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Biomedical subjects

William E Cooper

Publications and source records attributed to William E Cooper.

7 recordsLinked to original sources

Optimal flight initiation distance.

Decisions regarding flight initiation distance have received scant theoretical attention. A graphical model by Ydenberg and Dill (1986. The economics of fleeing from predators. Adv. Stud. Behav. 16, 229-249) that has guided research for the past 20 years specifies when escape begins. In the model, a prey detects a predator, monitors its approach until costs of escape and of remaining are equal, and then flees. The distance between predator and prey when escape is initiated (approach distance = flight initiation distance) occurs where decreasing cost of remaining and increasing cost of fleeing intersect. We argue that prey fleeing as predicted cannot maximize fitness because the best prey can do is break even during an encounter. We develop two optimality models, one applying when all expected future contribution to fitness (residual reproductive value) is lost if the prey dies, the other when any fitness gained (increase in expected RRV) during the encounter is retained after death. Both models predict optimal flight initiation distance from initial expected fitness, benefits obtainable during encounters, costs of escaping, and probability of being killed. Predictions match extensively verified predictions of Ydenberg and Dill's (1986) model. Our main conclusion is that optimality models are preferable to break-even models because they permit fitness maximization, offer many new testable predictions, and allow assessment of prey decisions in many naturally occurring situations through modification of benefit, escape cost, and risk functions.

Animals↗

History and the global ecology of squamate reptiles.

The structure of communities may be largely a result of evolutionary changes that occurred many millions of years ago. We explore the historical ecology of squamates (lizards and snakes), identify historically derived differences among clades, and examine how this history has affected present-day squamate assemblages globally. A dietary shift occurred in the evolutionary history of squamates. Iguanian diets contain large proportions of ants, other hymenopterans, and beetles, whereas these are minor prey in scleroglossan lizards. A preponderance of termites, grasshoppers, spiders, and insect larvae in their diets suggests that scleroglossan lizards harvest higher energy prey or avoid prey containing noxious chemicals. The success of this dietary shift is suggested by dominance of scleroglossans in lizard assemblages throughout the world. One scleroglossan clade, Autarchoglossa, combined an advanced vomeronasal chemosensory system with jaw prehension and increased activity levels. We suggest these traits provided them a competitive advantage during the day in terrestrial habitats. Iguanians and gekkotans shifted to elevated microhabitats historically, and gekkotans shifted activity to nighttime. These historically derived niche differences are apparent in extant lizard assemblages and account for some observed structure. These patterns occur in a variety of habitats at both regional and local levels throughout the world.

Adaptation, Biological↗

Foraging mode and evolution of strike-induced chemosensory searching in lizards.

Strike-induced chemosensory searching (SICS) in lizards and snakes is a means of relocating prey by scent-trailing. The two main components of SICS are an elevated tongue-flick rate for vomerolfactory sampling after biting prey (PETF) and searching movements. In combination, these behaviors permit scent-trailing. Prey chemical discrimination, which is a prerequisite for SICS, is present in active foragers, but not in ambush foragers. Using comparative data. I show that searching movements and SICS have undergone correlated evolution with foraging mode and with prey chemical discrimination in lizards. This suggests that active foraging selects for prey chemical discrimination, which is then employed to search for escaped prey using the typical movements and tongue-flicking behaviors of active foragers. SICS in lizards is simply heightened active foraging after biting prey. In nonvenomous snakes, SICS is similar to that in lizards but is not restricted to active foragers. Only highly venomous snakes voluntarily release dangerous prey upon envenomation, pause to let the venom incapacitate the prey, and then relocate the prey by scent-trailing. PETF was observed in two ambush foragers and is not evolutionarily correlated with foraging mode or searching movements. Because it occurs in species lacking prey chemical discrimination, such PETF may be a response to gustatory cues or to internal chemicals not encountered on surfaces or trails of uninjured prey.

Adaptation, Physiological↗

Behavioral responses to plant toxins by two omnivorous lizard species.

An ability to detect plant toxins and thereby avoid eating chemically defended plants would be very beneficial for omnivorous and herbivorous lizards. We studied the ability of the omnivorous Podarcis lilfordi to detect compounds belonging to three classes of common plant toxins, as well as responses indicating aversion. Solutions of the alkaloid quinine, saponin, and the phenolic coumarin, as well as distilled water (odorless control), were presented to lizards on cotton swabs. The lizards detected all three toxins as indicated by significantly decreased tongue-flick rates and tongue-flick attack scores in comparison with distilled water. Several other variables revealed aversion to saponin, including a low number of individuals that bit swabs, avoidance of swabs after tongue-flicking, performance after tongue-flicking the swab of repeated short-excursion tongue-flicks that were directed away from the swab and did not contact any substrate, failure to respond at all in the next trial, and wiping the snout on the floor of the terrarium. Reasons for apparent differences in tongue-flicking behavior between P. lilfordi and two other omnivorous lizard species are discussed. We also showed experimentally that saponin depresses the tongue-flick rate in the omnivorous Bonaire whiptail lizard, Cnemidophorus murinus. Tongue-flicking enables at least one lizard species to detect specific chemicals representing three major classes of plant toxins. It is hypothesized that this ability is a widespread adaptation to reduce ingestion of plant toxins.

