Behavioral evidence of hunting and foraging techniques by a top predator suggests the importance of scavenging for preadults

Scavenging may be a regular feeding behavior for some facultative raptor species occupying low quality habitats and/or with little experience in hunting techniques. However, its importance has been largely underestimated due to methodological limitations in identifying the real proportion in the diet. Here, through direct observations, we assessed the hunting and foraging success of the threatened Spanish imperial eagle Aquila adalberti determining the influence of age, sex, breeding status, habitat quality, prey type, and landscape characteristics. From 465 observations, Spanish imperial eagles used hunting in flight (42%), scavenging (30%), hunting from a perch (16%) and kleptoparasitism (12%). Our model suggests that Prey size and Prey type best explain hunting success, followed by Landscape and Sex. Our findings suggest that Spanish imperial eagles increase hunting success with age, with scavenging and kleptoparasitism regularly used as juveniles. The absence of relationships with any of the variables considered suggests that kleptoparasitism is an opportunistic behavior used sporadically. Scavenging is also independent of habitat quality and landscape characteristics. Accordingly, low prey density is not a driver of carrion use for preadult individuals, suggesting that a lack of hunting ability obliges this age‐class to use this alternative feeding technique regularly. As a result, the threatened Spanish imperial eagle population is also prone to mortality related to the illegal use of poison baits and, potentially, veterinary drugs (i.e., diclofenac).     

Population density of primates in a large fragment of the Brazilian Atlantic rainforest

We performed a line transect survey (352.4 km) of primates in the Serra de Paranapiacaba, at one of the largest relatively undisturbed fragments of the Atlantic rainforest of Southeastern Brazil (ca. 1400 km²), in August 1998. The brown capuchin, Cebus apella nigritus, was the most common species found in the area (20 groups, density estimate: 5.31 ± 2.05 individuals per km², mean ± SE). Nine groups of the brown howler monkey, Alouatta guariba clamitans, and eight of the woolly spider monkey, Brachyteles arachnoides arachnoides, were also recorded, with preliminary density estimates of 0.79 ± 0.40 and 2.33 ± 1.37 individuals per km², respectively. Density estimates for these species in other fragments of Atlantic rainforest are reviewed, showing that densities in Paranapiacaba are among the lowest reported. It is suggested that the higher densities reported for isolated populations in small forest patches (<50 km²) is related to the absence of main primate predators, the density compensation phenomenon and the ecological plasticity of some primate species. In contrast, local extinction in many small patches is probably related to hunting pressure. Given the important primate populations found in the Paranapiacaba fragment, conservation strategies for the studied species should give priority to effective protection of the largest remnant fragments from illegal hunting and deforestation, rather than translocation of individuals or captive breeding programs to introduce monkeys in small forest fragments vulnerable to hunting and of uncertain future.     

Feeding ecology and its influence on social organization in Brown hyenas (Hyaena brunnea, Thunberg) of the Central Kalahari Desert

(1) Observations are presented on the diet, feeding habits, hunting and foraging behaviour of Brown hyaenas of the Central Kalahari. (2) The remains of kills left by other predators are the single most important food item in the Brown hyaena's diet. The diet also consists of small scavenged items, small prey such as rodents which the hyaena itself kills, and wild fruits. (3) Brown hyaenas hunt and scavenge small items solitarily, but congregate for communal scavenging of the large kills left by other predators. (4) Individual hyaenas are not territorial and there is great overlap in home ranges. They use common pathways and frequently meet to socialize while foraging. (5) Resident adults form a group with a social hierarchy maintained through neck-biting, muzzle-wrestling, chasing, and other social interactions. Subadult hyaenas often leave the group when approximately 22 months old. (6) Brown hyaenas have a complex system of communication including visual displays, social interactions, vocalizations, and extensive pasting. These are described. (7) Since carrion is an important source of food, Brown hyaenas have developed distinct relationships with other predators and these are described. (8) In conclusion, the Brown hyaena exhibits a highly flexible social system, foraging and hunting small items solitarily and congregating for the common utilization of a large carcass. The social organization is therefore influenced by the feeding ecology.     

Are ecological compensation areas attractive hunting sites for common kestrels (Falco tinnunculus) and long-eared owls (Asio otus)?

