance Patterns and Nocturnal Foraging: A Test of the Foraging Efficiency Hypothesis

Moonlight is known to affect the nocturnal behaviour and activity rhythms of many organisms. For instance, predators active at night may take advantage from increased visibility afforded by the moon, while prey might regulate their activity patterns to become less detectable. Many species of pelagic seabirds attend their colony only at night, in complete darkness, avoiding approaching their nest sites under moonlight. This behaviour has been most often interpreted as an antipredator adaptation (‘predation avoidance’ hypothesis). However, it may also reflect a lower foraging efficiency during moonlit nights (‘foraging efficiency’ hypothesis). Indeed, moonlight may reduce prey availability because preferred seabird prey is known to occur at higher depths in moonlit nights. Using high‐accuracy behavioural information from data loggers, we investigated the effect of moonlight on colony attendance and at‐sea nocturnal foraging in breeding Scopoli's shearwaters Calonectris diomedea. We found that birds departing for self‐feeding trips around the full moon performed longer trips than those departing around the new moon. On nights when the moon was present only partly, nest burrow entrances took place largely in the moonless portion of the night. Moreover, contrary to predictions from the ‘foraging efficiency’ hypothesis, nocturnal foraging activity increased according to moonlight intensity, suggesting that birds increased their foraging activity when prey became more detectable. This study strengthens the idea that colony attendance behaviour is strictly controlled by moonlight in shearwaters, which is possibly related to the perception of a predation risk.     

Great tits take greater risk toward humans and sparrowhawks in urban habitats than in forests

Urban animals often take more risk toward humans than their non‐urban conspecifics do, but it is unclear how urbanization affects behavior toward non‐human predators. Responses to humans and non‐human predators may covary due to common mechanisms enforcing a phenotypic correlation. However, while increased tolerance toward humans may be advantageous for urban animals, reduced vigilance toward non‐human predators that can pose actual threat may be costly. Therefore, urban animals may benefit from showing specific responses to different threat levels, such as humans versus non‐human predators, or hostile versus non‐hostile humans. To test these alternatives, we compared responses (latencies to return to nest) of urban and forest‐breeding great tits (Parus major) to familiar hostile and unfamiliar humans as well as one of their common predators, the sparrowhawk (Accipiter nisus). We found that urban birds were more risk‐taking toward both humans and sparrowhawk than forest birds. However, responses to sparrowhawk did not correlate with responses to humans either within or across habitats. This suggests that higher risk‐taking of urban compared to forest‐dwelling great tits toward sparrowhawk may be threat‐specific response to lower predation risk rather than a spillover effect of increased tolerance to humans. Furthermore, birds responded similarly to unfamiliar and familiar (potentially dangerous) humans in both habitats, suggesting that great tits may not adjust their risk‐taking to the threat represented by individual humans. These findings indicate that urban birds may flexibly adjust their risk‐taking to certain, but not all, types of threat.     

Human settlement and baobab distribution in south-western Mali

Aim Human settlement establishment and reproduction of the baobab tree (Adansonia digitata) appear spatially and temporally dependent because baobabs are abundant in many settlement sites in Africa. This paper tests the spatiotemporal relationship between baobab and settlement distribution. Location South‐western Mali. Methods In an area of 183 km², 1240 baobabs were located and mapped, their diameters measured, and habitat characteristics recorded for each individual. All occupied (n = 9) and abandoned (n = 84) settlements were located and mapped, and occupation dates were determined through interviews. Chi‐squared analysis indicated baobab habitat preferences, and bivariate point‐pattern analysis tested baobab–settlement spatiotemporal independence. Results Baobabs and human settlements are positively spatially associated at most distances and for all baobab size‐class–settlement age‐class pairs. However, positive spatial association is significant only at distances < 500 m, and young settlements and large baobabs are not significantly associated. Positive association between small and large baobabs is marginally significant at <300 m, but observed significance is less than that for baobab–settlement positive association. Baobab abundance is not evenly distributed across the range of habitats it occupies; recruitment is strongest in settlements and fields, and on cliffs, while mortality is highest on cliffs. Ethnographic observations suggest that human settlement practices and fruit use are the main human factors contributing to baobab–settlement positive spatial association. Main conclusions There are three main conclusions: (1) Human settlement and baobab recruitment are spatially dependant because settlement leads directly and indirectly to the development of baobab groves at settlement sites. (2) The lower than expected abundance of mature individuals in natural habitats, and the habitat preferences of the observed population, suggest that baobabs were introduced to south‐western Mali, probably centuries ago. (3) Human mobility over decadal time‐scales is necessary to maintain baobab population structure in landscapes dominated by shifting land use, where baobabs are not purposefully planted. Baobab population processes in such landscapes occur at the scale of human settlement.