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1.
Actively foraging lizards are capable of identifying prey using only chemical cues sampled by tongue-flicking, and the relatively few omnivorous and herbivorous lizards tested similarly can identify both animal and plant foods from chemical cues. Whether lizards that eat plants respond to cues specific to preferred plant types and whether there is geographic variability in responses to cues from various plants correlated with the importance of those plants in local diets is unknown. In three populations of an omnivorous lacertid, the Balearic lizard Podarcis lilfordi, we studied chemosensory sampling and feeding responses to chemical cues from plant and animal foods presented on cotton swabs. Each lizard population is endemic to one islet off the coast of Menorca, Balearic Islands, Spain. Lizards in all three populations discriminated chemical cues from plant and animal foods from control substances. Our results extend findings of prey chemical discrimination and plant chemical discrimination in omnivores, increasing confidence that correlated evolution has occurred between plant diet and chemosensory response to palatable plants. There were no consistent differences among populations in tongue-flicking and biting responses to stimuli from flowers of syntopic and allopatric plant species. The lizards may respond to cues indicative of palatability in a wide range of plant species rather than exhibiting strong responses only to locally available plant species. Nevertheless, tongue-flicking and biting frequencies varied among plant species, perhaps indicating food preferences. In addition, there were differences among populations in tongue-flick rates, latency to bite, and licking behavior. Licking was observed in only one lizard population as a response to floral chemicals from only one of the plants species tested, raising the possibility of a population-specific linkage between identification of a particular plant species and performance of an appropriate feeding response.  相似文献   

2.
If tongue-flicking is important to lizards to sample chemical cues permitting identification of foods, tongue-flicking and subsequent feeding responses should be adjusted to match diet. This hypothesis can be examined for plant foods because most lizards are insectivores, but herbivory/omnivory has evolved independently in many lizard taxa. Here we present experimental data on chemosensory responses to chemical cues from animal prey and palatable plants by three species of the scincine lizards. When tested with chemical stimuli presented on cotton swabs, the insectivorous Eumeces fasciatus responded strongly to prey chemicals but not to chemicals from plants palatable to omnivorous lizards or to pungent or odorless control stimuli. Two omnivorous species, E. schneideri and Scincus mitranus, responded more strongly to chemical cues from both prey and food plants than to the control chemicals. All available data for actively foraging lizards, including these skinks, show that they are capable of prey chemical discrimination, and insectivores do not exhibit elevated tongue-flicking or biting responses to chemical cues from palatable plants. In all of the several species of herbivores/omnivores tested, the lizards show elevated responses to both animal and plant chemicals. We suggest two independent origins of both omnivory and plant chemical discrimination that may account for the evolution of diet and food chemical discriminations in the eight species of skinks studied, five of which are omnivores. All data are consistent with the hypothesis that acquisition of omnivory is accompanied by acquisition of plant chemical discrimination, but data on a broad diversity of taxa are needed for a definitive comparative test of the evolutionary hypothesis. J. Exp. Zool. 287:327-339, 2000.  相似文献   

3.
Experimental tests show that the ball python (Python regius) has the ability to discriminate prey chemicals from control substances by tongue-flicking and exhibits a poststrike elevation in tongue-flicking rate (PETF). Prey chemical discrimination was revealed by significantly higher number of tongue-flicks and tongue-flick attack score in response to integumental chemicals from mice than to cologne or distilled water and by a higher frequency of biting in response to prey than control chemicals. PETF was indicated by higher tongue-flicking rates after biting than in several control conditions. Concurrent movements of the body suggest the operation of strike-induced chemosensory searching (SICS). Ecological factors affecting responses to prey chemicals, including defensive behaviors and characteristics of foraging behavior related to reliance on different sensory modalities, are discussed. The presence of PETF and SICS in a henophidian snake and in scleroglossan lizards suggests that these behaviors are plesiomorphic in snakes.  相似文献   