Common kestrels (Falco tinnunculus) and long-eared owls (Asio otus) in intensively farmed areas in Switzerland decreased markedly as a result of declining vole (Microtus spp.) populations. In order to counteract the loss of biodiversity in intensively farmed areas, the Swiss agri-environment scheme stipulates several types of ecological compensation areas, which together should take up 7% of the farmland. Among them are wild flower and herbaceous strips, which are not mown every year and which in summer support up to 8 times more small mammals than ordinary fields and grassland. This study investigates whether kestrels and long-eared owls preferentially hunt on ecological compensation areas and whether preferred hunting areas are related to the density of small mammals or to the density and height of the vegetation. Both kestrels and long-eared owls mainly hunted on freshly mown low-intensity meadows and artificial grassland, despite low densities of small mammals. Therefore, vegetation structure was more important for the selection of hunting sites than prey abundance. However, both predators preferred to hunt on freshly mown grassland and meadows bordering a wild flower or herbaceous strip. Voles from these strips probably invaded the adjacent freshly mown grassland and became an easy prey for kestrels and owls. In intensively farmed regions, ecological compensation areas, particularly those not mown each year, are an important refuge for small mammals, although in summer the small mammals are not directly accessible to hunting birds. Hence, a mosaic of different habitat types with grassland mown at different times of the year together with undisturbed strips is best suited to provide a year-round supply of accessible food for vole hunters.     

Predator Presence Moves <Emphasis Type="Italic">Helicoverpa armigera</Emphasis> Larvae to Distraction

Displacement of herbivorous insects by the presence of predators on whole plants has rarely been studied. By semi-continuous observations of an externally feeding insect herbivore and a predator, we show how the mere presence of the predator, Geocoris lubra Kirkaldy (Hemiptera: Geocoridae), on a plant can have a strong influence on the movement and behaviors of Helicoverpa armigera (Hübner) (Lepidoptera: Noctuidae) larvae. The presence of predators, as opposed to mortality by predators, influenced the proportion of larvae feeding, resting and implementing avoidance activities. In addition, the proportion of time individual larvae allocated to feeding, resting and dropping off plants was affected when predators were present with and without contact between the two. Predators do more than just reduce numbers of herbivores; they influence feeding, displacement and subsequently the distribution of plant damage.     

Indirect cues in selecting a hunting site in a sit‐and‐wait predator

Sit‐and‐wait predators use relatively simple rules for their decisions to choose and leave a patch, such as using the direct presence of prey to select a hunting site. However, the direct presence of prey can only be used when there is a highly visited patch in the proximity of the predator. Therefore, it is plausible that sit‐and‐wait predators also exploit indirect cues of prey presence and, consequently, use associative learning to select a hunting site. The present study tests for the role of associative learning in a sit‐and‐wait predator species for which the ecology is well understood: Misumena vatia Clerck crab spiders. An ecologically relevant scenario is used by selecting flower colour as the conditioned stimulus and prey presence as the unconditioned stimulus. The results provide no evidence that M. vatia crab spiders use the association between flower colour and food presence for selecting a hunting site. After a training phase of being exposed to a colourful artificial flower highly visited by bees, spiders select a hunting site independently of its colour during the testing phase. Investigations of similar scope and ecological relevance are required with other sit‐and‐wait predators to identify the conditions promoting the use of associative learning for foraging site selection when animals face an unpredictable food supply.     

Which proximate factor determines sexual size dimorphism in tiger snakes?

Diverse interactions between factors that influence body size complicate the identification of the primary determinants of sexual size dimorphism. Using data from a long‐term field study (1997–2009), we examined the contributions of the main proximate factors potentially influencing sexual size dimorphism from birth to adulthood in tiger snakes (Notechis scutatus). Data on body size, body mass and body condition of neonates, juveniles and adults were obtained by mark–recapture. Frequent recaptures allowed us to monitor reproductive status, diet and food intake, and to estimate survival and growth rates in age and sex classes. Additional data from females held briefly in captivity enabled us to assess reproductive output and the body mass lost at parturition (proxies for reproductive effort). From birth to maturity, individuals of both sexes experienced similar growth and mortality rates. We found no difference in diet, feeding and survival rates between the sexes, nor between juveniles and adults. On maturity, despite comparable diet and food intake by both sexes, the high energy requirements of vitellogenesis and gestation were responsible for a depletion of body reserves and probably resulted in a marked decrease in growth rates. Males were largely exempt from such costs of reproduction, and so could grow faster than females and attain larger body sizes. The absence of niche divergence between the sexes (uniformity of habitat, lack of predators) suggests that the impact of differential energetic investment for reproduction on growth rate is probably the main proximate factor influencing sexual size dimorphism in this species. © 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, 103 , 668–680.     

