Resembling a Viper: Implications of Mimicry for Conservation of the Endangered Smooth Snake

The phenomenon of Batesian mimicry, where a palatable animal gains protection against predation by resembling an unpalatable model, has been a core interest of evolutionary biologists for 150 years. An extensive range of studies has focused on revealing mechanistic aspects of mimicry (shared education and generalization of predators) and the evolutionary dynamics of mimicry systems (co‐operation vs. conflict) and revealed that protective mimicry is widespread and is important for individual fitness. However, according to our knowledge, there are no case studies where mimicry theories have been applied to conservation of mimetic species. Theoretically, mimicry affects, for example, frequency dependency of predator avoidance learning and human induced mortality. We examined the case of the protected, endangered, nonvenomous smooth snake (Coronella austriaca) that mimics the nonprotected venomous adder (Vipera berus), both of which occur in the Åland archipelago, Finland. To quantify the added predation risk on smooth snakes caused by the rarity of vipers, we calculated risk estimates from experimental data. Resemblance of vipers enhances survival of smooth snakes against bird predation because many predators avoid touching venomous vipers. Mimetic resemblance is however disadvantageous against human predators, who kill venomous vipers and accidentally kill endangered, protected smooth snakes. We found that the effective population size of the adders in Åland is very low relative to its smooth snake mimic (28.93 and 41.35, respectively).Because Batesian mimicry is advantageous for the mimic only if model species exist in sufficiently high numbers, it is likely that the conservation program for smooth snakes will fail if adders continue to be destroyed. Understanding the population consequences of mimetic species may be crucial to the success of endangered species conservation. We suggest that when a Batesian mimic requires protection, conservation planners should not ignore the model species (or co‐mimic in Mullerian mimicry rings) even if it is not itself endangered. Implications of mimicry for Conservation of the endangered smooth snake     

Butterfly effects in mimicry? Combining signal and taste can twist the relationship of Müllerian co-mimics

Müllerian co-mimics are aposematic species that resemble each other; sharing a warning signal is thought to be mutually beneficial for the co-mimics by reducing per capita predation risk. In Batesian mimicry, edible mimics avoid predation by resembling an aposematic model species. The protection of both the model and the mimic is weakened when the mimics are abundant compared to the models. The quasi-Batesian view suggests that defended (Müllerian) co-mimics, when unequal in their defences, could also show a Batesian-like trend of increasing mortality with increasing abundance of a less defended “mimic”. We manipulated frequencies of unequally distasteful artificial co-mimics that were prey for great tits. The co-mimics had different signals (imperfect mimicry) but were equally preferred by the birds when palatable. Unexpectedly, when unpalatable, one of the signals was easier for the birds to learn to avoid. Consequently, during predator learning, the signal design of the prey strongly affected mortality of the co-mimics; there was an interaction between the signal and frequency treatments, but increasing the frequency of a less defended “mimic” did not increase co-mimic mortalities as predicted. In contrast, in a memory test that followed, the effect of signal design disappeared; if the birds had experienced high frequency of “mimics” during learning, co-mimic mortalities did subsequently increase. Since the effect of co-mimic frequencies on mortalities changed depending on the signal design of the prey and predator experience, the results suggest that mimetic relationship may be an unpredictable interplay of several factors in addition to taste and abundance.     

Facultative pheromonal mimicry in snakes: “she-males” attract courtship only when it is useful

Males of many animal species mimic females, and thereby deceive rival males. Facultative shifts in posture, color, or movement allow a male using visually-based mimicry to adopt and terminate mimicry rapidly. Pheromonal mimicry is rare in vertebrates perhaps because it is difficult to redeploy pheromones rapidly enough to adjust male tactics to local conditions. In Manitoba garter snakes (Thamnophis sirtalis parietalis), female mimicry benefits males immediately after they have emerged from hibernation. While the snakes are cold and slow, courtship warms them and protects them against predatory crows. This benefit disappears as soon as the snakes are warm. We show that (unlike females) she-male garter snakes attract courting males only when they are cold. Low temperatures may suppress volatility of “less attractive” components of the pheromones (saturated methyl ketones) that she-males use to attract courtship, allowing male snakes to function as transvestites only when this tactic is beneficial.     

