Marine subsidies change short‐term foraging activity and habitat utilization of terrestrial lizards

Resource pulses are brief periods of unusually high resource abundance. While population and community responses to resource pulses have been relatively well studied, how individual consumers respond to resource pulses has received less attention. Local consumers are often the first to respond to a resource pulse, and the form and timing of individual responses may influence how the effects of the pulse are transmitted throughout the community. Previous studies in Bahamian food webs have shown that detritivores associated with pulses of seaweed wrack provide an alternative prey source for lizards. When seaweed is abundant, lizards (Anolis sagrei) shift to consuming more marine‐derived prey and increase in density, which has important consequences for other components of the food web. We hypothesized that the diet shift requires individuals to alter their habitat use and foraging activity and that such responses may happen very rapidly. In this study, we used recorded video observations to investigate the immediate responses of lizards to an experimental seaweed pulse. We added seaweed to five treatment plots for comparison with five control plots. Immediately after seaweed addition, lizards decreased average perch height and increased movement rate, but these effects persisted for only 2 days. To explore the short‐term nature of the response, we used our field data to parametrize heuristic Markov chain models of perch height as a function of foraging state. These models suggest a “Synchronized‐satiation Hypothesis,” whereby lizards respond synchronously and feed quickly to satiation in the presence of a subsidy (causing an initial decrease in average perch height) and then return to the relative safety of higher perches. We suggest that the immediate responses of individual consumers to resource pulse events can provide insight into the mechanisms by which these consumers ultimately influence community‐level processes.     

Versuche zum Nachweis eines Riechvermögens von Anolis l. lineatopus (Rept., Iguanidae)

Various experiments with 56 captive-born lizards, Anolis lineatopus, involving up to four replicates strongly suggest that the species is capable of olfaction. The readiness to attack four diverse prey species (cricket, Mediterranean flour moth, German cockroach, wood ant) was neither influenced by odour cues nor by the degree of novelty of the prey item; odour cues were those of the prey offered in a trial and four pure non-prey odourous compounds. Approximately 80 % of the total variance of the two attack measures employed could be accounted for by the individuality of Ss and by fluctuations of attack readiness.     

Variation in the effect of profitability on prey size selection by the lacertid lizard Psammodromus algirus

Maximizing the average rate of energy intake (profitability) may not always be the optimal foraging strategy for ectotherms with relatively low energy requirements. To test this hypothesis, we studied the feeding behaviour of captive insectivorous lizards Psammodromus algirus, and we obtained experimental estimates of prey mass, handling time, profitability, and attack distance for several types of prey. Handling time increased linearly with prey mass and differed significantly among prey types when prey size differences were controlled for, and mean profitabilities differed among prey taxa, but profitability was independent of prey size. The attack distance increased with prey length and with the mobility of prey, but it was unrelated to profitability. Thus, lizards did not seem to take account of the rate of energy intake per second as a proximate cue eliciting predatory behavior. This information was combined with pitfall-trap censuses of prey (in late April, mid-June and late July) that allowed us to compare the mass of the prey captured in the environment with that of the arthropods found in the stomachs of sacrificed free-living lizards. In April, when food abundance was low and lizards were reproducing, profitability had a pronounced effect on size selection and lizards selected prey larger than average from all taxa except the least profitable ones. As the active season progressed, and with a higher availability of food, the number of prey per stomach decreased and their mean ize increased. The effect of profitability on size selection decreased (June) and eventually vanished (July–August). This variation is probably related to seasonal changes in the ecology of lizards, e.g. time minimization in the breeding season as a means of saving time for nonforaging activities versus movement minimization by selecting fewer (but larger) prey in the postbreeding season. Thus, the hypothesis that maximizing profitability could be just an optional strategy for a terrestrial ectothermic vertebrate was supported by our data.     

