Trophy hunting and perceived risk in closed ecosystems: Flight behaviour of three gregarious African ungulates in a semi‐arid tropical savanna

Although being an important conservation tool in Africa, trophy hunting is known to influence risk perception in wildlife species, thus affecting the behaviour and fitness of most targeted species. We studied the effects of trophy hunting on the flight behaviour of impala (Aepyceros melampus), greater kudu (Tragelaphus strepsiceros) and sable (Hippotragus niger) in two closed ecosystems, Cawston Ranch (hunting area) and Stanley and Livingstone Private Game Reserve (tourist area), western Zimbabwe. Using standardized field procedures, we assessed the flight behavioural responses of the three species in two seasons: non‐hunting (December–March) and hunting (April–November) between March 2013 and November 2014. We tested the effect of habitat, group size, sex, season, start distance and alert distance on flight initiation distance using linear mixed models. Habitat, group size sex and alert distance did not have any effect on flight initiation distance for the three species. The three species were more alert and displayed longer flight initiation distances in the hunting area compared with the tourist area. Flight initiation distances for the three species were higher during the hunting season for the hunting area and low during the non‐hunting season. Flight distances of the three species did not differ between the hunting area and the tourist area. We concluded that trophy hunting increased perceived risk of wild ungulates in closed hunting areas, whereas ungulates in non‐hunting areas are less responsive and somehow habituated to human presence. Management plans should include minimum approach distances by tourists as well as establishing seasonal restrictions on special zones to promote species viability. Research aimed at integrating behavioural responses with physiological aspects of target species should be promoted to ensure that managers are able to deal with the behavioural trade‐offs of trophy hunting at local and regional scale.     

Roadless wilderness area determines forest elephant movements in the Congo Basin

A dramatic expansion of road building is underway in the Congo Basin fuelled by private enterprise, international aid, and government aspirations. Among the great wilderness areas on earth, the Congo Basin is outstanding for its high biodiversity, particularly mobile megafauna including forest elephants (Loxodonta africana cyclotis). The abundance of many mammal species in the Basin increases with distance from roads due to hunting pressure, but the impacts of road proliferation on the movements of individuals are unknown. We investigated the ranging behaviour of forest elephants in relation to roads and roadless wilderness by fitting GPS telemetry collars onto a sample of 28 forest elephants living in six priority conservation areas. We show that the size of roadless wilderness is a strong determinant of home range size in this species. Though our study sites included the largest wilderness areas in central African forests, none of 4 home range metrics we calculated, including core area, tended toward an asymptote with increasing wilderness size, suggesting that uninhibited ranging in forest elephants no longer exists. Furthermore we show that roads outside protected areas which are not protected from hunting are a formidable barrier to movement while roads inside protected areas are not. Only 1 elephant from our sample crossed an unprotected road. During crossings her mean speed increased 14-fold compared to normal movements. Forest elephants are increasingly confined and constrained by roads across the Congo Basin which is reducing effective habitat availability and isolating populations, significantly threatening long term conservation efforts. If the current road development trajectory continues, forest wildernesses and the forest elephants they contain will collapse.     

Hunting and Plant Community Dynamics in Tropical Forests: A Synthesis and Future Directions

This synthesis builds on the preceding articles of this Special Section and has three goals. We first review the nascent literature that addresses indirect effects of hunting for tropical forest plant communities. Next, we highlight the potential indirect effects of hunting for other groups of organisms. Our final goal is to consider what could be done to ameliorate the demographic threats to harvest-sensitive game species caused by unsustainable hunting. Three conclusions are possible at this time concerning the impact of hunting for tropical forest plant communities: (1) Hunting tends to reduce seed movement for animal-dispersed species with very large diaspores; (2) Hunting reduces seed predation by granivorous vertebrates for species with large seeds; and (3) Hunting alters the species composition of the seedling and sapling layers. The cascading effects of hunting are already known to affect bruchid beetles and dung beetles and are likely to affect other, nongame taxa. To ameliorate these problems, several lines of research should be further explored to facilitate the development of game management plans including: (1) alternative use of sources of animal protein; (2) income supplementation for local people from sources other than wild meat; (3) outreach and extension activities for communities; (4) recognition and facilitation of the shifting of attitudes towards hunting; (5) implementation of community-based wildlife management programs in regulated-use areas such as extractive reserves; and (6) landscape-scale conservation planning that maximizes the source-sink dynamics of harvested and unharvested game populations and enforces game regulations in strictly protected areas.     