Does colour matter? The importance of colour in avoidance learning, memorability and generalisation

Aposematic species exploit the ability of predators to associate, for example, conspicuous colouration with the unprofitability of prey. We tested the importance of colour for avoidance learning, memory and generalisation in wild-caught great tits (Parus major). First, we determined the birds’ initial colour preferences for red, yellow, orange and grey artificial prey items. The birds showed some preferences, as they were more willing to eat grey prey as their first choice, but these were not strong preferences. We then trained birds to discriminate red, yellow or variable (red and yellow) signals from grey where colours signalled palatable and unpalatable food. In general, the birds learned the discrimination task equally well, irrespective of which colours signalled unpalatability, and subsequently remembered the distinction between previously palatable and previously unpalatable colours in the memorability test. We did not find strong evidence that variability in the signal affected learning or memory. Our results suggest that, in a task where birds must discriminate between palatable and unpalatable prey, it does not matter which specific colour signals unpalatability, although this might be context-dependent. To study whether training also affects responses to unconditioned stimuli, we included orange prey items in the memorability test. Although orange had been palatable in the initial preference test, the birds ate fewer orange prey items after they had been trained to avoid red, yellow or both colours (variable signal) as unpalatable prey, but did not change their preference when trained that these colourful signals were palatable. This indicates that generalisation occurred more readily after a negative experience than a positive experience, a situation that would potentially allow imperfect mimicry to occur.     

The anti-predator function of ‘eyespots’ on camouflaged and conspicuous prey

Animals utilise various strategies to reduce the risk of predation, including camouflage, warning colours and mimicry, and many of these protective signals promote avoidance behaviour in predators. For example, various species possess paired circular ‘eyespots’, which startle or intimidate predators, preventing or halting an attack. However, little is known of how the efficacy of such signals relates to the context in which they are found, and no studies have tested the relative effectiveness of anti-predator signals when on otherwise camouflaged and conspicuous prey. We find that the protective value of conspicuous wing spots, placed on artificial moth-like targets presented to wild birds in the field, is strongly affected by the attributes of the prey ‘animal’ on which they are found. Wing spots reduced predation when on conspicuous prey but were rendered ineffective when on otherwise camouflaged targets, indeed they increased the risk of predation compared to non-marked camouflaged controls. These results demonstrate how different anti-predator strategies may interact, and that protective signals can switch from being beneficial to costly under different contexts.     

A dynamic size-structured population model: does disturbance control size structure of a population of the massive coral Gardineroseris planulata in the Eastern Pacific?

Using a long-term data set of Gardineroseris planulata (Dana) on Uva Island reef, Panamá, we developed a simulation model that relates size-specific schedules of growth and partial mortality to predation by Acanthaster planci (Linnaeus) and El Niño–Southern Oscillation (ENSO)-related elevation of water temperature. We compared the model predictions to field observations for both the subpopulation of colonies that was used for model development and for the entire population. No statistically significant differences in the size-frequency distributions of the real and modeled coral populations were found for the subpopulation during any of 9 yr or for the entire population during 4 yr. These results suggested that the model relationships were reflecting field conditions. Longer-term (100 yr) simulations were conducted to assess the relative importance of predation and ENSO-related colony losses in determining the size structure of the coral population. Predation by A. planci was of overwhelming importance due to both stronger effects of predation (larger transitions) and the frequency of predation (yearly) compared to ENSO (episodically). Even the least-frequent predation scenario skewed the distribution toward smaller colonies, while simulations where populations were subjected to frequent ENSO events (≥3 yr) still maintained colonies in the largest size class. The model results suggested that sea star predators may not have been in the present abundance on this reef prior to the last 30 yr; with predators present, the model predicts that the distribution would be skewed toward smaller colonies.     