Plant feeding of yellow baboons (Papio cynocephalus) in Mikumi national park,Tanzania, and the relationship between seasonal feeding and immature survival

Focal-animal feeding data obtained from 64 adult baboons during a 3-year period were used together with equivalent data from 46 infants to evaluate hypotheses predicting selection for a birth peak and to study the baboon’s eclectic/selective feeding adaptation, with emphasis on differential feeding by sex, developmental trends, and seasonal use of food classes (fruit, leaf, flower, grass, etc.). The findings suggest that feeding conditions are better in the wet season than in the dry season. Despite large sexual dimorphism, estimates of total amounts eaten were virtually identical for males and females. Infants used all of the same plant-food classes as adults, but proportional differences occurred for some food classes in amounts eaten. Foods eaten proportionately less by infants were probably harder for them to obtain and process or were chosen through inexperience or for exploration. There was considerable between-year variation in amounts of food classes eaten, but the within-year standard deviations were similar, as were also the mean amounts eaten per year. An eclectic/selective feeding adaptation has the advantage of permitting long-run acquisition of adequate nutrition within a context of high feeding variation from season to season and year to year. Mixed results were obtained from hypotheses about selection for a birth peak. Although a peak occurred in the early dry season, this was not the optimal time of birth for survival. Survival was highest for individuals born in the late wet season, when the availability and probably the quality of food for lactating mothers were greatest.     

Facilitated predation through interaction between life stages in the stinkbug predatorperillus bioculatus (Hemiptera: Pentatomidae)

Competitive interactions in arthropod predators are well-known, but positive interactions have received less attention. The two-spotted stinkbugPerillus bioculatus often feeds gregariously on leaf beetle larvae and caterpillar prey. Consequences of prey sharing amongP. bioculatus conspecifics of dissimilar size (instar) was studied using Colorado potato beetle (CPB) prey. Rearing second-instar (N2) nymphs ofP. bioculatus with an N5 conspecific facilitated early feeding on L4 CPB larvae (a difficult prey to handle by N2 nymphs but not by N5’s), thus increasing survival and accelerating development. One in every 20 cases ofP. bioculatus foraging in the field was accounted for by pairs or small groups of mostly feeding individuals. CPB egg masses and L4’s represented a disproportionate number of cases of aggregated feeding byP. bioculatus, compared to feeding singly. Small CPB larvae decreased in the diet of aggregated stinkbugs compared to L4 larvae and egg masses, suggesting that sharing these prey may be favorable or unavoidable. In a field test measuring residence/survival of N2’s limited to L4 prey, the N2’s rate of residence/survival increased significantly when large nymphs acting as food providers were also present. The function of communal feeding inP. bioculatus is discussed, as well as the potential for greater impact on prey density that may be expected from tolerance to opportunistic feeding by conspecifics in slightly gregarious predators.     

Rabbit (Oryctolagus cuniculus) abundance and protected areas in central-southern Spain: why they do not match?

European rabbits (Oryctolagus cuniculus), a keystone species in the Iberian Mediterranean ecosystem, are the staple prey of the Iberian lynx (Lynx pardinus) and the Spanish imperial eagle (Aquila adalberti). These predators require medium to high rabbit densities and a low degree of human disturbance. We compared rabbit abundances in areas of central-southern Spain under three levels of protection and management: protected areas, intensively managed (nonprotected) hunting estates, and other nonprotected areas. We used pellet abundance indices to estimate rabbit density in 118 surveys conducted during the summers of 2002 and 2003. We observed greater rabbit abundance in intensively managed hunting estates compared to protected areas and other nonprotected areas, perhaps because policy makers did not consider rabbit numbers when selecting priority areas. Alternatively, differences in game management practices (e.g., predator control or habitat management) may explain the higher rabbit densities observed in managed hunting estates. Our results suggest that the best feeding conditions for the Iberian lynx and the Spanish imperial eagle occur in intensively managed hunting areas, where such predators are frequently persecuted. The conservation of these endangered predators may require efforts to increase rabbit densities in protected areas.     