Study on the foraging behaviour of the European nightjar Caprimulgus europaeus reveals the need for a change in conservation strategy in Belgium

Effective nature conservation requires coherent actions based on the best available evidence concerning protected species. Recent studies have suggested that European nightjars Caprimulgus europaeus forage outside their recognized breeding habitats, yet, for Flanders (northern Belgium) information on nightjar foraging behaviour and key foraging habitats is lacking. To assess whether the foraging ecology of nightjars in Flanders is similar to that observed in other parts of Europe, we studied the crepuscular behaviour of this species in Bosland (northeastern Flanders) during a five‐year radio telemetry study. Tracking of 48 individuals within a coniferous forest was standardized and home ranges were calculated using a kernel density estimator (fixed kernel). Habitat use was investigated by comparing kernel placement to available habitat. Average maximal foraging distance was 2603 ± 1094 m and home ranges extended up to 691 ha. We identified the key foraging habitats to be extensively‐cultivated grasslands and recreational areas, areas that were previously assumed unsuitable for Belgian nightjars. Our results indicate the importance of foraging sites outside the breeding territory, confirming the findings of previous studies performed elsewhere in Europe. Incorporating our findings into future conservation plans could, therefore, lead to improved efficiency of EU conservation measures, designed for the protection of this bird species in Flanders.     

Aboriginal overkill

Prior to European influence, predation by Native Americans was the major factor limiting the numbers and distribution of ungulates in the Intermountain West. This hypothesis is based on analyses of (1) the efficiency of Native American predation, including cooperative hunting, use of dogs, food storage, use of nonungulate foods, and hunting methods; (2) optimal-foraging studies; (3) tribal territory boundary zones as prey reservoirs; (4) species ratios, and sex and age of aboriginal ungulate kills; (5) impact of European diseases on aboriginal populations; and (6) synergism between aboriginal and carnivore predation. Native Americans had no effective conservation practices, and the manner in which they harvested ungulates was the exact opposite of any predicted conservation strategy. Native Americans acted in ways that maximized their individual fitness regardless of the impact on the environment. For humans, conservation is seldom an evolutionarily stable strategy. By limiting ungulate numbers and purposefully modifying the vegetation with fire, Native Americans structured entire plant and animal communities. Because ecosystems with native peoples are entirely different than those lacking aboriginal populations, a “hands-off” or “natural regulation” approach by today’s land managers will not duplicate the ecological conditions under which those ecosystems developed. The modern concept of wilderness as areas without human influence is a myth. North America was not a “wilderness” waiting to be discovered, instead it was home to tens of millions of aboriginal peoples before European-introduced diseases decimated their numbers.     

Effects of hunting on the behaviour and spatial distribution of farmland birds: importance of hunting-free refuges in agricultural areas

Hunting is one of the human activities that directly affect wildlife and has received increasing attention given its socioeconomic dimensions. Most studies have been conducted on coastal and wetland areas and showed that hunting activity can greatly affect bird behaviour and distribution. Hunting-free reserves for game species are zones where birds find an area of reduced disturbance. We evaluated the effect of hunting activities on the behaviour and use of hunting-free areas of lapwings Vanellus vanellus , golden plovers Pluvialis apricaria and little bustards Tetrax tetrax in agricultural areas. We compared the habitat use and behaviour of birds on days before, during and after hunting took place. All three studied species showed strong behavioural responses to hunting activities. Hunting activity increased flight probability and time spent vigilant (higher on hunting days than just before and after a hunting day), to the detriment of resting. We also found distributional (use of hunting-free reserve) responses to hunting activities, with hunting-free reserves being used more frequently during hunting days. Thus, reserves can mitigate the disturbance caused by hunting activities, benefiting threatened species in agricultural areas. Increasing the size or number of hunting-free areas might be an important management and conservation tool to reduce the impacts of hunting activities.     