Are sea snakes pertinent bio-indicators for coral reefs? a comparison between species and sites

Classical sampling methods often miss important components of coral reef biodiversity, notably organisms that remain sheltered within the coral matrix. Recent studies using sea kraits (sea snakes) as bio-indicators suggest that the guild of predators represented by anguilliform fish (Congridae, Muraenidae, Ophichthidae, henceforth “eels” for simplicity) were far more abundant and diverse than previously suspected. In the current study, eel diversity (similarity and species richness indices) estimated via sea snake sampling (SSS) was compared among six areas of one of the main oceanic biodiversity hotspot of the Pacific Ocean (southwest lagoon of New Caledonia). Based on the eel diversity in the snakes’ diet, the results obtained in six areas, in two snake species, and using different estimates (ANOSIM, Shannon index…) were consistent, suggesting that SSS provided robust information. Analyses also suggested subtle, albeit significant, differences in the eel assemblages among islets. Such spatial differences are discussed in light of local management practices. As SSS is easy to use, cost-effective, and provides the best picture of eel assemblages to date, it can be employed to monitor the eel assemblages in addition to the snakes themselves in many areas of the Indo-Pacific Ocean, thereby providing an index of the top predator biodiversity of many coral reefs.     

Delayed plumage maturation in Lazuli buntings: tests of the female mimicry and status signalling hypotheses

The evolutionary importance of delayed plumage maturation (DPM) in passerines, the condition when more than 1 year is required to achieve adult-like coloration, remains highly contentious. Adaptive hypotheses propose that aggression from after 2nd-year (ASY) males or predation favors DPM in 2nd-year (SY) males, thereby increasing SY male survivorship or reproductive success. However, each hypothesis suggests a distinct selective mechanism explaining “how” this is accomplished. Alternatively, DPM may be a consequence of a nonadaptive molt constraint. We tested the female mimicry and status signalling hypotheses in territorial ASY male lazuli buntings (Passerinaamoena) using three sets of model presentation experiments. The female mimicry hypothesis proposes that dull SY male plumage deceptively mimics female plumage, and predicts that ASY males can not distinguish SY male from female plumage. The status signalling hypothesis proposes that dull SY male plumage honestly signals low competitive threat, and predicts that ASY males respond less aggressively to dull versus bright, ASY-like plumage. Contrary to the female mimicry hypothesis, ASY males distinguished between SY male and female plumage, as they were aggressive to SY male models exclusively and attempted to copulate with female models. Supporting the status signalling hypothesis, ASY males were significantly less aggressive to SY versus ASY male plumage. While DPM may result from a physiological constraint on bright SY male plumage, our results support the idea that dull plumage in an SY male's first breeding season may be maintained by selection to reduce aggression from ASY males, serving as a signal of competitive status. Received: 21 February 1997 / Accepted after revision: 16 June 1997     

Sexual conflict in the snake den

Red-sided garter snakes (Thamnophis sirtalis parietalis) court and mate in spring, soon after they emerge from large communal overwintering dens in south-central Manitoba. Because of a massive bias in the operational sex ratio, every female attracts intense courtship from dozens to hundreds of males. We suggest that this courtship constitutes significant ”harassment,” because it delays the females’ dispersal from the den and hence increases their vulnerability to predation. Small females may face the greatest costs, because they are less able to escape from amorous males (who court all females, even juvenile animals). Our measurements show that males are stronger and faster than females. Experimental trials confirm that the locomotor ability of females (especially small females) is greatly reduced by the weight of a courting male. Arena trials show that intense courtship stimulates females to attempt to escape. Remarkably, some females that are too small to produce offspring may nonetheless copulate. This precocious sexual receptivity may benefit juvenile females because copulation renders them unattractive to males, and thus allows them to escape more easily from the den. Female ”tactics” to escape male harassment may explain other puzzling aspects of garter snake biology including size-assortative mating, temporal patterns in dispersal from the den, avoidance of communal dens by young-of-the-year snakes, and female mimicry. Hence, sexual conflict may have influenced important features of the mating system and behavioral ecology of these animals. Received: 8 May 2000 / Revised: 28 July 2000 / Accepted: 30 July 2000     