4.
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  相似文献   

5.
Cologne has been used extensively as a pungency control in experiments on chemosensory behavior to assess responses to an odorous, readily detectable stimulus that is irrelevant to the adaptive response being studied. However, undiluted cologne may be aversive, its effects might differ among brands, it might suppress responses to simultaneously presented chemical stimuli, and might affect subsequent responsiveness to other stimuli. We present experimental data showing that undiluted cologne can be aversive, but that aversion can be eliminated by dilution. We also show that the utility of cologne as a pungency control varies among brands, that cologne does not suppress responses to food chemicals in some species, but does in others, and that prior exposure to cologne does not affect later response to food chemicals. In 60 s swab trials with the Balearic lizard (Podarcis lilfordi), the main chemosensory responses were unaffected by cologne concentration. However, one-fourth of lizards exhibited slight to moderate aversion to undiluted cologne and 3:1 water:cologne, but not to a readily detected lower concentration (9:1). Two cologne brands did not affect responses, but a third brand induced increased tongue-flick rates. Colognes that stimulate increased tongue-flicking might mask real experimental effects. Cologne presented simultaneously with cricket chemicals did not affect tongue-flicking and biting responses in two species, but caused increased tongue-flicking in a third species and weaker response to cricket chemicals in a fourth. Prior testing with cologne did not affect responsiveness to cricket chemicals in a subsequent trial. Although cologne is a useful pungency control, pilot tests are needed to verify its utility for unstudied cologne types and animal species. Electronic Publication  相似文献   

6.
When important ecological factors change predictably during the life of an organism, the ontogeny of related behaviors must be timed to maintain appropriate behavioral responsiveness to current ecological conditions. In the brown iguana, Ctenosaura pectinata , hatchlings in natural populations eat primarily insects, consuming little plant matter, whereas adults eat primarily plants, consuming some insects as well. We conducted laboratory experiments on diet preferences and responses to chemical cues that the lizards sampled by tongue-flicking and used to identify food. All hatchlings ate crickets, but only one of six ate romaine lettuce. They responded strongly to chemical cues from prey, as indicated by elevated tongue-flick rates, but not from romaine lettuce. All older individuals ate both crickets and romaine lettuce. They responded much more strongly to chemical cues from both crickets and romaine lettuce than to control chemicals, as indicated by higher proportions of individuals that bit and higher tongue-flick attack scores.
Thus, an ontogenetic change to increased responsiveness to plant chemical stimuli was coordinated with an ontogenetic change to an herbivorous diet. The mechanisms underlying these ontogenetic changes are unknown, but folivory may be unprofitable before juveniles acquire intestinal flora that degrade cellulose by ingestion of feces of adult conspecifics. Possible mechanisms are discussed, including the detection of chemical cues from appropriate food plants during consumption of feces from older individuals. Studies of other squamate reptiles suggest that exposure to these chemicals might affect both future responsiveness to the chemical cues and a tendency to eat the corresponding plants.  相似文献   

7.
Use of chemical senses to detect prey is believed to be an important component of foraging behavior in actively foraging lizards. Ability to detect prey odors and discriminate them from control odors by tongue-flicking was studied in representatives of two families of lizards having highly forked, elongated, retractile tongues. Responses of gila monsters (Heloderma suspectum) and savannah monitors (Varanus exanthematicus) to deionized water, a control for pungency (cologne), and mouse odor on cotton swabs were studied in experiments using repeated-measures designs and employing the tongue-flick attack score (TFAS) as the primary measure of response strength. TFAS differed among treatments for gila monsters and monitors. Both species had greater TFAS to mouse odors than to either of the control stimuli, but responses to cologne were not statistically distinguishable from those to water. Numbers of tongue-flicks elicited by prey odors were greater than those for control stimuli in V. exanthematicus, but not for H. suspectum. Gila monsters, but not savannah monitors, bit in a significantly greater proportion of tests with prey odors than with control stimuli. Details of responses, including frequency of biting, apparent search behavior in the vicinity of applicators bearing mouse odors, and responses to control stimuli are discussed in relation to the foraging behavior of these two species and their relatives. Evidence from this and other studies suggests that the vomeronasal sense (and perhaps other chemical senses) is an important means of locating and recognizing prey in widely foraging autarchoglossan lizards.  相似文献   

8.
Specialist predators may respond strongly to sensory cues from preferred prey, but responses by generalist predators, although predicted to be less specific, are poorly known. Among squamate reptiles, diet and strength of response to chemical prey cues covary geographically in snakes that are specialist predators. There have been no previous studies of correspondence between diet and chemosensory response in lizards that are prey generalists. Actively foraging lizards discriminate between prey chemicals and control substances. It has been speculated that differential responses among prey species are unlikely in typical species that are dietary generalists. We examined this relationship in Podarcis lilfordi, an omnivorous lacertid that consumes a wide variety of animal prey. In experiments in which chemical stimuli were presented on cotton swabs, lizards responded more strongly to chemicals from a broad spectrum of prey types than to deionized water, an odorless control. These findings plus previous data showing that P. lilfordi is capable of prey chemical discrimination suggest that P. lilfordi can identify a wide range of potential prey using chemical cues. However, there was no evidence of differential response to stimuli among prey species, even in comparisons of prey included in the natural diet and potential prey not in the diet. The results, although limited to a single species, are consistent with the hypothesis that lizard species that are prey generalists do not exhibit the differential response strengths to chemical prey cues observed in snakes that have more specialized diets. Received in revised form: 17 July 2001 Electronic Publication  相似文献   