Potential relocation of climatic environments suggests high rates of climate displacement within the North American protection network

Ongoing climate change may undermine the effectiveness of protected area networks in preserving the set of biotic components and ecological processes they harbor, thereby jeopardizing their conservation capacity into the future. Metrics of climate change, particularly rates and spatial patterns of climatic alteration, can help assess potential threats. Here, we perform a continent‐wide climate change vulnerability assessment whereby we compare the baseline climate of the protected area network in North America (Canada, United States, México—NAM) to the projected end‐of‐century climate (2071–2100). We estimated the projected pace at which climatic conditions may redistribute across NAM (i.e., climate velocity), and identified future nearest climate analogs to quantify patterns of climate relocation within, among, and outside protected areas. Also, we interpret climatic relocation patterns in terms of associated land‐cover types. Our analysis suggests that the conservation capacity of the NAM protection network is likely to be severely compromised by a changing climate. The majority of protected areas (~80%) might be exposed to high rates of climate displacement that could promote important shifts in species abundance or distribution. A small fraction of protected areas (<10%) could be critical for future conservation plans, as they will host climates that represent analogs of conditions currently characterizing almost a fifth of the protected areas across NAM. However, the majority of nearest climatic analogs for protected areas are in nonprotected locations. Therefore, unprotected landscapes could pose additional threats, beyond climate forcing itself, as sensitive biota may have to migrate farther than what is prescribed by the climate velocity to reach a protected area destination. To mitigate future threats to the conservation capacity of the NAM protected area network, conservation plans will need to capitalize on opportunities provided by the existing availability of natural land‐cover types outside the current network of NAM protected areas.     

Winter home range and habitat use by porcupines in Alaska

Habitat selection results from trade-offs between availability and use of resources under constraints of predation, competition, or other threats, which can vary spatially and temporally. For northern herbivores, winter food availability and quality can limit population size and may drive habitat preference. North American porcupines (Erethizon dorsatum) are widespread generalist herbivores that range from Mexico to the northern reaches of Alaska. During the long Alaskan winter, porcupines deal with high energetic demands resulting from low ambient temperatures while subsisting on low quality forage. We tracked free-ranging porcupines over 3 winters in southcentral Alaska to determine habitat selection and home range size in relation to diet. Porcupines maintained larger than expected home ranges, and selected for conifer-hardwood forests at the home range level. Individual variation among porcupines was too large to determine a pattern of microhabitat selection among trees. Regardless, direct observations revealed that porcupines used only white spruce and paper birch trees for foraging. White spruce may provide some nutritional and thermoregulatory advantage over paper birch; however, porcupines did feed on paper birch cambium, suggesting some nutritional requirement is met by eating paper birch. Porcupines most likely feed on paper birch cambium when detoxification pathways used to process plant toxins in white spruce needles are saturated. Maintaining mixed conifer-hardwood forests in southcentral Alaska would provide suitable winter habitat for porcupines and may alleviate damage to single species stands of conifers or hardwoods that are preferred by commercial forestry operations. © 2012 The Wildlife Society.     