Experimental evolution of a microbial predator's ability to find prey

Foraging theory seeks to explain how the distribution and abundance of prey influence the evolution of predatory behaviour, including the allocation of effort to searching for prey and handling them after they are found. While experiments have shown that many predators alter their behaviour phenotypically within individual lifetimes, few have examined the actual evolution of predatory behaviour in light of this theory. Here, we test the effects of prey density on the evolution of a predator's searching and handling behaviours using a bacterial predator, Myxococcus xanthus. Sixteen predator populations evolved for almost a year on agar surfaces containing patches of Escherichia coli prey at low or high density. Improvements in searching rate were significantly greater in those predators that evolved at low prey density. Handling performance also improved in some predator populations, but prey density did not significantly affect the magnitude of these gains. As the predators evolved greater foraging proficiency, their capacity diminished to produce fruiting bodies that enable them to survive prolonged periods of starvation. More generally, these results demonstrate that predators evolve behaviours that reflect at least some of the opportunities and limitations imposed by the distribution and abundance of their prey.     

The effect of prey movement on attack behavior and patch residence decision rules of wolf spiders (Araneae: Lycosidae)

We used a video imaging technique to test the effects of prey movement on attack behavior and foraging patch residence time decision rules of wolf spiders. TwelveSchizocosa ocreata (Hentz) (Lycosidae) were tested in an artificial foraging patch stimulus chamber consisting of a microscreen television displaying a computer digitized, animated image of a cricket. Four prey movement treatments were used: (1) a blank screen, (2) a stationary cricket control, (3) a cricket moving for 1 min, and (4) a cricket moving for 10 min. Spiders stayed significantly longer in treatments with higher cricket activity. Spiders also stayed longer when they attacked the stimulus than when they did not. The distribution of patch residence times of spiders indicates a decision rule based on a fixed probability of leaving.     

Resource Management in the Orb-Weaving Spider Zygiella x-notata: I. Influence of Spider Behaviour and Available Prey

Management of resource exploitation by the orb-weaving spider Zygiella x-notata was investigated in relation to available prey. Prey capture is studied according to prey type (cricket or fly), and the spider's behaviour (waiting without prey or consuming other prey), at the moment when a prey item was placed onto the web. Characteristics of the spider's exploitation of the previous item (feeding time, and quantities ingested) were analysed according to the type and quantity of prey presented, (1 or 2 items at 1-h intervals). Capture time did not vary in relation to prey type, nor spider activity. However, duration and quantities of certain behavioural components (biting and wrapping) did vary according to prey type and the spider's behaviour, at the moment of presentation of prey. Whatever the type of prey received, if the first item consumed was a cricket, the spiders reduced time spent feeding on this prey without any decrease in the quantities ingested.     

Wall lizards combine chemical and visual cues of ambush snake predators to avoid overestimating risk inside refuges

The threat sensitivity hypothesis assumes that multiple cues from a predator should contribute in an additive way to determine the degree of risk-sensitive behaviour. The ability to use multiple cues in assessing the current level of predation risk should be especially important to prey exposed to multiple predators. Wall lizards, Podarcis muralis, respond to predatory attacks from birds or mammals by hiding inside rock crevices, where they may encounter another predator, the smooth snake, Coronella austriaca. We investigated in the laboratory whether chemical cues may be important to wall lizards for detection of snakes. The greater tongue-flick rate and shorter latency to first tongue-flick in response to predator scents indicated that lizards were able to detect the snakes' chemical cues. We also investigated the use of different predatory cues by lizards when detecting the presence of snakes within refuges. We simulated successive predator attacks and compared the propensity of lizards to enter the refuge and time spent within it for predator-free refuges, refuges containing either only visual or chemical cues of a snake, or a combination of these. The antipredatory response of lizards was greater when they were exposed to both visual and chemical cues than when only one cue was presented, supporting the threat sensitivity hypothesis. This ability may improve the accuracy of assessments of the current level of predation risk inside the refuge. It could be especially important in allowing lizards to cope with threats posed by two types of predators requiring conflicting prey defences.     