9.
Iguanid lizards are known for visual acuity and a diminished vomeronasal organ, which has led to mixed conclusions on whether iguanids use chemical cues. The collared lizard, Crotaphytus collaris, is a territorial iguanid that lives in open rocky habitats. Fecal pellets placed prominently on open rocky perches may provide an ideal mechanism for intraspecific chemical signaling. In order to determine whether collared lizards can discriminate between chemical stimuli found in conspecific fecal pellets, we collected 24 males and 25 females to analyze sex-specific behavioral responses via tongue-flicks and a newly observed behavior for the species, gular pumps, to cotton swabs containing water, cologne, chemical stimuli from conspecific male and female fecal pellets, and the lizard’s own fecal pellet. Both sexes were able to discriminate chemical stimuli from water via at least one behavior. Male collared lizards exhibited greater rates of response (tongue-flick and gular pumps) toward male fecal pellets when compared to the negative water control. Our results also suggest individuals may be able to discriminate between fecal pellets, as indicated by generally greater (but non-significant) counts of male tongue-flick responses to male fecal pellets when compared to their own. Collared lizard chemical discrimination appears to utilize tongue-flick and gular pump behaviors, possibly associated with distinct chemosensory modes (vomerolfaction and olfaction). Based on this study, we suggest that chemical signals may play a greater role in intraspecific communication than previously thought in this highly visual lizard.  相似文献   

10.
《Animal behaviour》1986,34(6):1805-1813
Behavioural cues were used to assay the capacity of common lizards to detect chemical deposits of snakes. The lizards were observed in cages that had been previously inhabited either by one of two species of snake that feed on lizards (the viper Vipera berus and the smooth snake Coronella austriaca), or the grass snake (Natrix natrix), which does not feed on lizards. As a control, the lizards were tested both in a clean cage and in one sprayed with a pungent odorant. The lizards responded to the snakes' chemicals by increased tongue-flick rates, with the highest rates being given in response to the deposits of their predators. The chemosensory examination of the snakes' odours induced a shift in general behaviour in response to the predator, but not to the non-predator chemical cues. This behavioural response consisted mainly of a disruption of the locomotor patterns. Our findings strongly suggest that lizards detected and distinguished between the chemicals deposited by three species of snake. Behavioural performances were highly variable among individual lizards in all trials, but the relative scores of individuals tended to be similar in response to different stimuli.  相似文献   

11.
A post-biting elevation in tongue-flicking rate was demonstrated experimentally in neonatal, ingestively naive garter snakes (Thamnophis radix). That the snakes also exhibited apparent searching movements suggests that strike-induced chemosensory searching occurs in nonvenomous snakes lacking previous experience with food or prey chemicals. Two litters of neonates differed in numbers of tongue-flicks emitted, but had similar relative magnitudes of response across experimental conditions. The existence of post-bite elevation in tongue-flick rate (and presumably strike-induced chemosensory searching) argues for a genetic basis for these chemosensory behaviors in a nonvenomous species of snake, extending the recent finding that strike-induced chemosensory searching is fully developed in ingestively naive neonatal rattlesnakes. Possible patterns of evolution of post-bite elevation in tongue-flick rate, and the strike-release-trail strategy of highly venomous snakes are discussed.  相似文献   

12.
Many lizards and all snakes flick their tongues. It is known that this unique behavioral pattern serves to collect airborne and substrate chemicals which give the animal information via Jacobson's Organ about the location of food, conspecifics, and possibly other environmental factors. However, a comparative topographic analysis of tongue movements in squamate reptiles is lacking, and it might shed light on the evolution of this behavior. In this study, a survey was made of the lizards and snakes which tongue-flick. Observations and films were made of 25 lizard species representing 10 families and 30 snake species representing 5 families. The information from observations and film analyses of representative species was used to hypothesize the steps of the evolution of tongue-flicking from the simple downward extensions of primitive lizards to the complex multiple oscillations of snakes.  相似文献   