Food and feeding habits of Clarias mossambicus from four areas in the Lake Victoria basin,East Africa

Synopsis The food and feeding habits of Clarias mossambicus in four habitat types in the northern Lake Victoria basin were studied. The importance of certain food types to this species varies from area to area. In the lake, rivers and swamps, C. mossambicus fed mainly on fish, particularly Haplochromis species. In the sewage ponds, Crustacea and insects were the main food items. The extent of ingestion by C. mossambicus of a given food type presumably reflects the availability and abundance of the food in the habitat. This constitutes a shift in the food habits which is of vital importance in this migratory fish. By employing several feeding methods, i.e. hunting, bottom feeding, surface feeding and filter feeding, C. mossambicus exploits a broad spectrum of food resources in the various habitats.Many modern workers consider Clarias mossambicus conspecific with C. gariepinus and a junior synonym of the latter (Editor)     

Use of grit supplements by waterbirds: an experimental assessment of strategies to reduce lead poisoning

The ingestion of spent Pb shot due to confusion with grit or inadvertently with food particles causes Pb poisoning in a large number of waterbirds, this being one of the main causes of mortality for some species. Lead ammunition for hunting is being progressively banned in more countries, while grit supplementation has been proposed as a management measure to reduce the ingestion of deposited Pb shot. Studies of grit selection with waterfowl in semi-captivity and in the wild were undertaken to evaluate preferences in the colour and geochemical composition of grit, whether it was available dry or in water, its position within the wetland and the relationship between grit ingestion and feeding behaviour. Grit ingestion was higher when food was included in the treatments. In the absence of food, red grit was taken in higher amounts than grey in semi-captivity but not in the wild. Siliceous grit was taken in a higher amount than calcareous when offered dry, but not in water. No differences in the amount of ingested grit were found among different positions within the wetland. The number of feeding attempts in plots supplemented with grit was higher than in those without grit, although the highest numbers of feeding birds were found in plots supplemented with food. Grit ingestion in waterfowl is intimately associated with feeding behaviour. To optimize the effectiveness of grit supplementation to reduce the risk of Pb poisoning in waterfowl, calcareous and siliceous grit may be combined and applied in feeding sites or mixed with bait to attract birds.     

Does illegal hunting affect density and behaviour of African grassland birds? A case study on ostrich (Struthio camelus)

Widespread bushmeat hunting represents one of the major threats to many mammals and birds in Africa. We studied the influence of illegal bushmeat hunting on large grassland birds in the Serengeti National Park (SNP) and adjoining protected areas, by using the ostrich (Struthio camelus) as a case study. First, we documented illegal hunting of both small and large birds by using a questionnaire in the villages on the western and eastern side of the SNP. Second, we studied the effect of illegal hunting on density by driving 4,659 km of transects inside SNP and on the adjacent protected areas, where the data were analysed by DISTANCE sampling. Last, we used flight initiation distance (FID, i.e. the distance between an approaching predator (human) and prey when flight is started), to assess possible impacts on behaviour from illegal hunting. We found that people from the western side of the SNP admitted to hunting both small and large grassland birds, and collect ostrich feathers and eggs. Although the Maasai also hunted small birds, only ostrich feathers and eggs of the large grassland birds were used. Surprisingly, we found no significant differences in densities between the SNP and adjoining partially protected areas, but ostriches had longer FID to an approaching human outside the SNP. Currently illegal hunting does not appear to affect the ostrich population, but given the extensive use of birds for consumption more awareness educational programs accompanied by provision of agricultural incentives within the protected areas are needed.     

