Development of Antipredator Responses in Snakes

Individual and litter differences in the number of attacks directed by Mexican garter snakes (Thamnophis melanogaster) at a threatening stimulus were studied over the first year of life. Newborn snakes born to wildcaught females from the same population were tested the day after they were born and at 7, 16, 31, and 54 weeks of age on reactivity to a potent nonmoving and moving stimulus (a human finger). There were significant individual and interlitter differences in number of strikes directed at both stimuli but no significant effect of sex. Intercorrelations among the strike scores of individuals across test days averaged .54 for the nonmoving stimulus presentations and .63 for trials with the moving stimulus. Strike scores to the moving and nonmoving stimuli were significantly correlated at each age, however, attack responses to the moving stimulus declined over the year while those to the nonmoving stimulus remained constant. Measures of concordance showed that both individual strike scores and litter means were highly consistent, whereas consistency within litters was reduced. The results indicate that litter and individual differences among newborn snakes in the tendency to engage in antipredator behavior (“defensive temperament”) remain relatively stable from the first day after birth throughout the first year of life.     

孵化温度对王锦蛇胚胎代谢和幼体适合度的影响

孵化温度所驱动的爬行动物的表型变异是生理生态学研究的热点。本研究以王锦蛇(Elaphe carinata)为实验动物,检验了24℃和28℃孵化温度对王锦蛇胚胎代谢速率、孵化过程中的卵重量、孵出幼体代谢和行为的影响。研究结果显示:卵重和胚胎的呼吸代谢均与孵化时间呈正相关;28℃下胚胎代谢速率大于24℃;幼蛇孵出15 d内体重随着生长时间的延长而减小,24℃孵出幼体的代谢速率大于28℃孵出幼体,两温度下孵出幼体的呼吸代谢速率和生长时间无显著关系;28℃孵出幼体的疾游速和吐信频次均大于24℃;两孵化温度孵出幼体的选择体温无显著差异,但在消耗完体内的剩余卵黄后28℃孵出幼体有60%的个体摄食,而24℃孵出幼体无摄食个体。总体而言,王锦蛇28℃孵出幼体适合度优于24℃孵出幼体。     

Family Effects on Antipredator Behavior in the Field Cricket, Gryllus Integer

Most animals have antipredator mechanisms. These may include a variety of behaviors. One such behavior is “freezing”, i.e. becoming immobile in response to a predator. This paper is focused on freezing behavior in a species of field cricket, Gryllus integer. We studied this behavior in both nymphs and adults, and looked for family effects, as well as the effects of sex. Our research shows that there are no family effects in nymphs, but there are family effects in adults. No sex differences occur between the sexes at either stage of development. Because all of the crickets were reared under identical conditions in the laboratory, such family effects are likely due to genetic effects, maternal effects, or both.     

Inter‐individual Variation in Antipredator Hiding Behavior of Spanish Terrapins Depends on Sex,Size, and Coloration

Behavioral responses to predation risk are critical for survival but as antipredator behavior is costly, prey animals should flexibly modulate their optimum defensive responses by considering both costs and benefits, which are partly influenced by the individual characteristics of the prey. Turtles have the shell as a morphological structure that may provide partial protection against predators, but hiding into the shell may entail some high costs, and turtles should decide when to switch to an active escape strategy to safe refuges. Here, we examined how gender, body size, and sexual coloration influence inter‐individual variability of antipredatory hiding behavior into the shell of Spanish terrapins (Mauremys leprosa). We simulated predatory attacks under different conditions and measured the time that the turtles spent hidden entirely inside the shell (i.e., appearance times) and from then until the turtle started to flee actively (i.e., waiting times). Our results showed that when risk increased, appearance times increased but waiting times decreased. When turtles were in a prone position, their hiding behavior was related with their body weight with heavier turtles having longer appearance times. Also, the conspicuousness of limb coloration was important for the appearance times of males, but not for females. Thus, males with brighter coloration of the limb stripes had longer appearance times than duller ones. In addition, when turtles were overturned, males appeared out of the shell earlier than females and heavier turtles started to right sooner, but only when risk was low. However, when turtles were overturned and risk was high, they should assume that they have already been detected, making inter‐individual differences in size and coloration apparently unimportant for deciding hiding behavior.     