13.
In lizards and snakes, foraging mode (active vs. ambush) is highly correlated with the ability to detect prey chemical cues, and the way in which such cues are utilized. Ambush-foraging lizards tend not to recognize prey scent, whereas active foragers do. Prey scent often elicits strikes in actively-foraging snakes, while ambushers use it to select profitable foraging sites. We tested the influence of foraging ecology on the evolution of squamate chemoreception by gauging the response of Burton's legless lizard ( Lialis burtonis Gray, Pygopodidae) to prey chemical cues. Lialis burtonis is the ecological equivalent of an ambush-foraging snake, feeding at infrequent intervals on relatively large prey, which are swallowed whole. Captive L. burtonis did not respond to prey odour in any manner: prey chemical cues did not elicit elevated tongue-flick rates or feeding strikes, nor were they utilized in the selection of ambush sites. Like other ambushing lizards, L. burtonis appears to be a visually oriented predator. In contrast, an active forager in the same family, the common scaly-foot ( Pygopus lepidopodus ), did tongue-flick in response to odours of its preferred prey. These results extend the correlation between lizard foraging mode and chemosensory abilities to a heretofore-unstudied family, the Pygopodidae.  相似文献   

14.
Luisa Amo 《Animal behaviour》2004,67(4):647-653
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.  相似文献   

15.
Corallus grenadensis is an arboreal boa endemic to the Grenada Bank. Thirty-five encounters with boas resulted in 17.65 hours of observations, including 6.3 hours of video-tape (which included two acts of predation). Boas under 100 cm are largely active foragers that move slowly through bushes and trees and tongue-flick leaf and branch surfaces apparently seeking chemosensory evidence of nocturnally quiescent lizard (Anolis) prey. Significantly more search time was directed to branches below the snake rather than to either the branches supporting the snake or to those above the snake, and tongue-flick rates were significantly higher for moving snakes than for those that were stationary. Smaller snakes prey on nocturnally quiescent lizards and they spent more time moving than did large snakes that feed on nocturnally active rodents and often employ an ambush foraging strategy. Once visual and, presumably, thermal information was received from a sleeping anole, C. grenadensis adopted a lengthy stalking process devoid of tongue-flicks. Snakes approached inactive lizards from adjacent branches with great stealth, moving at a rate of about 1 cm/min. The strike was made from close range (within 3 cm), and the prey was never released once contact was made. We conclude that, if chemosensory cues successfully lead a treeboa to a visual encounter with a sleeping lizard, subsequent behavior ensures a high rate of predation success.  相似文献   

16.
The chemical senses are crucial for squamates (lizards and snakes). The extent to which squamates utilize their chemosensory system, however, varies greatly among taxa and species’ foraging strategies, and played an influential role in squamate evolution. In lizards, ‘Scleroglossa’ evolved a state where species use chemical cues to search for food (active foragers), whereas ‘Iguania’ retained the use of vision to hunt prey (ambush foragers). However, such strict dichotomy is flawed as shifts in foraging modes have occurred in all clades. Here, we attempted to disentangle effects of foraging ecology from phylogenetic trait conservatism as leading cause of the disparity in chemosensory investment among squamates. To do so, we used species’ tongue‐flick rate (TFR) in the absence of ecological relevant chemical stimuli as a proxy for its fundamental level of chemosensory investigation, that is baseline TFR. Based on literature data of nearly 100 species and using phylogenetic comparative methods, we tested whether and how foraging mode and diet affect baseline TFR. Our results show that baseline TFR is higher in active than ambush foragers. Although baseline TFRs appear phylogenetically stable in some lizard taxa, that is a consequence of concordant stability of foraging mode: when foraging mode shifts within taxa, so does baseline TFR. Also, baseline TFR is a good predictor of prey chemical discriminatory ability, as we established a strong positive relationship between baseline TFR and TFR in response to prey. Baseline TFR is unrelated to diet. Essentially, foraging mode, not phylogenetic relatedness, drives convergent evolution of similar levels of squamate chemosensory investigation.  相似文献   