Mortality is associated with social rank in the clown anemonefish (<Emphasis Type="Italic">Amphiprion percula</Emphasis>)

Elucidating the causes of post-recruitment mortality is a vital step toward understanding the population dynamics of coral reef fishes. Predation is often considered to be the primary proximate cause of mortality. It has, however, proven difficult to discern the relative contributions of predation and other processes, such as competition for food, shelter, or mates, to patterns of mortality. To determine which other processes might be important drivers of mortality patterns, factors related to mortality in the clown anemonefish Amphiprion percula (Lacepède, 1802) were examined. Patterns of mortality will not be driven by predation in A. percula, because these fish are well protected from predators by their close association with sea anemones. Mortality rates were based on the disappearance of known individuals from a population of 201, in 57 groups, during a 1-year field study (in 1997), in Madang Lagoon, Papua New Guinea. Mortality rate of A. percula was low (14% per annum) compared to other coral reef fish, probably due to the protection from predators afforded by the anemone. Six factors (reef, depth, anemone diameter, number of individuals, density, and standard length) showed no association with the probability of mortality (P>0.05). Rank was the only factor associated with the probability of mortality (P<0.03); low-rank individuals (ranks 4–6) suffered a higher mortality rate than high-rank individuals (ranks 1–3) (P<0.01). The most likely explanation for this pattern was that competition for rank, amongst individuals within an anemone, resulted in some individuals evicting their subordinates. Individuals probably competed for rank because it conferred access to reproduction, and not because it conferred access to food or shelter. Such competition for reproduction will be intense whenever some individuals obtain a greater share of reproduction than others do, and it may be an important process influencing the dynamics of coral reef fish populations.Electronic Supplementary Material Supplementary material is available for this article if you access the article at . A link in the frame on the left on that page takes you directly to the supplementary material.Communicated by J.P. Grassle, New Brunswick     

Bright plumage in the magpie: does it increase or reduce the risk of predation?

Conspicuous color patterns in birds may lead to increased risk of predation. Alternatively, bright birds may be aposematic or unprofitable prey, which leads to decreased predation. During four autumns, I examined whether the black-and-white plumage of (stuffed) magpies Pica pica increases or reduces the risk of attack by migrating goshawks Accipiter gentilis. Attack risk was higher for normal magpies than for cryptic, brown magpies whether the mounts were placed near one another or far apart. However, the brown magpie may have been avoided because of its novelty. For magpies and cryptic jays Garrulus glandarius exposed together, the attack risk was similar. In 2 years, magpie and jay mounts were also exposed far apart. In 1994, with an invasion of migrating jays, attack risk was much higher for jays than for magpies. In 1995, with a normal density of jays, hawks attacked the magpies more often. The results demonstrate frequency-dependent prey selection by goshawks, which would influence any predation cost of bright plumage. The attacks on normally colored magpie mounts suggest that magpies are not aposematic. Trials with photographs and human observers indicated that normal magpies were somewhat easier to detect than jays. The plumage of the magpie possibly increases the risk of predation, but may be favoured by sexual or social selection. Received: 18 January 1996 / Accepted after revision: 6 October 1996     

Circadian shelter occupancy patterns and predator–prey interactions of juvenile Caribbean spiny lobsters in a reef lagoon