Behavioural and life history effects of predator diet cues during ontogeny in damselfly larvae

A central issue in predator–prey interactions is how predator associated chemical cues affect the behaviour and life history of prey. In this study, we investigated how growth and behaviour during ontogeny of a damselfly larva (Coenagrion hastulatum) in high and low food environments was affected by the diet of a predator (Aeshna juncea). We reared larvae in three different predator treatments; no predator, predator feeding on conspecifics and predator feeding on heterospecifics. We found that, independent of food availability, larvae displayed the strongest anti-predator behaviours where predators consumed prey conspecifics. Interestingly, the effect of predator diet on prey activity was only present early in ontogeny, whereas late in ontogeny no difference in prey activity between treatments could be found. In contrast, the significant effect of predator diet on prey spatial distribution was unaffected by time. Larval size was affected by both food availability and predator diet. Larvae reared in the high food treatment grew larger than larvae in the low food treatment. Mean larval size was smallest in the treatment where predators consumed prey conspecifics, intermediate where predators consumed heterospecifics and largest in the treatment without predators. The difference in mean larval size between treatments is probably an effect of reduced larval feeding, due to behavioural responses to chemical cues associated with predator diet. Our study suggests that anti-predator responses can be specific for certain stages in ontogeny. This finding shows the importance of considering where in its ontogeny a study organism is before results are interpreted and generalisations are made. Furthermore, this finding accentuates the importance of long-term studies and may have implications for how results generated by short-term studies can be used.     

Multiple intraguild predators reduce mortality risk of a mutual agricultural pest prey in simple,but not in complex,experimental settings

Multiple predator species that coexist with each other and their mutual prey can have combined effects on prey mortality that are similar to the sum of each predator's individual impact (linear effects), greater than the sum of each predator's individual impact (risk enhancement), or less than the sum of each predator's individual impact (risk reduction). Understanding multiple predator effects is important to determine the impact of predators on pest prey in agroecosystems. If two predators share the same broad spatial domain and hunting mode and engage in intraguild predation, then their combination is expected to result in risk reduction for a mutual prey. We tested this hypothesis using both additive and replacement experimental designs on two species of generalist wolf spider predators (Tasmanicosa leuckartii and Hogna crispipes) that hunt in the same domain, and a mutual insect prey (cotton bollworm Helicoverpa armigera). We used two types of enclosures: a small simple laboratory enclosure, and a larger more complex cotton plant enclosure. We found that in the small simple laboratory enclosures, the presence of two spiders led to risk reduction of Helicoverpa larva mortality as expected, but in larger more complex cotton plant enclosures the presence of both species resulted in linear effects rather than risk reduction on Helicoverpa mortality. Furthermore, intraguild predation did not change multiple predator effects in laboratory or plant enclosures. This study has implications for managing arthropod predators in agroecosystems; contrary to predictions of ecological frameworks, coexistence of predators that share the same hunting mode and hunting domain may not lead to risk reduction on a mutual prey in more complex environments, where encounters among predators can be lower. Conservation of multiple predators of a single guild can play an essential role on biological control of insect pests.     

Age bias in the bag of pink-footed geese <Emphasis Type="Italic">Anser brachyrhynchus</Emphasis>: influence of flocking behaviour on vulnerability

In pink-footed goose (Anser brachyrhynchus) wintering in Denmark, The Netherlands and Belgium, the proportion of juveniles in the hunting bag is consistently higher than that observed in the autumn population. Such juvenile bias in the bag is usually ascribed to young geese lacking experience with hunting or disruption of juveniles from families. An alternative explanation may be that flocking behaviour of families make juveniles more vulnerable. Observations of morning flights of pink-footed geese to the feeding grounds from two of the major autumn-staging areas showed that geese were distributed in many small flocks (median flock size = 9). This was not significantly different from the flock size distribution shot at by hunters (median = 8), suggesting that hunters targeted goose flock size in proportion to the general probability of encounter. The rate at which hunters downed geese was independent of flock size. The ratio between juveniles and adults in flocks decreased with flock size and flocks of <60 individuals primarily comprised family groups. The likelihood of being shot at was 2.4 times higher for juveniles and 3.4 times higher for older birds in small flocks (<10 individuals) compared to larger flocks. The observations suggest that both juveniles as well as successful adult breeding birds were more vulnerable than non-breeding/failed breeding birds as a result of flocking behaviour.     

Food partitioning between breeding White-tailed Kites (Elanus leucurus; Aves; Accipitridae) and Barn Owls (Tyto alba; Aves; Tytonidae) in southern Brazil.