History of canids in Chile and impacts on prey adaptations

Artiodactyl prey species of Chile, especially guanacos (Lama guanicoe), are reported to be very susceptible to predation by pack‐hunting feral dogs. It has been previously suggested that guanacos and endemic South American deer may have evolved in the absence of pack‐hunting cursorial predators. However, the paleoecology of canid presence in southern South America and Chile is unclear. Here, we review the literature on South American and Chilean canids, their distributions, ecologies, and hunting behavior. We consider both wild and domestic canids, including Canis familiaris breeds. We establish two known antipredator defense behaviors of guanacos: predator inspection of ambush predators, for example, Puma concolor, and rushing at and kicking smaller cursorial predators, for example, Lycalopex culpaeus. We propose that since the late Pleistocene extinction of hypercarnivorous group‐hunting canids east of the Andes, there were no native species creating group‐hunting predation pressures on guanacos. Endemic deer of Chile may have never experienced group‐hunting selection pressure from native predators. Even hunting dogs (or other canids) used by indigenous groups in the far north and extreme south of Chile (and presumably the center as well) appear to have been used primarily within ambush hunting strategies. This may account for the susceptibility of guanacos and other prey species to feral dog attacks. We detail seven separate hypotheses that require further investigation in order to assess how best to respond to the threat posed by feral dogs to the conservation of native deer and camelids in Chile and other parts of South America.     

Responses of a top and a meso predator and their prey to moon phases

We compared movement patterns and rhythms of activity of a top predator, the Iberian lynx Lynx pardinus, a mesopredator, the red fox Vulpes vulpes, and their shared principal prey, the rabbit Oryctolagus cuniculus, in relation to moon phases. Because the three species are mostly nocturnal and crepuscular, we hypothesized that the shared prey would reduce its activity at most risky moon phases (i.e. during the brightest nights), but that fox, an intraguild prey of lynx, would avoid lynx activity peaks at the same time. Rabbits generally moved further from their core areas on darkest nights (i.e. new moon), using direct movements which minimize predation risk. Though rabbits responded to the increased predation risk by reducing their activity during the full moon, this response may require several days, and the moon effect we observed on the rabbits had, therefore, a temporal gap. Lynx activity patterns may be at least partially mirroring rabbit activity: around new moons, when rabbits moved furthest and were more active, lynxes reduced their travelling distances and their movements were concentrated in the core areas of their home ranges, which generally correspond to areas of high density of rabbits. Red foxes were more active during the darkest nights, when both the conditions for rabbit hunting were the best and lynxes moved less. On the one hand, foxes increased their activity when rabbits were further from their core areas and moved with more discrete displacements; on the other hand, fox activity in relation to the moon seemed to reduce dangerous encounters with its intraguild predator.     

Insights into the Management of Large Carnivores for Profitable Wildlife-Based Land Uses in African Savannas

Large African predators, especially lions (Panthera leo) and leopards (Panthera pardus), are financially valuable for ecotourism and trophy hunting operations on land also utilized for the production of other wildlife species for the same purpose. Predation of ungulates used for trophy hunting can create conflict with landholders and trade off thus exists between the value of lions and leopards and their impact on ungulate populations. Therefore productionist and conservation trade-offs are complexly graded and difficult to resolve. We investigated this with a risk-benefit analysis on a large private wildlife production area in Zimbabwe. Our model showed that lions result in substantial financial costs through predation on wild ungulates that may not be offset by profits from hunting them, whereas the returns from trophy hunting of leopards are projected to exceed the costs due to leopard predation. In the absence of additional income derived from photo-tourism the number of lions may need to be managed to minimize their impact. Lions drive important ecological processes, but there is a need to balance ecological and financial imperatives on wildlife ranches, community wildlife lands and other categories of multiple use land used for wildlife production. This will ensure the competitiveness of wildlife based land uses relative to alternatives. Our findings may thus be limited to conservancies, community land-use areas and commercial game ranches, which are expansive in Africa, and should not necessarily applied to areas where biodiversity conservation is the primary objective, even if hunting is allowed there.     