The effects of prey species on food conversion efficiency and growth of an insectivorous lizard

Little is known about the effects of different prey species on lizard growth. We conducted a 6‐week study to determine the relative effects of prey species on growth parameters of hatchling western fence lizards, Sceloporus occidentalis. Lizards were fed house cricket nymphs, Acheta domesticus, or mealworm larvae, Tenebrio molitor. The effects of prey species on growth were determined by measuring prey consumption, gross conversion efficiency of food [gain in mass (g)/food consumed (g)], gain in mass, and gain in snout–vent length. Lizards grew well on both the prey species. However, lizards that fed on crickets consumed a significantly higher percentage of their body mass per day than those fed mealworms. Nevertheless, lizards that consumed mealworms ingested significantly more metabolizable energy, had significantly higher food conversion efficiencies, significantly higher daily gains in mass, and significantly greater total growth in mass than lizards that fed on crickets. Zoo Biol 27:181–187, 2008. © 2008 Wiley‐Liss, Inc.     

Influence of gaze and directness of approach on the escape responses of the Indian rock lizard, Psammophilus dorsalis (Gray, 1831)

Animals often evaluate the degree of risk posed by a predator and respond accordingly. Since many predators orient their eyes towards prey while attacking, predator gaze and directness of approach could serve as conspicuous indicators of risk to prey. The ability to perceive these cues and discriminate between high and low predation risk should benefit prey species through both higher survival and decreased energy expenditure. We experimentally examined whether Indian rock lizards (Psammophilus dorsalis) can perceive these two indicators of predation risk by measuring the variation in their fleeing behaviour in response to type of gaze and approach by a human predator. Overall, we found that the gaze and approach of the predator influenced flight initiation distance, which also varied with attributes of the prey (i.e. size/sex and tail-raise behaviour). Flight initiation distance (FID) was 43% longer during direct approaches with direct gaze compared with tangential approaches with averted gaze. In further, exploratory, analyses, we found that FID was 23% shorter for adult male lizards than for female or young male (FYM) lizards. In addition, FYM lizards that showed a tail-raise display during approach had a 71% longer FID than those that did not. Our results suggest that multiple factors influence the decision to flee in animals. Further studies are needed to test the generality of these factors and to investigate the proximate mechanisms underlying flight decisions.     

Wind-sensitive interneurones in the spider CNS (Cupiennius salei ): directional information processing of sensory inputs from trichobothria on the walking legs

Spiders can use air particle movements to localize moving prey. We studied the responses of 32 wind-sensitive interneurones in the hunting spider Cupiennius salei to prey stimuli. Stimulation with a tethered flying fly or with artificial air pulses activated plurisegmental interneurones that responded to changes in air movement velocity and were thus well suited to represent the highly fluctuating air stream typical of prey stimuli. In most interneurones (n = 18) the responses to the stimulation of different legs were not significantly different from each other. Different interneurones had different response characteristics and their latencies largely overlapped suggesting that there is parallel processing of the signals by populations of interneurones with different response characteristics. In two interneurones the number of spikes and the spiking pattern elicited by stimulation of each of the eight legs markedly differed depending on the leg stimulated. These neurones may play an important role in directional information processing. Stimulation of the adjacent legs from front to back or from back to front revealed two interneurones sensitive to the direction of successive stimulation of the legs. These neurones may be able to detect the motion of an air movement source in a preferred direction and thus act as nearfield motion detectors to localize a moving prey item. Accepted: 28 September 1996     

Chemosensory responses to foods by an herbivorous acrodont lizard, <Emphasis Type="Italic">Uromastyx aegyptius</Emphasis>

Actively foraging lizards use the lingual-vomeronasal system to identify prey by chemical cues, but insectivorous ambush foragers do not. The major clade Iguania includes numerous herbivores and omnivores; among them, two iguanid and one agamine species identify plant and animal foods by tongue flicking, and data suggest that the leiolepidine Uromastyx acanthinurus may as well. We conducted experiments on chemosensory response to food by the herbivorous U. aegyptius. When chemical stimuli were presented on cotton balls in experiment 1, the lizards exhibited greater responsiveness (tongue-flick attack scores) to chemical stimuli from crickets and a preferred plant food (dandelion flowers) than from deionized water. When chemical stimuli were on ceramic tiles in experiment 2, the lizards exhibited greater total tongue flicks to cricket stimuli than to any other stimuli, and to dandelion than to deionized water. Lizards bit more frequently in response to cricket and dandelion cues than to stimuli from a nonpreferred plant (carrot) and deionized water. Tongue-flick attack scores were greater in response to cricket and dandelion stimuli than to carrot or water stimuli. These findings are consistent with the hypothesis that herbivores, even those having ambush-foraging ancestors, use chemical cues to identify potential foods. The data support the hypothesis that chemosensory responses correspond to diet. Because most lizards are generalist predators, studies of herbivorous species can provide important information on possible evolutionary adjustment of chemosensory response to dietary shifts. Electronic Publication     