Blastobotrys serpentis sp. nov., isolated from the intestine of a Trinket snake (Elaphe sp., Colubridae)

Asporogenus yeast strains W113AT and W113B were isolated from the intestine of a dead Trinket snake. The two isolates showed 100% sequence similarity in the D1/D2 domain of the large-subunit (LSU) rRNA gene, internal transcribed spacer (ITS) 1-5.8S rRNA gene-ITS2 region and mitochondrial small-subunit rRNA gene and the cytochrome oxidase II gene sequence and also showed similar phenotypic characteristics. The nearest phylogenetic neighbors of W113AT and W113B based on the sequence of the D1/D2 domain of the LSU rRNA gene were Blastobotrys chiropterorum NRRL Y-17017T and Blastobotrys terrestris NRRL Y-17704T with about 98% similarity. The close affiliation of W113AT and W113B with B. chiropterorum NRRL Y-17017T and B. terrestris NRRL Y-17704T was also evident from the high similarity observed in the nucleotide sequences of the mitochondrial small subunit rRNA (96-97.8%) and the cytochrome oxidase II (95.5-95.6%) genes. In the neighbor-joining phylogenetic trees constructed based on the D1/D2 domain or cytochrome oxidase gene, the isolates clustered with the above-mentioned species. However, the isolates showed a number of differences in their phenotypic properties with B. chiropterorum NRRL Y-17017T and B. terrestris NRRL Y-17704T and hence are regarded as representing a novel member of the genus Blastobotrys, for which the name Blastobotrys serpentis sp. nov. is proposed.     

The Effects of Relaxed and Reversed Selection by Predators on the Antipredator Behavior of the Threespine Stickleback, Gasterosteus aculeatus

Isolation from predators can lead to the reduction or loss of ancestral behavioral defenses in prey, but does not always do so. Predators introduced to populations that have experienced relaxed selection from some ancestral predators can favor the evolution of antipredator behavior that has been lost. We examined these possibilities by eliciting antipredator behavior in three populations of threespine stickleback fish, Gasterosteus aculeatus : an oceanic population thought to resemble the ancestral form, and two populations historically (up to 20 000 yr) devoid of piscine predators (relaxed selection), one of which has been stocked with salmonids for nearly 25 yr (reversed selection). We used three kinds of predator models: a sculpin (ambush predator), a rainbow trout (chasing predator), and an overhead silhouette of an arctic tern. Stickleback reacted differently to the three models, indicating that they distinguished among them. Individuals from all populations responded similarly to the tern model. The ancestral population showed the weakest response to the sculpin model despite being the only population that encounters these predators naturally. Stickleback from the trout-free population displayed slightly reduced responses to the trout model, and recovery times like those in the ancestral population providing only weak evidence for loss of the ancestral antipredator repertoire. Fish from the reverse-selected population exhibited fascinating, elevated responses to both the trout and sculpin models relative to the other two populations. These findings offer initial evidence of (1) a partial alteration of the ancestral behavioral repertoire during a long period of relaxed selection from piscine predators, and (2) rapid acquisition of extreme responses to piscine predators under reverse selection.     

Contextual Flexibility: Reassessing the Effects of Prey Size and Status on Prey Restraint Behaviour of Macrostomate Snakes

Contextual flexibility in prey restraint behaviour has been documented in advanced snakes (Colubroidea), but the degree of flexibility for earlier snake lineages has been largely unstudied. We document the prey restraint behaviour of five snake species belonging to three early macrostomate lineages: Loxocemidae, Erycinae and Boidae. Species from these lineages were chosen for this study because they utilize similar prey resources but exhibit different ecological habits that may have important consequences on prey restraint behaviour. Snakes (n = 27) were studied in a systematic experimental design assessing the effects of mouse size (small and large) and status (live and dead) across a total of 216 feeding trials. Loxocemus and Erycine snakes were highly flexible in their prey restraint behaviour patterns and these varied across prey category. Individuals of Boa constrictor exhibited very little contextual flexibility in feeding behaviour, confirming earlier reports. Flexibility in prey restraint behaviour corresponded with loop application pattern, whether the snake bent laterally or ventrally when forming a loop around prey. Our study is the first to show that early macrostomate snakes exhibit flexible prey restraint behaviours. Thus, our results suggest that flexibility in predatory behaviour may be more widespread across snake taxa than previously thought and we offer hypotheses for the observed interspecific differences in snake feeding behaviour.     