Legal subsistence hunting trends in the Ugalla ecosystem of western Tanzania

Sustainable legal subsistence hunting has a place in conservation. Nonetheless, the long-term success of such schemes depends on them being well managed. We assessed the effectiveness of legal subsistence hunting in the Ugalla ecosystem of western Tanzania using data from the local legal hunting scheme. The hunting in the ecosystem is conducted within the partially protected areas around Ugalla Game Reserve. The Wildlife Division of Tanzania supervises hunting activities in the area via local conservation authorities. We analysed hunting success (animals shot per quota per licence) across species in the period from 1997 to 2004. Our results revealed that 10,511 and 5,991 animals were licenced and shot, respectively. There were considerable variations in hunting success across wildlife species. With the exception of common duiker (Sylvicapra grimmia), hunting success trends for most of the species were declining. The documented decline in wildlife off-take should be further investigated to ensure the sustainable management of this area.     

The Impact of Hunting on the Mammalian Fauna of Tropical Asian Forests

People have hunted mammals in tropical Asian forests for at least 40,000 yr. This period has seen one confirmed global extinction (the giant pangolin, Manis palaeojavanica ) and range restrictions for several large mammals, but there is no strong evidence for unsustainable hunting pressure until the last 2000–3000 yr, when elephants, rhinoceroses, and several other species were progressively eliminated from the large parts of their ranges. Regional declines in most species have occurred largely within the last 50 yr. Recent subsistence hunting has typically focused on pigs and deer (hunted with dogs and spears or with snares), monkeys and other arboreal mammals (often caught with blowpipes), and porcupines and other rodents (smoked or dug out of burrows). Over the last 50 yr, the importance of hunting for subsistence has been increasingly outweighed by hunting for the market. The hunted biomass is dominated by the same species as before, sold mostly for local consumption, but numerous additional species are targeted for the colossal regional trade in wild animals and their parts for food, medicines, raw materials, and pets. Many populations of mammalian dispersers of large seeds and understory browsers have been depleted or eliminated, while seed predators have had a more variable fate. Most of this hunting is now illegal, but the law enforcement is generally weak. However, examples of successful enforcement show that hunting impacts can be greatly reduced where there is sufficient political will. Ending the trade in wild animals and their parts should have the highest regional conservation priority.     

The non-impact of hunting on moose <Emphasis Type="Italic">Alces alces</Emphasis> movement,diurnal activity,and activity range

Previous studies on moose Alces alces have suggested that interactions with humans may trigger anti-predator behaviors and generate a demographical cost. Therefore, we hypothesized that disturbances from small and big game hunting may have negative effects on moose movements, diurnal activity, and activity range. Using location data from 64 moose equipped with GPS collars from three populations (Low Alpine, Inland, Coastal) with different temporal human presence and spatial accessibility, we evaluated the impact of hunting on moose activity rhythms. On average, female moose in the low human population density (Low Alpine) area (<0.5/km²) had significantly lower movement rates during moose hunting season, but variation in movement rates among individuals were higher compared with female moose in regions with denser human populations (6–24/km²). We found no evidence that reproductive status influenced female moose sensitivity to disturbance. As expected, females used smaller activity ranges and were less active nocturnally than males. The high within-group variation suggests that current hunting disturbance levels do not alter moose population behavior in general. Our data indicate that alterations in movement were related to rutting activity, not human disturbance induced by hunting. In line with behavioral theory, our study suggests that some individuals were more sensitive to hunting disturbance than the general population. Our work suggests that individual moose may perceive human predation risk to be similar to other predation risks.     

Do intensive drive hunts affect wild boar (Sus scrofa) spatial behaviour in Italy? Some evidences and management implications