Resource Management in the Orb-Weaving Spider Zygiella x-notata II. Ability to Manage a Set of Various Prey

Management of a set of various prey received ‘simultaneously’ by the orb-weaving spider Zygiella x-notata is investigated in relation to prey species, weight, order of arrival in the web, and factors concerning the spider's state of nourishment. When the spiders encountered several different prey ‘simultaneously’, they subdued all of them, took one of them to begin consumption, and stored the others in the web. The first item for consumption was chosen according to the characteristics of the available prey. Factors concerning previous feeding influenced the spiders's choice of the first item to be ingested. Exploitation of the first item, estimated by the quantity of food extracted and the time necessary for this extraction, depended on the type and the quantity of prey at the spider's disposal. When the spiders received several prey, they spent less time feeding on the first cricket compared with spiders receiving only one cricket, while no difference was found when the first prey item exploited was a fly.     

Dangerous food: lacking venom and constriction,how do snake‐like lizards (Lialis burtonis,Pygopodidae) subdue their lizard prey?

Snakes are renowned for their ability to subdue and swallow large, often dangerous prey animals. Numerous adaptations, including constriction, venom, and a strike-and-release feeding strategy, help them avoid injury during predatory encounters. Burton's legless lizard ( Lialis burtonis Gray, Pygopodidae) has converged strongly on snakes. It is functionally limbless and feeds at infrequent intervals on relatively large prey items (other lizards) capable of inflicting a damaging bite. However, L. burtonis possesses neither venom glands, nor the ability to constrict prey. We investigated how L. burtonis subdues its prey without suffering serious retaliatory bites. Experiments showed that lizards modified their strike precision according to prey size; very large prey were always struck on the head or neck, preventing them from biting. In addition, L. burtonis delayed swallowing large lizards until they were incapacitated, whereas smaller prey were usually swallowed while still struggling. Lialis burtonis also displays morphological adaptations protecting it from prey retaliation. Its long snout prevents prey from biting, and it can retract its lidless eyes out of harm's way while holding onto a food item. The present study further clarifies the remarkable convergence between snakes and L. burtonis , and highlights the importance of prey retaliatory potential in predator evolution.  © 2007 The Linnean Society of London, Biological Journal of the Linnean Society, 2007, 91 , 719–727.     

Functional and Numerical Responses of Predators: Where Do Vipers Fit in the Traditional Paradigms?

Snakes typically are not considered top carnivores, yet in many ecosystems they are a major predatory influence. A literature search confirmed that terrestrial ectotherms such as snakes are largely absent in most discussions of predator‐prey dynamics. Here, we review classical functional and numerical responses of predator‐prey relationships and then assess whether these traditional views are consistent with what we know of one group of snakes (true vipers and pitvipers: Viperidae). Specifically, we compare behavioural and physiological characteristics of vipers with those of more commonly studied mammalian (endothermic) predators and discuss how functional and numerical responses of vipers are fundamentally different. Overall, when compared to similar‐sized endotherms, our analysis showed that vipers have: (i) lower functional responses owing primarily to longer prey handling times resulting from digestive limitations of consuming large prey and, for some adults, tolerance of fasting; (ii) stronger numerical responses resulting from higher efficiency of converting food into fitness currency (progeny), although this response often takes longer to be expressed; and (iii) reduced capacity for rapid numerical responses to short‐term changes in prey abundance. Given these factors, the potential for viperids to regulate prey populations would most likely occur when prey populations are low. We provide suggestions for future research on key issues in predator‐prey relationships of vipers, including their position within the classical paradigms of functional and numerical responses.     