Prey-handling Behaviour of Newly Hatched Snakes in Two Species of the Genus Elaphe with Comparison to Adult Behaviour

The prey-handling behaviour of two species of Asian ratsnakes was studied in the laboratory. In experiment 1, effects of prey size and type (mouse, lizard and frog) on capture position, direction of ingestion, condition of prey at ingestion, and prey-handling method were investigated using newly hatched Elaphe quadrivirgata. Prey size did not affect these variables except prey-handling method in frog trials: large frogs were more frequently constricted after a delay of more than 1 s than small frogs, and small frogs were usually simply seized. On the other hand, prey type affected prey-handling method: mice and lizards were more frequently constricted or pinioned than frogs. When hatchlings of E. quadrivirgata tried to coil around a prey animal immediately after striking, they frequently failed or released their coils within 10 s after striking. In experiment 2, prey-handling behaviour of adult E. climacophora was examined with mice of various sizes. Adult E. climacophora tended to constrict large mice immediately after striking, whereas they simply seized small mice. Only one out of 38 mice which were constricted immediately after striking was released from the coils within the subsequent 10 s. Large mice were killed prior to ingestion, whereas small mice were swallowed alive. Large mice tended to be swallowed head first. Experiment 3 was conducted to investigate effects of mouse size on prey-handling behaviour of newly hatched E. climacophora. Mouse size affected condition of prey at ingestion and prey-handling method but not capture position and direction of ingestion. Possible relationships between feeding ecology and the differences of prey-handling behaviour among Elaphe are discussed.     

Phylogeny of Courtship and Male-Male Combat Behavior in Snakes

Background

Behaviors involved in courtship and male-male combat have been recorded in a taxonomically broad sample (76 species in five families) of snakes in the clade Boidae + Colubroidea, but before now no one has attempted to find phylogenetic patterns in such behaviors. Here, we present a study of phylogenetic patterns in such behaviors in snakes.

Methodology/Principal Findings

From the literature on courtship and male-male combat in snakes we chose 33 behaviors to analyze. We plotted the 33 behaviors onto a phylogenetic tree to determine whether phylogenetic patterns were discernible. We found that phylogenetic patterns are discernible for some behaviors but not for others. For behaviors with discernible phylogenetic patterns, we used the fossil record to determine minimum ages for the addition of each behavior to the courtship and combat behavioral repertoire of each snake clade.

Conclusions/Significance

The phylogenetic patterns of behavior reveal that male-male combat in the Late Cretaceous common ancestors of Boidae and Colubridae involved combatants raising the head and neck and attempting to topple each other. Poking with spurs was added in Boidae. In Lampropeltini the toppling behavior was replaced by coiling without neck-raising, and body-bridging was added. Phylogenetic patterns reveal that courtship ancestrally involved rubbing with spurs in Boidae. In Colubroidea, courtship ancestrally involved chin-rubbing and head- or body-jerking. Various colubroid clades subsequently added other behaviors, e.g. moving undulations in Natricinae and Lampropeltini, coital neck biting in the Eurasian ratsnake clade, and tail quivering in Pantherophis. The appearance of each group in the fossil record provides a minimum age of the addition of each behavior to combat and courtship repertoires. Although many gaps in the story of the evolution of courtship and combat in snakes remain, this study is an important first step in the reconstruction of the evolution of these behaviors in snakes.     

