Predator Detection is Limited in Microhabitats with High Light Intensity: An Experiment with Brown‐Headed Cowbirds

Variations in ambient light conditions across different microhabitats can modify the detectability of predators and prey. Prey have been shown to be more visible in sunlit than in shaded patches, leading to higher predation risk and more investment in vigilance (predation risk hypothesis). Additionally, prey have been hypothesized to take longer to detect predators in sunlit compared to shaded patches because of the excess of sunlight causing glare effects (disability glare hypothesis). We tested the predictions of these two non‐mutually exclusive hypotheses in a seminatural experiment with brown‐headed cowbirds by measuring vigilance behavior and detection of a ground predator in patches under the shade of vegetation and in the open. Light intensity and achromatic contrast were higher in the sunlit patches, which could enhance glare effects, but chromatic contrast was higher in the shaded patches. Brown‐headed cowbirds took longer to show alert reactions to and flee from a ground predator in sunlit compared to shaded patches. However, the two parameters associated with perceived predation risk (vigilance prior to the predator exposure and time to resume foraging after the attack) did not differ between sunlit and shaded patches. Our findings support to a greater extent the disability glare hypothesis than the predation risk hypothesis. Overall, ambient light conditions can affect two critical components of behavioral predator–prey interactions in terrestrial habitats: detection of and escape from predators. The effects of disability glare are expected to be more pronounced in bird species with wider visual fields or without sun‐shading structures; however, species may compensate through various behaviors (e.g. avoidance of sunlit patches and changes in head orientation).     

Effects of multiple vertebrate predators on grasshopper habitat selection: trade-offs due to predation risk, foraging, and thermoregulation

Predation risk can influence habitat use and activity of potential prey. I explored how the risk of predation by vertebrates influenced the behavior of grasshoppers. I monitored the height in vegetation and the frequency of resting, moving, and feeding behaviors of both tethered and free-ranging grasshoppers under exposure to various predators. Grasshoppers protected from birds remained high in the vegetation, while those protected only from small mammals and lizards remained low in the vegetation. Grasshoppers exposed to all predators occupied an intermediate height. Lower positions in the vegetation were associated with cooler thermal conditions, lower feeding rates, and lower food availability. My results are consistent with the hypothesis that grasshoppers utilize different microhabitats to balance the trade-off between reducing mortality from predators and experiencing greater food availability, and warmer conditions. This revised version was published online in July 2006 with corrections to the Cover Date.     

Loss of body mass under predation risk: cost of antipredatory behaviour or adaptive fit-for-escape?

Predation risk may compromise the ability of animals to acquire and maintain body reserves by hindering foraging efficiency and increasing physiological stress. Locomotor performance may depend on body mass, so losing mass under predation risk could be an adaptive response of prey to improve escape ability. We studied individual variation in antipredatory behaviour, feeding rate, body mass and escape performance in the lacertid lizard Psammodromus algirus. Individuals were experimentally exposed to different levels of food availability (limited or abundant) and predation risk, represented by reduced refuge availability and simulated predator attacks. Predation risk induced lizards to reduce conspicuousness behaviourally and to avoid feeding in the presence of predators. If food was abundant, alarmed lizards reduced feeding rate, losing mass. Lizards supplied with limited food fed at near-maximum rates independently of predation risk but lost more mass when alarmed; thus, mass losses experienced under predation risk were higher than those expected from feeding interruption alone. Although body mass of lizards varied between treatments, no component of escape performance measured during predator attacks (endurance, speed, escape strategy) was affected by treatments or by variations in body mass. Thus, the body mass changes were consistent with a trade-off between gaining resources and avoiding predators, mediated by hampered foraging efficiency and physiological stress. However, improved escape efficiency is not required to explain mass reduction upon predator encounters beyond that expected from feeding interruption or predation-related stress. Therefore, the idea that animals may regulate body reserves in relation to performance demands should be reconsidered.     