Not All Signals are Equal: Male Brown Anole Lizards (Anolis sagrei) Selectively Decrease Pushup Frequency Following a Simulated Predatory Attack

The use of conspicuous communication signals often increases a signaler's risk of predation. Many species communicate with a repertoire of signals that may differ in their conspicuousness to predators. Few studies have examined the ability of prey to selectively decrease the use of individual signals in their displays under heightened predation risk. Here, I examined the behavior of male brown anole lizards (Anolis sagrei) in response to a simulated predatory attack from a model kestrel. This species communicates with three major visual signal types, the head‐bob, pushup, and dewlap extension, which vary in their motion and spectral characteristics. I predicted that lizards would decrease frequencies of the dewlap extension and pushup following the attack, but not the head‐bob. Males modulated their use of individual signals by decreasing pushup rates, but not head‐bob rates. Decreases in dewlap frequency were marginally significant. One explanation for these results is that lizards decrease frequencies of signal types based partly on their conspicuousness. The energetic cost of each signal type may be an equally important factor that determines the signaler's response to predators, particularly if a predatory attack is perceived as imminent.     

Invertebrate prey and predatory behaviour of the omnivorous African tortoise Kinixys spekii

Invertebrates form an important part of the diet in the omnivorous African tortoises of the genus Kinixys. Millipede prey of Kinixys spekii in Zimbabwe had a mean volume of 0.9 ml, and made up 64.7% of invertebrate food by volume; beetles made up most of the remainder and had a mean volume of 1.4 ml. The mean mass of invertebrate prey was 0.19% of tortoise mass, a similar value of relative prey mass to many insectivorous lizards. Tortoises preferentially attacked moving millipedes at or just behind the head, but predatory behaviour was otherwise unspecialized, with tortoises killing millipedes by ingesting them in pieces. Handling times of millipede prey varied significantly with relative prey mass, defensive behaviour, and direction of ingestion. The profitability (mass intake/handling time) of millipedes was maximal at a relative prey mass of 0.2%; the basis on which prey are selected is discussed.     

Are predatory birds effective secondary seed dispersers?

We have studied the unusual phenomenon of secondary seed dispersal of Lycium intricatum seeds on a small oceanic Atlantic island (Alegranza, Canarian Archipelago) in which a small frugivorous lizard ( Gallotia atlantica ) and two different predatory birds participate, a shrike ( Lanius excubitor ) and a kestrel ( Falco tinnunculus ). Endemic lizards that are common prey of both bird species consume Lycium fruits. Lizard remains were significantly matched with the presence of Lycium fruits in the regurgitation pellets of the two predatory birds. Seeds were found in 7.3% of the lizard droppings, 31.0% of kestrel pellets and 55.7% of shrike regurgitations. The mean number of seeds per dropping or pellet was 4.8 ±4 in lizard, 20.2 ±34.5 in shrike and 6.7 ±8.1 in kestrel. The percentage of viable seeds showed significant differences among all four treatments, decreasing in the following direction: seeds collected directly from plants (98.0%), shrikes (88.0%), lizards (72.3%), and kestrels (31.7%). Seeds from Lycium fresh fruits and shrike pellets showed significantly higher germination rates than those from lizard droppings and kestrel pellets. While lizards and shrikes are effective seed dispersers, kestrel gut treatment decreases seed viability. Seed viability is always higher than seed germination in each of the four treatments. In this island environment, Lycium seeds are under an important random influence during the seed dispersal process. Secondary seed dispersal seems to acquire a relevant dimension in small and remote insular environments or isolated continental systems where interactions among the different elements involved are intense, all of them are abundant native residents, and they have been coexisting for a long time. © 2002 The Linnean Society of London, Biological Journal of the Linnean Society , 2002, 75 , 345–352.     