The spiny lobster, Panulirus argus, is predominantly nocturnal, remaining inside shelters during the day and foraging outside at night, presumably to minimize predation risk. Predation risk generally decreases with increasing lobster size. Therefore, this study examined the hypothesis that size would influence this basic circadian pattern. Video cameras continuously recorded the shelter occupancy of juvenile lobsters (n = 72) having a carapace length (CL) of 30–62 mm that were tethered to shelters in a shallow reef lagoon. The lobsters’ shelter occupancy was 100% during the day, but declined linearly from shortly before sunset to a minimum of 50% shortly after midnight and then increased linearly, reaching 100% by 1 h after sunrise. The percent time the lobsters spent in the shelters followed a similar trend, but there was wide variability at night (0–100%) for individual lobsters. Lobsters left their shelters 2–30 times night⁻¹, with a majority of excursions lasting <10 min. These results suggest that juvenile P. argus minimize predation risk by remaining in their shelters as long as possible but offset the energetic cost of this behavior by foraging close to their shelters for several short periods at night. This emergence pattern contrasts with those of early benthic phase lobsters (<15 mm CL), which seldom leave their shelters, and adults (>80 mm CL), which have a dusk/early evening peak in activity and leave the shelter for extended periods of time during the night. Furthermore, a minimum shelter occupancy in the middle of the night appears especially well adapted to avoid exposure to daytime predators. Videotaped observations also included interactions between lobsters and two dominant lobster predators, the triggerfish, Balistes capriscus, and the octopus Octopus cf. vulgaris. Lobsters responded differently to these predators: remaining in the shelter when attacked by a triggerfish and fleeing the shelter when attacked by an octopus. Triggerfish were nearly twice as likely to attack a lobster that was outside of the shelter than inside. Once under attack, however, a lobster had nearly the same chance of surviving if it was inside or outside. Results suggest that the patterns of shelter use and emergence change as lobsters grow, probably reflecting the interplay between perception of predation risk and the need to forage. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Predation risk makes bees reject rewarding flowers and reduce foraging activity

In the absence of predators, pollinators can often maximize their foraging success by visiting the most rewarding flowers. However, if predators use those highly rewarding flowers to locate their prey, pollinators may benefit from changing their foraging preferences to accept less rewarding flowers. Previous studies have shown that some predators, such as crab spiders, indeed hunt preferentially on the most pollinator-attractive flowers. In order to determine whether predation risk can alter pollinator preferences, we conducted laboratory experiments on the foraging behavior of bumble bees (Bombus impatiens) when predation risk was associated with a particular reward level (measured here as sugar concentration). Bees foraged in arenas containing a choice of a high-reward and a low-reward artificial flower. On a bee’s first foraging trip, it was either lightly squeezed with forceps, to simulate a crab spider attack, or was allowed to forage safely. The foragers’ subsequent visits were recorded for between 1 and 4 h without any further simulated attacks. Compared to bees that foraged safely, bees that experienced a simulated attack on a low-reward artificial flower had reduced foraging activity. However, bees attacked on a high-reward artificial flower were more likely to visit low-reward artificial flowers on subsequent foraging trips. Forager body size, which is thought to affect vulnerability to capture by predators, did not have an effect on response to an attack. Predation risk can thus alter pollinator foraging behavior in ways that influence the number and reward level of flowers that are visited.     

Why do snakes have eyes? The (non-)effect of blindness in island tiger snakes (Notechis scutatus)

Large (to >1 m), diurnally active tiger snakes (Notechis scutatus) are abundant on Carnac Island, near the coast of Western Australia. Our behavioural and mark-recapture studies provide the first ecological data on this population, and reveal a surprising phenomenon. Many adult tiger snakes have had their eyes destroyed, apparently during nest defence, by silver gulls (Larus novaehollandiae). This loss of vision did not reduce the snakes' body condition (mass relative to length), or their rates of growth or survival (measured over a 12-month period). Blind male snakes trail-followed females, and mated successfully. Thus, destruction of a major sensory modality had no detectable effect on these predators. This result is strongly counter-intuitive, but mirrors an earlier report of congenital blindness (without ill-effects) in American viperid snakes. Similarities between the two systems (island populations, highly venomous snakes, reliance on sessile prey) clarify the circumstances under which the loss of vision does not reduce an organism's viability. These natural experiments support Gans' hypothesis of “momentarily excessive construction” in that the snakes possess a complex organ system that they do not actually require for successful feeding, survival or reproduction. Received: 30 November 1998 / Received in revised form: 24 April 1999 / Accepted: 9 May 1999     