Escape tactics and effects of perch height and habituation on flight initiation distance in two Jamaican anoles (Squamata: polychrotidae)

Escape by Anolis lizards is influenced by microhabitats and fight initiation distance increases with predation risk. Differences in microhabitat use among ecomorphs affect escape behavior, but only two studies have reported ecomorphological differences in flight initiation distance among Greater Antillean species. I studied effects of predation risk and microhabitats on escape behavior by conducting field experiments using two species of anoles, Anolis lineatopus and A. grahami, on the campus of the University of the West Indies at Mona, Jamaica. Because ecomorphological variation of anoles has evolved independently within each island of the Greater Antilles, but relationships between ecomorphs and escape behaviors are poorly known, I characterized microhabitat use and escape tactics, and determined relationships between flight initiation distance and two risk factors, habituation to human presence and perch height, in Anolis lineatopus, a trunk-ground anole and A. grahami, a trunk-crown anole. Sample sizes for A. lineatopus and A. grahami were 214 and 93, for microhabitat use and escape destinations, 74 and 34 for human presence and 125 and 34 for perch height. The two species occurred in similar microhabitats and exhibited similar escape tactics, but exhibited key differences expected for their ecomorphs. Both species were sighted frequently on the ground and on trees, but A. lineatopus were more frequently on ground and were perched lower than A. grahami. Both species escaped from ground to trees and when on trees hid on far sides and escaped without changing climbing direction with equal frequency. The frequency of fleeing upward was greater for A. grahami than A. lineatopus. Both species exhibited habituation by having shorter flight initiation distances in areas with more frequent exposure to people. In both species flight initiation distance increased as perch height decreased because, lizards had to climb farther to be out of reach when perched lower. The relationship between flight initiation distance and perch height may apply to other anole ecomorphs that flee upward when low perched on trees.     

Effects of tail loss on the movement patterns of the lizard, Psammodromus algirus

1. Many lizards use caudal autotomy as a defensive strategy. However, subsequent costs related to the alteration of locomotor abilities might decrease the fitness of individuals. In this paper, the movement patterns of spontaneously moving Psammodromus algirus lizards and their escape performance running at high speed were compared before and after tail loss. A control tailed group was also studied to assess the repeatability of locomotor patterns between trials.
2. Tail loss had a significant effect on spontaneous movement patterns. Tailless individuals moved at significantly slower speeds during bursts of locomotion, and distances moved within bursts were significantly reduced. The overall time spent pausing increased, and, as a result, overall speeds decreased to an even greater extent than burst speeds. However, mean durations of individual locomotor bursts and mean pause durations did not change significantly after tail loss.
3. Loss of the tail decreased mean stride length, although the positive relation between stride length and speed was retained.
4. Escape performance was also greatly affected; loss of the tail resulted in substantially reduced attained, maximal and overall escape speeds. These changes resulted in shorter escape distances (the time of the first pause after the initiation of the escape response) because the mean duration of escape responses did not change.
5. The relevance of these alterations for the ecology of this species, and how individuals may compensate for the costs of tail loss, favouring autotomy as an escape strategy, are discussed.     

Vigilance and Collective Detection of Predators in Degus (Octodon degus)

Individuals of social and partially social species typically reduce their vigilance activity when foraging in groups. As a result, per capita risk of predation decreases and individuals allocate more time to foraging and other fitness rewarding activities. Reduction of per capita risk is hypothesized to occur because there are more individuals to detect potential predators. If so, collective (i.e. total) vigilance is expected to increase with foraging group size. Increased surveillance during group foraging may occur if group members scan independently of one another, or sequentially to avoid the overlapping of their vigilance bouts. Intriguingly, such coordinated vigilance assumes that individuals monitor not only the presence, but the vigilance behaviour of group mates. We used seasonal records on time budget and grouping patterns of individually marked degus (Octodon degus), a social rodent, to examine if (a) individual vigilance decreases and/or foraging increases with group size, (b) collective vigilance increases with group size and (c) foraging degus coordinate their vigilance. When foraging, degus decreased their individual vigilance and increased their foraging time when in larger groups. Despite this, degus in larger groups increased their collective vigilance, supporting the hypothesis that socially foraging degus decrease predation risk through an improved ability to detect and escape potential predators. Additionally, patterns of collective vigilance suggested that degus scan independently of each other and so, they do not coordinate their vigilance to prevent its temporal overlapping. This finding does not support that foraging degus monitor the vigilance activity of group mates.     