Animals↗

Pheromonal discriminations of sex, reproductive condition, and species by the lacertid lizard Podarcis hispanica.

In some vertebrate taxa, pheromones provide important information about species, sex, reproductive condition, kinship, and even individual identity. Because they possess highly developed nasal chemosensory systems, lizards are capable of many chemical discriminations, but many aspects of their pheromonal communication remain poorly understood even in major families. We report that males of a lacertid lizard, Podarcis muralis, are capable of differential response to surface chemical cues from conspecific males and females, from gravid and nongravid females, and from conspecific females and females of the closely related sympatric congener, P. bocagei carbonelli. In 60 sec trials in which stimuli from the femoral, cloacal, lateral, and upper body surfaces were presented to males on cotton swabs, males tongue-flicked at significantly higher rates to stimuli from conspecific females than males, from conspecific nongravid than gravid females, and from conspecific than heterospecific females. Responses to stimuli from conspecific males did not differ from those to distilled water. Together with previous findings that males can distinguish between chemical cues from familiar and unfamiliar males, these findings suggest that pheromones provide male P. hispanica important information regarding the presence of sexual rivals and the reproductive condition of potential mates without visually encountering other lizards. A growing body of literature indicates that lacertids are capable of sophisticated pheromonal discriminations that may play important roles in their social behavior.

Animals↗

Responses to major categories of food chemicals by the lizard Podarcis lilfordi.

Many lizards are capable of identifying food using only chemical cues from food surfaces, but almost nothing is known about the types of compounds that are effective stimuli. We experimentally studied lingual and biting responses by a lacertid lizard, Podarcis lilfordi, to single representatives of three major categories of food chemicals, sucrose as a carbohydrate, pure pork fat as a mixture of lipids, and bovine gamma globulin as a protein. In 60-sec trials in which stimuli were presented on cotton swabs, the lizards detected all three stimuli, exhibiting more tongue-flicks, licks, or bites, or a greater tongue-flick attack score (TFAS; overall measure of response strength to prey stimuli) than to deionized water. The initial response to all stimuli was tongue-flicking, but the lizards discriminated among the types of chemical stimuli. After preliminary tongue-flicks, the lizards responded to sucrose solutions by licking at high rates, to pure pork fat by biting, and to protein by a combination of additional tongue-flicks and biting. Biting is a feeding response to prey or solid plant material. Licking is a feeding response to sugars in nectar or ripe fruit. Its frequency increased with sucrose concentration. Our data suggest that lizards can identify several types of chemicals associated with food and direct feeding attempts to sources of such chemicals in the absence of visual cues.

Analysis of Variance↗

Lingual and biting responses to selected lipids by the lizard Podarcis lilfordi.

Many lizards can identify food using chemical cues, but very little is known about the chemical constituents used for this purpose. We experimentally investigated responses to several lipid stimuli by the omnivorous lacertid lizard Podarcis lilfordi, which had been shown previously to be capable of identifying prey using only chemical cues and to respond to pork fat by tongue-flicking and biting. In 60-s trials in which stimuli were presented on cotton swabs, the lizards responded very strongly to pure pork fat and to oleic acid, but not to cholesterol or glycerol. Latency to bite swabs, the number of individuals that bit swabs, and the tongue-flick attack score, TFAS(R), which combines effects of tongue-flicks and bites, showed stronger responses to fat than to cholesterol, glycerol, and distilled water but did not differ significantly from responses to oleic acid. Several lines of evidence show that oleic acid elicited strong chemosensory and feeding responses. For individuals that did not bite, the number of tongue-flicks was significantly greater for oleic acid than for distilled water or glycerol, and nearly so for cholesterol. Latency to bite was significantly shorter for oleic acid than for distilled water, and TFAS(R) was significantly greater for oleic acid than for distilled water and glycerol. In combination with pilot data indicating no strong response to the waxy, saturated palmitic acid, these findings suggest that oleic acid in particular and probably other unsaturated fatty acids found in animal fat contribute strongly to the food-related responses to lipids.

Animals↗