Learning affects prey selection in larvae of a generalist coccinellid predator

Under natural conditions, generalist predatory insects have to cope with a variety of potential prey species that are not all equally suitable. Under these circumstances, learning may be adaptive if it allows adjustment to variations in resource quality and availability. Under laboratory conditions, we examined the learning ability and memory in the prey selection process of larvae of the predatory coccinellid Coleomegilla maculata ssp. lengi Timberlake (Coleoptera: Coccinellidae). Using choice tests, we studied prey rejection behaviour of C. maculata fourth instars towards prey of different quality and we also tested the influence of hunger and prior experience with other food types on the prey rejection behaviour of coccinellid larvae. Coleomegilla maculata larvae gradually changed their behaviour and rejected low‐quality hosts more frequently, whereas high‐quality hosts were nearly always accepted. After 48 h, the learned behaviour appeared to be partially forgotten. Hunger and experience with other food types prior to the test had little effect on the gradual change of behaviour but the quality of the food ingested influenced the initial level of prey rejection. Our results demonstrate that (1) C. maculata larvae can adjust their prey selection behaviour with experience to reject progressively less suitable prey, and (2) previous experience with other prey types can influence their initial preference.     

Can Wall Lizards Combine Chemical and Visual Cues to Discriminate Predatory from Non‐Predatory Snakes Inside Refuges?

The ability to use multiple cues in assessing predation risk is especially important to prey animals exposed to multiple predators. Wall lizards, Podarcis muralis, respond to predatory attacks from birds in the open by hiding inside rock crevices, where they may encounter saurophagous ambush smooth snakes. Lizards should avoid refuges with these snakes, but in refuges lizards can also find non‐saurophagous viperine snakes, which lizards do not need to avoid. We investigated in the laboratory whether wall lizards used different predator cues to detect and discriminate between snake species within refuges. We simulated predatory attacks in the open to lizards, and compared their refuge use, and the variation in the responses after a repeated attack, between predator‐free refuges and refuges containing visual, chemical, or visual and chemical cues of saurophagous or non‐saurophagous snakes. Time to enter a refuge was not influenced by potential risk inside the refuge. In contrast, in a successive second attack, lizards sought cover faster and tended to increase time spent hidden in the refuge. This suggests a case of predator facilitation because persistent predators in the open may force lizards to hide faster and for longer in hazardous refuges. However, after hiding, lizards spent less time in refuges with both chemical and visual cues of snakes, or with chemical cues alone, than in predator‐free refuges or in refuges with snake visual cues alone, but there were no differences in response to the two snake species. Therefore, lizards could be overestimating predation risk inside refuges. We discuss which selection pressures might explain this lack of discrimination of predatory from similar non‐predatory snakes.     

Experience with methyl salicylate affects behavioural responses of a predatory mite to blends of herbivore-induced plant volatiles

Many natural enemies of herbivorous arthropods use herbivore‐induced plant volatiles to locate their prey. These foraging cues consist of mixtures of compounds that show a considerable variation within and among plant–herbivore combinations, a situation that favours a flexible approach in the foraging behaviour of the natural enemies. In this paper, we address the flexibility in behavioural responses of the predatory mite Phytoseiulus persimilis Athias‐Henriot (Acari: Phytoseiidae) to herbivore‐induced plant volatiles. In particular, we investigated the effect of experience with one component of a herbivore‐induced volatile blend: methyl salicylate (MeSA). We compared the responses of three groups of predatory mites: (1) those reared from egg to adult on Tetranychus urticae Koch (Acari: Tetranychidae) on lima bean plants (Phaseolus lunatus L. that produces MeSA), (2) those reared on T. urticae on cucumber (Cucumus sativus L. that does not produce MeSA), and (3) those reared on T. urticae on cucumber in the presence of synthetic MeSA. Exposure to MeSA during the rearing period (groups 1 and 3) resulted in an attraction to the single compound MeSA in a Y‐tube olfactometer. Moreover, exposure to MeSA affected the choice of predatory mites between two volatile blends that were similar, except for the presence of MeSA. Predators reared on lima bean plants preferred the volatile blend from T. urticae‐induced lima bean (including MeSA) to the volatile blend from jasmonic‐acid induced lima bean (lacking MeSA), but predators reared on cucumber preferred the volatile blend from the latter. Predatory mites reared on cucumber in the presence of synthetic MeSA did not discriminate between these two blends. Exposure to MeSA for 3 days in the adult phase, after rearing on cucumber, also resulted in attraction to the single compound MeSA. We conclude that a minor difference in the composition of the volatile blend to which a predatory mite is exposed can explain its preferences between two odour sources.