More Ornamented Males Exhibit Increased Predation Risk and Antipredatory Escapes,but not Greater Mortality

Secondary sexual traits not only confer benefits to their bearer through increased mate acquisition, but may also have inherent costs, including the attraction of predators. Here, we examined the relationship between conspicuous secondary sexual traits and predation costs using two male morphs of Schizocosa wolf spiders: brush‐legged and non‐ornamented. In the field, we ran two predation experiments using artificial enclosures to directly test mortality costs of predation on the two male morphs. Using a natural predator, a larger wolf spider in the genus Hogna, we found no difference in predation on brush‐legged vs. non‐ornamented males. However, predation was depends on environmental conditions. More individuals were preyed upon at night (vs. during the day) and on rock litter (vs. leaf litter), but the two male morphs were preyed upon equally to each other across environmental treatments. A laboratory experiment incorporated staged interactions between a single predator (Hogna) and each male morph to examine finer details of predation events. Again, we found no differential mortality between brush‐legged and non‐ornamented males. However, brush‐legged males were attacked sooner and were more likely to escape the attack. Our results show an association between sexual ornamentation and predation risk as well as escape behavior.     

Morphological correlates of burst speed and field movement patterns: the behavioural adjustment of locomotion in wall lizards (Podarcis muralis)

Locomotion of lizards has clear morphological determinants and is important for developing activities such as feeding, social interaction and predator avoidance. Thus, morphological variation is believed to have fitness consequences through affecting locomotor performance. This paper firstly evaluates the dependence of burst speed on morphology, and secondly examines the movement patterns of free-ranging undisturbed wall lizards ( Podarcis muralis ) engaged in several kinds of activity. Body size was the most important correlate of burst speed as performed at the optimal temperature for running in the laboratory. After removing size effects from performance and morphological traits, the length of some particular limb segments had positive influence on burst speed, but these effects were weak, each trait explaining less than 16% of variance in burst speed. Free-ranging P. muralis exhibited intermittent locomotion, with movement sequences interrupted by frequent short pauses. Field movement patterns greatly differed depending upon the kind of activity and were in most aspects independent of the size and sex of the animal. P. muralis involved in thermoregulation performed short and low-speed displacements; exploratory activities were characterized by frequent, slow and short movements. On the contrary, lizards involved in intraspecific pursuits and predator escape developed comparatively high speeds, although only exceptionally did they attain the size-specific burst speed predicted from the laboratory trials. Speed of escape increased with distance to the refuge and the animals are able to assess predation risks to modulate approach distance, speed and pauses, so maximum exertion is seldom required. The evolution of locomotor capacities exceeding routine needs is discussed in the context of the principle of 'excessive construction'.  © 2003 The Linnean Society of London, Biological Journal of the Linnean Society , 2003, 80 , 135–146.     

Red kangaroos (<Emphasis Type="Italic">Macropus rufus</Emphasis>) receive an antipredator benefit from aggregation

For species that cannot seek cover to escape predators, aggregation becomes an important strategy to reduce predation risk. However, aggregation may not be entirely beneficial because aggregated animals may compete for access to limited resources and might even attract predators. Available evidence suggests that foraging competition influences time allocation in large-bodied macropodid marsupials, but previous studies have focused primarily on species in areas with protective cover. We studied red kangaroos, a species often found in open country without noticeable cover, to determine whether they experienced a net benefit by aggregation. Red kangaroos varied their time allocation as a function of group size and, importantly, more variation in time allocation to vigilance and foraging was explained by non-linear models than by linear models. This suggests red kangaroos directly translated the reduction of predation risk brought about by aggregation into greater time foraging and less time engaged in vigilance. We infer that red kangaroos received a net benefit by aggregation. Social species living in the open may be generally expected to rely on others to help manage predation risk. Communicated by K. Kotrschal     

Locomotion and Escape Modes in Rodents of the Monte Desert (Argentina)