The Influence of Hunting on Antipredator Behavior in Central African Monkeys and Duikers

Many animals can adjust their behavioral strategies to reduce predation risk. We investigated whether rain forest monkeys and duikers alter their antipredatory behavior in response to hunting by humans in southwestern Gabon. We compared monkey and duiker responses to human observers in an area where hunting is prohibited, to those in a nearby area where hunting pressure is moderate but spatially variable. The results of our study indicate that monkeys become more secretive when hunted, commencing alarm calls only when at a certain distance (typically > 50 m) from humans. We found no difference in monkey group size between hunted and no-hunting areas. In no-hunting areas, duikers often freeze in response to approaching observers, but in hunted areas they abandon this strategy and rapidly flee from humans. Duikers also whistle more often in areas where they are hunted frequently. Our findings have at least two important implications. First, behavioral observations of monkeys and duikers may be useful in gauging local hunting intensity in African rain forests. Second, duiker densities are likely to be overestimated in hunted areas, where they more readily flee and whistle, and underestimated in no-hunting areas, where they rely on freezing behavior to avoid detection. Because behavioral adaptations to hunting vary both among species and localities, these differences should be considered when attempting to derive population-density estimates for forest wildlife.     

Antipredator Behavior of Brown Howlers Attacked by Black Hawk-eagle in Southern Brazil

In field studies between 2002 and 2004 in southern Brazil, we recorded antipredatory behaviors (N=9) of brown howlers (Alouatta guariba clamitans) under attack by the black hawk-eagle (Spizaetus tyrannus). We observed 2 types of behavioral responses: the descent to the understory followed by the dispersion of the group and immobility and silence. Brown howlers displayed behaviors specific to evading aerial predators, which are similar to the ones in other Neotropical primates.The article is contribution number 1578 of Departamento de Zoologia, Universidade Federal do Paraná     

摄食水平对瓦氏黄颡鱼餐后代谢特征的影响

以新鲜泥鳅(Misgurus anguillicanndotus)肉块为饵料,采用麻醉灌喂的方法,在(25.0±0.5)℃条件下,研究了瓦氏黄颡鱼(Pelteobagrus vachelli)幼鱼[体重(7.03±0.15)g,n=42]不同摄食水平(饵料分别为体重的0%、1%、2%、4%和8%)对餐后代谢反应的影响.结果显示,在不同摄食水平下,瓦氏黄颗鱼摄食代谢均呈现先上升后下降的整体变化趋势;摄食水平由1%增加到8%,实验鱼的SDA耗能、SDA时间和摄食代谢峰值(PMR)分别从3.09 kJ/kg、8 h和56.08 mg O_2/kg·h增加到47.21 kJ/kg、36 h和97.25 mgO_2/kg·h;其中,8%(饱足摄食水平)和4%摄食水平组的代谢率在峰值水平能够持续20 h左右.瓦氏黄颡鱼最大运动代谢率(MMR)为166.5 mg O_2/kg·h,显著大于饱食组PMR(P＜0.05).本研究还表明,随摄食水平上升,瓦氏黄颡鱼通过PMR的增加和SDA时间的延长来满足SDA耗能增加的需求;从摄食4%组和8%组相似的梯形摄食代谢曲线,可以看出瓦氏黄颡鱼在高摄食水平条件(＞4%)下限制了摄食代谢的上升;PMR相对于静止代谢上升倍率较小,暗示其摄食占据的代谢空间较小,进而保留了大量剩余代谢空间用于运动和其他生理活动,这可能与其活跃的觅食活动习性有关.     

Effects of Doubling the Portion Size of Fruit and Vegetable Side Dishes on Children's Intake at a Meal