Wild boar have been increasing in numbers all over Western Europe in the last 30 years. The species is a major pest for agriculture, but it has a high value as a game species, and in Italy, as in several other countries, it is traditionally hunted in drive hunts by hunting teams with several dogs. This hunting method can have disruptive effects on the demography and spatial behaviour of wild boar, especially family groups. We conducted a 2-year study (2003 and 2004) to determine the effects of drive hunt disturbance on the spatial behaviour of wild boar family groups in the Northern Apennines (central Italy). Twenty wild boar belonging to ten family groups were ear tagged with a radio device. We located resting sites daily and used intensive tracking sessions during drive hunts. Three seasons were determined: pre-hunting, hunting and post-hunting. A general pattern of increased spatial instability during the hunting season was shown. Resting ranges were larger, and resting sites were more interspersed. Distances between consecutive resting sites were greater during the hunting season and, especially, on hunting days. The displacement of family groups caused by drive hunts was generally short lived except for those groups that were repeatedly hunted and so abandoned their pre-hunt (native) range. During drive hunts, wild boar showed a moderate tolerance to hunting disturbance, and only family groups which were directly chased by dogs escaped or altered their behaviour. The response of wild boar to hunting disturbance seemed to be highly related to the degree of hunting pressure combined with individual variability. The impact on wild boar behaviour should be reduced, above all by avoiding repeated hunts in the same areas within a short period and by employing well-trained hounds.     

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.     

Decline of sable antelope in one of its key conservation areas: the greater Hwange ecosystem,Zimbabwe

Land use has major effects on wildlife conservation. We studied variations of sable antelope Hippotragus niger densities between 1990 and 2001 in comparison with various land uses in and around Hwange National Park, Zimbabwe. Trends of other ungulates, including elephant Loxodonta africana, were examined simultaneously, because sable may be sensitive to forage and apparent competition and to high elephant densities. Sable densities declined in the whole region, very likely because of adverse rainfall conditions. Densities were constantly higher in the hunting areas and forestry lands than in the national park. Interestingly, elephant densities showed the opposite, with higher densities in the national park than in the adjacent areas. Whether these results reflect a negative effect of high elephant numbers on sable must still be tested directly. Likewise, while habitat characteristics and lion predation did not appear responsible for the higher sable densities outside the national park, they could not be discounted as an influence on the differing sable densities in different land‐use areas. It is clear, however, that high protection status is not always sufficient to ensure adequate conservation of flagship species. We therefore call for further investigations of ecological interactions within protected areas.     

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.     

Duiker demography and dispersal under hunting in Northern Congo

Duikers are the most heavily hunted species across forested Central and West Africa. Although these species form a vital food resource for people, little is known about their ecology and demography. The information available to date was collected from populations protected from hunting. However, hunting can profoundly alter demography and behaviour, such as dispersal, which in turn determine the sustainability of harvest. To address this research gap, I used radiotelemetry to examine duiker demography under hunting in Congo-Brazzaville. Among blue duikers ( Philantoba monticola , n = 17) I found annual survival rates (0.94), immature dispersal rates (0.25), and average home ranges (5.5 ha) that largely coincided with previous research from unhunted regions. Neighbouring animals all showed some home range overlap, in contrast to previous studies which found blue duikers to be strictly territorial. Although limited by sample size, immature animals' dispersal distances were relatively high (1.5 km, n = 2). Animals' dispersal rates do not appear to be greatly increased by hunting pressure, although source-sink theory depends upon high rates of dispersal to maintain the elevated and localized hunting offtakes observed around settlements. Building a fuller understanding of duiker demography under hunting, including dispersal, will be essential for conservation and management efforts.     

Timing of vole hunting in aerial predators: RAPTOR GROUP RUG/RIJP*

In a short but intense field investigation, surface activity of common voles in a cropped lucerne field was assayed by live-trapping while, simultaneously, hunting activity and yield of three species of raptors were recorded by continuous observation. Pronounced short-term rhythms in trappability of the vole population ran parallel with fluctuations in yield per hour of flight-hunt of hen harriers and kestrels. These raptors, as well as Rough-legged buzzards, hunted more at times of increased vole surface activity; hen harriers saved c. 15 hours of flight-hunt per day by such temporal adjustment, corresponding to about 12% of their daily energy intake. Voles suffered a predation rate of an estimated 0 2% per day; under such heavy predation and temporal concentration of raptor hunting at times of increased vole activity, surface feeding in synchrony with the vole majority was associated with increased risk of predation.