17.
Numerous studies on the feeding behavior of snakes have reported the consistency of tongue-flick responses with their natural diets. For representatives of widely distributed, dietary generalist species from particular localities, we can expect that their tongue-flick responses to potential prey unavailable in their original habitats have been reduced whereas those to prey common in the habitats have been enhanced. To test this hypothesis, intraspecific variation in tongue-flick responses to prey chemicals was examined using ingestively naive snakes (Elaphe quadrivirgata) from dietarily different populations: populations from the main Japanese island, where the snakes' diet predominantly consists of sympatric frogs, and from Mikura-jima Island, where no frogs occur and the snakes thus chiefly prey on lizards. We presented chemical stimuli from six items including those from their natural and potential prey (fish, frog, lizard, mouse, water, and cologne) to newborn snakes. Significant effects of stimuli on the tongue-flick responses were detected. On the other hand, effects of population and interaction between stimuli and population were not significant, and individual comparisons revealed no significant interlocality differences in responses to either frog or lizard chemicals. Thus, our hypothesis was not supported. However, in the Mikura-jima sample, significantly fewer snakes responded to frog chemicals than in the main island sample. The significance of the inconsistency between prey recognition ability and prey availability in the Mikura-jima population are discussed. Received: October 17, 2000 / Accepted: December 14, 2000  相似文献   

18.
Sensory abilities must allow efficient detection of prey, but the senses used and their relative importance may vary with hunting methods. In lizards, ambush foragers locate prey visually and active foragers use a combination of vision and vomerolfaction, the chemical sense associated with the vomeronasal system. Active foragers, but not ambush foragers, discriminate between prey chemicals and other chemical stimuli sampled by tongue-flicking. In active foragers, features of the tongue that may improve chemical sampling, such as elongation and forking are more pronounced and density of vomeronasal chemoreceptors is greater, than in ambush foragers. Foraging mode is fixed in most lizard families, and correlated evolution has been demonstrated among foraging mode, discrimination of prey chemicals, and lingual-vomeronasal morphology by interfamilial comparisons. Here I present information on a rare case of an intrageneric difference in foraging mode in the genus Mabuya . Laboratory experiments on the discrimination of prey chemicals showed that the active forager M . striata sparsa exhibits prey chemical discrimination, but the ambush forager M . acutilabris does not. The active forager also has a slightly more elongated tongue with deeper notching at the tip than the ambush forager, which might be a response to a change in foraging behavior or a reflection of unrelated differences in head shape. These findings confirm predictions based on correlated evolution between the hunting method and use of the chemical sense to locate food. They further show that chemosensory behavior is adjusted to change in foraging mode more rapidly than was previously known and suggest that behavioral changes may occur more rapidly than associated modifications of chemosensory morphology.  相似文献   

19.
The Japanese lacertid lizard Takydromus tachydromoides and the praying mantis Tenodera aridifolia are sympatric generalist predators feeding on similar prey. To confirm reciprocal predation between them, we observed the behavioural interactions between the lizards and the mantises of different sizes in a laboratory condition. The lizards caught small mantises (from first to fifth instars), but sometimes escaped from large mantises (from sixth instar to adult). Large mantises occasionally showed catch responses to the lizards. The lizards sometimes caught the mantis without a tongue-flick response (sampling of chemical cues), and they sometimes did not catch the small mantises showing immobile or cryptic responses that prevent visual detection. These results suggested the primary role of vision on recognition of the mantis as a prey. The lizards spent a longer time to approach larger mantises. The time from orienting to catch was longer when the lizards showed tongue-flick responses. The lizard also spent a longer time before deciding to escape from the mantis than to catch it. Biological significance of these differences in timing was discussed.  相似文献   

20.
The ability of broad-headed skinks (Eumeces laticeps) to distinguish familiar from unfamiliar individuals of the opposite sex and themselves from other individuals of the same sex using only chemical stimuli was examined experimentally. Cloacal chemical stimuli were presented to lizards on moistened cotton swabs and numbers of tongue-flicks performed in 60 s were recorded. Males emitted significantly greater numbers of tongue-flicks in response to chemical cues from unfamiliar females than of female cagemates. Response rates of males housed with females and males housed alone to chemical stimuli from unfamiliar females did not differ. Chemical stimuli from unfamiliar males elicited significantly more tongue flicks from males than their own or deionized water, but their own stimuli elicited no more than the odourless control. Females tongue-flicked significantly more in response to cloacal chemical stimuli from unfamiliar males than from male cagemates. The findings indicate that both sexes of broad-headed skinks can discriminate between chemical stimuli of familiar and unfamiliar individuals of the opposite sex and that males can distinguish their own cloacal chemicals from those of unfamiliar males. The possible functions of these chemosensory capacities, including location of mates by scent-trailing and assessment of the presence of sexual competitors, are discussed in relation to the social behaviour of E. laticeps.  相似文献   

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