Increasing the portion size of energy‐dense entrées has been shown to increase children's energy intake during a meal. It remains to be investigated whether serving larger portions to children can be used to promote intake of more healthful foods, such as fruits and vegetables (F&V). The aim of the present study was to examine the effects of increasing the portion size of F&V side dishes on children's intake. Forty‐three children (22 boys, 21 girls), aged 5–6 years, were served dinner once a week for 2 weeks. Each dinner consisted of pasta with tomato sauce, three F&V side dishes (broccoli, carrots, and applesauce), and milk. The portion size of the F&V was doubled between experimental conditions whereas the size of the pasta remained constant. Doubling the portion size of the side dishes resulted in a 43% increase in children's intake of the fruit side dish (P = 0.001), but did not affect children's intake of the two vegetable side dishes (P > 0.60). Further, when the portion size of F&V side dishes was doubled, children ate significantly less of the pasta (P = 0.04). The difference in meal energy intake between portion size conditions (19.5 ± 16.3 kcal) was not significant (P = 0.24). Although more studies are needed to understand whether increases in portion size can influence vegetable intake, children did eat more in response to a large quantity of a preferred low energy‐dense fruit side dish at meals. Thus variations in portion size can be used strategically to help children achieve the recommended intake of fruits.     

群体规模对越冬灰鹤警戒行为的影响

2005年1—3月,在云南丽江拉市海就群体规模对越冬灰鹤(Grusgrus)警戒行为的影响进行了研究。用扫描取样记录群体的规模和警戒个体的数量、用焦点取样记录群体中个体警戒行为的频次和持续时间,结果显示灰鹤群体和个体的警戒力均随群体规模增加而降低,但集群个体数超过30只后,群体警戒力便不会再下降(P>0.05)、成体的警戒持续时间也会增加(P<0.01);当群体规模超过50只后,成体的警戒频次也会上升(P<0.05)。推测亚成体维持低警戒的群体规模上限值要高于成体,单从警戒行为分析,20—30只个体的集群可能代表越冬灰鹤的最适群体大小。     

Antipredator Behavior of American Bullfrogs (Lithobates catesbeianus) in a Novel Environment

Invasive species capable of recognizing potential predators may have increased establishment rates in novel environments. Individuals may retain historical predator recognition and invoke innate responses in the presence of taxonomically or ecologically similar predators, generalize antipredator responses, or learn to avoid risky species in novel environments. Invasive amphibians in aquatic environments often use chemical cues to assess predation risk and learn to avoid novel predators via direct experience and/or associated chemical cues. Ontogeny may also influence recognition; experience with predators may need to occur at certain developmental stages for individuals to respond correctly. We tested predator recognition in invasive American bullfrog ( Lithobates catesbeianus) tadpoles that varied in experience with fish predators at the population and individual scale. We found that bullfrog tadpoles responded to a historical predator, largemouth bass ( Micropterus salmoides), only if the population was locally sympatric with largemouth bass. Individuals from a population that did not co‐occur with largemouth bass did not increase refuge use in response to either largemouth bass chemical cues alone or chemical cues with diet cues (largemouth bass fed bullfrog tadpoles). To test whether this behavioral response was generalized across fish predators, we exposed tadpoles to rainbow trout ( Oncorhynchus mykiss) and found that tadpoles could not recognize this novel predator regardless of co‐occurrence with other fish species. These results suggest that environment may be more important for predator recognition than evolutionary history for this invasive species, and individuals do not retain predator recognition or generalize across fish predators.     

Correlations between Ontogenetic Change in Color Pattern and Antipredator Behavior in the Racer, Coluber constrictor

Differences in snake color pattern have been demonstrated to affect behaviors involved in antipredator defense. Snakes with blotched or banded color patterns are concealed when not moving, and tend to rely on concealment and aggression for defense. In contrast, snakes with uniform or striped color patterns are easily seen when stationary, but their speed and direction are difficult to track when moving. They tend to rely on flight for protection. Some snake taxa exhibit ontogenetic change in color pattern, but the behavioral consequences of this change have not been investigated. I present results of a behavioral study in the racer, Coluber constrictor, which has a blotched juvenile color pattern but is uniformly colored as an adult. Hatchling racers were significantly more likely than adults to show aggressive behavior when confronted with a model predator, whereas adults were more likely to flee. This supports the hypothesis that changes in behavior and color pattern are correlated in this species to provide effective antipredator defense at different stages of life history. I also examined sprint speed, which may be an important factor in antipredator defense. Juvenile and adult racers showed a similar relationship between length and speed, a pattern also seen in other species that lack color change. This result suggests that sprint speed is not a causal factor in the evolution of ontogenetic color change.