Modes of locomotion and escape tactics are attributes that affect the structure of animal communities, promoting exploitation of different microhabitats and the coexistence of different species. Bipedal locomotion is considered to be more effective than a quadrupedal gait in escaping attacks by predators because it allows for higher speed, a faster response to attack, sudden changes of direction and better detection of aerial raptors. The aim of this study was to determine the type of locomotion used at the moment of escape by three rodent species of the Monte desert –Eligmodontia typus, Akodon molinae and Graomys griseoflavus. The study was carried out in three plant communities of the Ñacuñán Reserve (Mendoza). All three species showed differences in both mode of escape and locomotory pattern. Graomys griseoflavus exhibited the highest proportion of escapes using quadrupedal saltation. The mode of locomotion employed by E. typus varied according to the type of plant communities it inhabited. Those occurring at open sites (Medanal community) exhibited a greater propensity to jump during escapes than those from more sheltered habitats (Algarrobal community). Akodon molinae relied primarily on a quadrupedal gait when fleeing from predators, which would explain its greater dependence on plant cover. Therefore, the morphological and behavioural characteristics of these species are related to their mode of locomotion and the strategies they employ to diminish the risk of predation.     

Locomotion patterns in two South American gymnophthalmid lizards: Vanzosaura rubricauda and Procellosaurinus tetradactylus

We quantified gait and stride characteristics (velocity, frequency, stride length, stance and swing duration, and duty factor) in the bursts of locomotion of two small, intermittently moving, closely related South American gymnophthalmid lizards: Vanzosaura rubricauda and Procellosaurinus tetradactylus. They occur in different environments: V. rubricauda is widely distributed in open areas with various habitats and substrates, while P. tetradactylus is endemic to dunes in the semi-arid Brazilian Caatinga. Both use trot or walking trot characterised by a lateral sequence. For various substrates in a gradient of roughness (perspex, cardboard, sand, gravel), both species have low relative velocities in comparison with those reported for larger continuously moving lizards. To generate velocity, these animals increase stride frequency but decrease relative stride length. For these parameters, P. tetradactylus showed lower values than V. rubricauda. In their relative range of velocities, no significant differences in stride length and frequency were recorded for gravel. However, the slopes of a correlation between velocity and its components were lower in P. tetradactylus on cardboard, whereas on sand this was only observed for velocity and stride length. The data showed that the difference in rhythmic parameters between both species increased with the smoothness of the substrates. Moreover, P. tetradactylus shows a highly specialised locomotor strategy involving lower stride length and frequency for generating lower velocities than in V. rubricauda. This suggests the evolution of a central motor pattern generator to control slower limb movements and to produce fewer and longer pauses in intermittent locomotion.     

Detecting predators and locating competitors while foraging: an experimental study of a medium-sized herbivore in an African savanna

Vigilance allows individuals to escape from predators, but it also reduces time for other activities which determine fitness, in particular resource acquisition. The principles determining how prey trade time between the detection of predators and food acquisition are not fully understood, particularly in herbivores because of many potential confounding factors (such as group size), and the ability of these animals to be vigilant while handling food. We designed a fertilization experiment to manipulate the quality of resources, and compared awareness (distinguishing apprehensive foraging and vigilance) of wild impalas (Aepyceros melampus) foraging on patches of different grass height and quality in a wilderness area with a full community of predators. While handling food, these animals can allocate time to other functions. The impalas were aware of their environment less often when on good food patches and when the grass was short. The animals spent more time in apprehensive foraging when grass was tall, and no other variable affected apprehensive behavior. The probability of exhibiting a vigilance posture decreased with group size. The interaction between grass height and patch enrichment also affected the time spent in vigilance, suggesting that resource quality was the main driver when visibility is good, and the risk of predation the main driver when the risk is high. We discuss various possible mechanisms underlying the perception of predation risk: foraging strategy, opportunities for scrounging, and inter-individual interference. Overall, this experiment shows that improving patch quality modifies the trade-off between vigilance and foraging in favor of feeding, but vigilance remains ultimately driven by the visibility of predators by foragers within their feeding patches.     