Anguilliform fishes and sea kraits: neglected predators in coral-reef ecosystems

Despite intensive sampling efforts in coral reefs, densities and species richness of anguilliform fishes (eels) are difficult to quantify because these fishes evade classical sampling methods such as underwater visual census and rotenone poisoning. An alternative method revealed that in New Caledonia, eels are far more abundant and diverse than previously suspected. We analysed the stomach contents of two species of sea snakes that feed on eels (Laticauda laticaudata and L. saintgironsi). This technique is feasible because the snakes return to land to digest their prey, and (since they swallow their prey whole) undigested food items are identifiable. The snakes’ diet consisted almost entirely (99.6%) of eels and included 14 species previously unrecorded from the area. Very large populations of snakes occur in the study area (e.g. at least 1,500 individuals on a small coral islet). The snakes capture approximately 36,000 eels (972 kg) per year, suggesting that eels and snakes play key roles in the functioning of this reef ecosystem.     

Individual differences in nest defense in the colonial breeding Black-tailed Gulls

Often in colonial seabirds, all colony members are believed to defend against nest predators and experience equal nest predation risk. However, the variation of defense behavior among members and its reproductive consequences are largely unknown. We investigated (1) individual variation in the nest defense of breeding Black-tailed Gulls Larus crassirostris against a natural egg predator, the Jungle Crow Corvus macrorhynchos and (2) how this behavioral variation affects an individual’s own nest predation risk and that of their neighbors. Results were compared between 2 years where crow attack levels were manipulated to average 5 and 22 times normal rates (“low” and “high” predation risk years, respectively) by the placement of varying numbers of artificial nests containing unguarded eggs at the perimeter of the gull colony. In both years, 23–38% of parents, mostly males, showed “aggressive” defense behavior (strikes or chases) against crows and decoys. Other “non-aggressive” gulls showed no defense. In the year of low predation risk, intrusion rates by crows (landing within 0.5 m of an individual gull’s nest) were similar for aggressive and non-aggressive gulls. In the year of high predation risk, however, the rates of intrusion for aggressive gulls (4%) and for non-aggressive gulls with an aggressive neighbor (37%) were significantly lower than for non-aggressive gulls without an aggressive neighbor (76%). These results indicate that aggressive individuals reduce nest predation risk for themselves and conspecific neighbors in a colonially breeding species.     

Inter-specific variation in anti-predator behavior in sympatric species of kangaroo rat

Kangaroo rats, Dipodomys, occupy desert habitats with little cover and thus are under high predation risk from diverse predators. The behavior used to assess predation risk or to escape capture is unknown. We therefore compared anti-predator behavior of two sympatric species of kangaroo rat of different sizes, D. merriami and D. spectabilis. We first examined whether kangaroo rats use olfaction as a first line of defense against snake predation and tested the rats for their responses to scent extracted from two species of snake that live sympatrically with the kangaroo rats, the Mojave rattlesnake (Crotalus scutulatus) and the gopher snake (Pitophis melanoleucus). We also tested for species differences in anti-predator behavior through 15-min interactions between the kangaroo rats and free-moving gopher snakes. We found that D. spectabilis actively approached the scent of both rattlesnakes and gopher snakes more than controls of vegetable oil and evaporated solvent (Fig. 1). In contrast, D. merriami did not differentiate snake odors from controls in the experimental arena, but they sniffed the sand where a free-moving snake had passed more than D. spectabilis. Both species successfully avoided predation in encounters with live snakes. Although total numbers of approaches and withdrawals were similar (Fig. 2), D. spectabilis spent significantly more time within striking distance of the snake than D. merriami. D. spectabilis approached the head of the snake in 93% of its approaches and often engaged in nose to snout contact with the snake. If the snake struck, D. spectabilis jumped directly backward to avoid a strike and footdrummed at a safe distance. In contrast, D. merriami oriented to the snake more than D. spectabilis, but approached the head in only 41% of the approaches and rarely engaged in nose-to-snout contact. The snakes struck, hissed and decreased predatory approaches with D. spectabilis but not with D. merriami (Fig. 3). These results show that kangaroo rats can behaviorally influence the risk of being preyed on by snakes. The two species differ, however, in how they react to snakes. The larger D. spectabilis confronts snakes while the smaller D. merriami monitors snakes from a safe distance and avoids them.     