I examined the diet of breeding White-tailed Kites (Elanus leucurus; Aves; Accipitridae) and Barn Owls (Tyto alba; Aves; Tytonidae) in an agrarian area of southern Brazil by analyzing regurgitated prey remains. The objective was to evaluate how these raptors, which differ markedly in their hunting activity periods (owls are nocturnal and kites diurnal), share their mammalian food component. 2,087 prey consumed by Barn Owls and 1,276 by White-tailed Kites were identified. They presented a high overlap of food-niches (Piankas index was 0.98). Based on the daily activity period of their main small mammal prey, a lower overlap would be expected. The crepuscular/nocturnal Mus musculus was the main prey for the diet of breeding Barn Owls (81%) and White-tailed Kites (63%). This small exotic rodent provided 63% of the small mammal biomass ingested by owls and 44% by kites. Larger native small mammals were also considered important for the diet of kites, mainly because of their biomass contribution. Although these raptors differ markedly in their hunting activity periods, Barn Owls and White-tailed Kites are very similar predators in southern Brazil, overlapping their diets.     

Relationship Between Small-game Hunting and Carnivore Diversity in Central Spain

Small-game hunting plays an important economic role in central Spain and some game species are key food resources for threatened predators. Small carnivores may reduce numbers of important game species. To alleviate predation pressure on game populations, management is often focused on predator control. Control methods can be non-selective, and so could potentially have a negative impact on non-target carnivore species, affecting carnivore guild composition and diversity. Using data on carnivore diversity in 70 quadrants (5 × 5 km) in a large area of central Spain with two main land uses (small-game hunting and a mixture of big-game hunting and forestry), we assessed how game management was related to carnivore diversity. Carnivore species richness was significantly lower in those areas managed for small-game hunting than in areas where other land uses predominated. The apparently least affected species by predator control was the red fox (Vulpes vulpes), the main target species of predator control. Results suggested that low carnivore species richness was linked to small-game hunting activities, which was probably associated with non-selective predator control practices. Therefore, in order to reconcile carnivore conservation and economic goals it is imperative to develop and to evaluate selective predator control techniques.     

Annual and seasonal changes in diets of martens: evidence from stable isotope analysis

Theory predicts that generalist predators will switch to alternative prey when preferred foods are not readily available. Studies on the feeding ecology of the American marten (Martes americana) throughout North America suggest that this mustelid is a generalist predator feeding largely on voles (Microtus sp.; Clethrionomys sp.). We investigated seasonal and annual changes in diets of martens in response to the changing abundance of small rodents (Peromyscus keeni, and Microtus longicaudus) on Chichagof Island, Southeast Alaska, using stable isotope analysis. We hypothesized that martens would feed primarily on small rodents during years with high abundance of these prey species, whereas during years of low abundance of prey, martens would switch to feed primarily on the seasonally available carcasses of salmon. We also hypothesized that home-range location on the landscape (i.e., access to salmon streams) would determine the type of food consumed by martens, and martens feeding on preferred prey would exhibit better body condition than those feeding on other foods. We live-captured 75 martens repeatedly, from mid-February to mid-December 1992–1994. We also obtained marten carcasses from trappers during late autumn 1991 and 1992, from which we randomly sub-sampled 165 individuals. Using stable isotope ratios and a multiple-source mixing model, we inferred that salmon carcasses composed a large portion of the diet of martens in autumn during years of low abundance of rodents (1991 and 1992). When small rodents were available in high numbers (1993 and 1994), they composed the bulk of the diet of martens in autumn, despite salmon carcasses being equally available in all years. Selection for small rodents occurred only in seasons in which abundance of small rodents was low. Logistic regression revealed that individuals with access to salmon streams were more likely to incorporate salmon carcasses in their diet during years of low abundance of small rodents. Using stable isotope analysis on repeated samples from the same individuals, we explored some of the factors underlying feeding habits of individuals under variable ecological conditions. We were unable to demonstrate that body weights of live-captured male and female martens differed significantly between individuals feeding on marine-derived or terrestrial diets. Therefore, martens, as true generalist predators, switched to alternative prey when their principal food was not readily available on a seasonal or annual basis. Although salmon carcasses were not a preferred food for martens, they provided a suitable alternative to maintain body condition during years when small rodents were not readily available. Received: 1 May 1996 / Accepted: 24 February 1997