Associations between Non‐Lethal Injury,Body Size,and Foraging Ecology in an Amphibian Intraguild Predator

Theoretical treatments of intraguild predation and its effects on behavioral interactions regard the phenomenon as a size‐structured binary response wherein predation among competitors is completely successful or completely unsuccessful. However, intermediate outcomes occur when individuals escape intraguild (IG) interactions with non‐lethal injuries. While the effects of wounds for prey include compromised mobility and increased predation risk, the consequences of similar injuries among top predators are not well understood, despite the implications for species interactions. Using an amphibian IG predator, Ambystoma opacum (Caudata: Ambystomatidae), we examined associations between non‐lethal injuries and predator body size, foraging strategy, microhabitat selection, and intraspecific agonistic interactions. Wounds were common among IG predators, generally increasing in frequency throughout larval ontogeny. Non‐lethal injuries were associated with differences in predator body size and behavior, with injured predators exhibiting smaller body sizes, increased use of benthic microhabitats, reduced agonistic displays, and increased risk of intraspecific aggression. While such effects were not ultimately associated with reduced foraging success, non‐lethal injury could contribute to niche partitioning between injured and healthy predators via habitat selection, but injured predators likely continue to exert predatory pressure on IG and basal prey populations. Our results indicate that studies of top‐down population regulation should incorporate injury‐related modifications to both prey and predator behavior and size structure.     

Development of anuran locomotion: Ethological and neurophysiological considerations

There are dramatic quantitative and qualitative differences in the locomotor behavior of larval and juvenile frogs. Larvae (tadpoles) are primarily herbivourous and rely heavily on locomotion via undulations to acquire food and avoid predation. After metamorphosis, juvenile frogs adopt a carnivorous lifestyle and capture prey and avoid predators by remaining motionless in a place of concealment. When they must move, frogs locomote by means of ballistic hops or by more conventional walking. However, locomotion of both tadpoles and frogs can be considered of two fundamental functional types: (a) startle and escape; and (b) sustained locomotion. Neural mechanisms underlying startle responses and sustained locomotion in larvae and juveniles are described and possible ontogenetic relationships those behaviors are proposed. The role of different parts of the nervous system in the ontogeny of locomotion, as well as nonneuronal factors, are described. Results show that the transition from tadpole-like behavior to frog-like behavior is not a simple function of maturation of central locomotor controls. Rather, it results from a complex interaction of central nervous system maturation, morphological change, and a change in habitat preference. Examples of similar multidimensional control of behavioral ontogeny in other species are described, and it is argued that to understand the ontogeny of behavior, one must investigate contributions made at all levels, from the neuronal to the environmental. © 1992 John Wiley & Sons, Inc.     

Predator-prey shell games: large-scale movement and its implications for decision-making by prey

Many classical models of food patch use under predation risk assume that predators impose patch-specific predation risks independent of prey behavior. These models predict that prey should leave a chosen patch only if and when the food depletes below some critical level. In nature, however, prey individuals may regularly move among food patches, even in the apparent absence of food depletion. We suggest that such prey movement is part of a predator-prey "shell game", in which predators attempt to learn prey location, and the prey attempt to be unpredictable in space. We investigate this shell game using an individual-based model that allows predators to update information about prey location, and permits prey to move with some random component among patches, but with reduced energy intake. Our results show the best prey strategy depends on what the predator does. A non-learning (randomly moving) predator favors non-moving prey – moving prey suffer higher starvation and predation. However, a learning predator favors prey movement. In general, the best prey strategy involves movement biased toward, but not completely committed to, the richer food patch. The strategy of prey movement remains beneficial even in combination with other anti-predator defenses, such as prey vigilance.     

Development of anuran locomotion: ethological and neurophysiological considerations.

There are dramatic quantitative and qualitative differences in the locomotor behavior of larval and juvenile frogs. Larvae (tadpoles) are primarily herbivourous and rely heavily on locomotion via undulations to acquire food and avoid predation. After metamorphosis, juvenile frogs adopt a carnivorous lifestyle and capture prey and avoid predators by remaining motionless in a place of concealment. When they must move, frogs locomote by means of ballistic hops or by more conventional walking. However, locomotion of both tadpoles and frogs can be considered of two fundamental functional types: (a) startle and escape; and (b) sustained locomotion. Neural mechanisms underlying startle responses and sustained locomotion in larvae and juveniles are described and possible ontogenetic relationships those behaviors are proposed. The role of different parts of the nervous system in the ontogeny of locomotion, as well as nonneuronal factors, are described. Results show that the transition from tadpole-like behavior to frog-like behavior is not a simple function of maturation of central locomotor controls. Rather, it results from a complex interaction of central nervous system maturation, morphological change, and a change in habitat preference. Examples of similar multidimensional control of behavioral ontogeny in other species are described, and it is argued that to understand the ontogeny of behavior, one must investigate contributions made at all levels, from the neuronal to the environmental.     