The ghost of predation future: threat-sensitive and temporal assessment of risk by embryonic woodfrogs

Amphibians are able to learn to recognize their future predators during their embryonic development (the ghost of predation future). Here, we investigate whether amphibian embryos can also acquire additional information about their future predators, such as the level of threat associated with them and the time of day at which they would be the most dangerous. We exposed woodfrog embryos (Rana sylvatica) to different concentrations of injured tadpole cues paired with the odor of a tiger salamander (Ambystoma tigrinum) between 1500 and 1700 hours for five consecutive days and raised them for 9 days after hatching. First, we showed that embryos exposed to predator odor paired with increasing concentrations of injured cues during their embryonic development subsequently display stronger antipredator responses to the salamander as tadpoles, thereby demonstrating threat-sensitive learning by embryonic amphibians. Second, we showed that the learned responses of tadpoles were stronger when the tadpoles were exposed to salamander odor between 1500 and 1700 hours, the time at which the embryos were exposed to the salamander, than during earlier (1100–1300 hours) or later (1900–2100 hours) periods. Our results highlight the amazing sophistication of learned predator recognition by prey and emphasize the importance of temporal considerations in experiments examining risk assessment by prey.     

Defensive secretion of a flightless grasshopper: Failure to prevent lizard attack

Summary We tested the role of a grasshopper defensive secretion in deterring lizard predation. Adults, but not young larvae, of the chemically defended lubber grasshopperRomalea guttata (=microptera) froth a volatile secretion when attacked by predators. The lizardAnolis carolinensis failed to strike juvenile lubbers (which lack secretion) in laboratory trials. Survivorship of palatable crickets loaded with secretion offered toA. carolinensis was not significantly different from survivorship of control crickets. In experiments designed to investigate if lizards learn an aversion to the secretion, striking times forSceloporus undulatus fed wax worms coated with secretion were not significantly different over three days of trials. Three primary conclusions are drawn from these data. First, the secretion may not be necessary for lubber protection from lizards. Second, lubber secretion does not appear to deter lizards from attacking or eating prey items. Third, lizards do not appear to develop an aversion to the secretion.     

Predation risk and foraging behavior of the hoary marmot in Alaska

Summary I observed hoary marmots for three field seasons to determine how the distribution of food and the risk of predation influenced marmots' foraging behavior. I quantified the amount of time Marmota caligata foraged in different patches of alpine meadows and assessed the distribution and abundance of vegetation eaten by marmots in these meadows. Because marmots dig burrows and run to them when attacked by predators, marmot-toburrow distance provided an index of predation risk that could be specified for different meadow patches.Patch use correlated positively with food abundance and negatively with predation risk. However, these significant relationships disappeared when partial correlations were calculated because food abundance and risk were intercorrelated. Using multiple regression, 77.0% of the variance in patch use was explained by a combination of food abundance, refuge burrow density, and a patch's distance from the talus where sleeping burrows were located. Variations in vigilance behavior (look-ups to search for predators while feeding) according to marmots' ages, the presence of other conspecifics, and animals' proximity to their sleeping burrows all indicated that predation risk influenced foraging.In a forage-manipulation experiment, the use of forage-enhanced patches increased six-fold, verifying directly the role of food availability on patch used. Concomitant with increased feeding, however, was the intense construction of refuge burrows in experimental patches that presumably reduced the risk of feeding. Thus, I suggest that food and predation risk jointly influence patch use by hoary marmots and that both factors must be considered when modeling the foraging behavior of species that can be predator and prey simultaneously.