Field use of maximal sprint speed by collared lizards (Crotaphytus collaris): compensation and sexual selection

To understand how selection acts on performance capacity, the ecological role of the performance trait being measured must be determined. Knowing if and when an animal uses maximal performance capacity may give insight into what specific selective pressures may be acting on performance, because individuals are expected to use close to maximal capacity only in contexts important to survival or reproductive success. Furthermore, if an ecological context is important, poor performers are expected to compensate behaviorally. To understand the relative roles of natural and sexual selection on maximal sprint speed capacity we measured maximal sprint speed of collared lizards (Crotaphytus collaris) in the laboratory and field-realized sprint speed for the same individuals in three different contexts (foraging, escaping a predator, and responding to a rival intruder). Females used closer to maximal speed while escaping predators than in the other contexts. Adult males, on the other hand, used closer to maximal speed while responding to an unfamiliar male intruder tethered within their territory. Sprint speeds during foraging attempts were far below maximal capacity for all lizards. Yearlings appeared to compensate for having lower absolute maximal capacity by using a greater percentage of their maximal capacity while foraging and escaping predators than did adults of either sex. We also found evidence for compensation within age and sex classes, where slower individuals used a greater percentage of their maximal capacity than faster individuals. However, this was true only while foraging and escaping predators and not while responding to a rival. Collared lizards appeared to choose microhabitats near refugia such that maximal speed was not necessary to escape predators. Although natural selection for predator avoidance cannot be ruled out as a selective force acting on locomotor performance in collared lizards, intrasexual selection for territory maintenance may be more important for territorial males.     

Influence of predation risk and plant structure on vigilance and intermittent locomotion in <Emphasis Type="Italic">Microcavia australis</Emphasis> (Rodentia,Caviidae)

The aim of this study was to analyze and compare vigilance behavior and intermittent locomotion at two sites (El Leoncito and Ñacuñán, Argentina) that differ in predation risk, plant structure, and plant resource availability. Subjects were lesser cavies (Microcavia australis), a social species that is semi-fossorial, diurnal, and native to South America. Continuous focal sampling was conducted during the day, at times of food shortage, food abundance, and reproduction from 2003 to 2005. The proportion of time spent vigilance was significantly higher at Ñacuñán, where vigilance peaked at midday and reached a minimum in the evening. This midday peak of vigilance at Ñacuñán was associated with a midday peak of danger from raptors as indicated by a raptor activity peak at that time. In contrast, both vigilance and predator activity at El Leoncito were constant through the day. Records of intermittent locomotion and number and duration of pauses in locomotion were significantly higher at El Leoncito, a difference that may have been due to the need for greater vigilance while moving across areas of less protective cover at this site.     

Vigilance in Przewalski's gazelle: effects of sex, predation risk and group size

Vigilance in social animals is often aimed at detecting predators. Many social and environmental factors influence vigilance, including sex, predation risk and group size. During the summer of 2007, we studied Przewalski's gazelle Procapra przewalskii , an endemic ungulate to the Qinghai-Tibet Plateau, to test whether and how these three factors affect vigilance. We distinguished groups consisting of males, mothers with lambs and females without lambs making observations on groups in the presence or absence of nearby predators. We assessed the group-size effect on vigilance and how this varied with levels of predation risk and sex. Males and mothers scanned longer and with a higher frequency than females without lambs. Individuals were more vigilant under direct predation threat. Although vigilance generally decreased with group size, the extent of the decrease was independent of predation risk and was not significant in males. The results suggest that mothers are more vigilant suggesting greater vulnerability and that males may have increased their vigilance to compete for higher social ranks. The positive correlation between vigilance and predation risk and the negative correlation between vigilance and group size are consistent with earlier findings, but we failed to find an interaction between group size and predation risk on vigilance perhaps because vigilance levels are low even in small groups, thus making similar vigilant upward adjustments in both small and large groups.