Juvenile Collared Lizards Adjust Tail Display Frequency in Response to Variable Predatory Threat

Size: px
Start display at page:

Download "Juvenile Collared Lizards Adjust Tail Display Frequency in Response to Variable Predatory Threat"

Transcription

1 Ethology Juvenile Collared Lizards Adjust Tail Display Frequency in Response to Variable Predatory Threat Joshua R. York* & Troy A. Baird * Department of Biology, University of Oklahoma, Norman, OK, USA Department of Biology, University of Central Oklahoma, Edmond, OK, USA Correspondence Joshua R. York, Department of Biology, University of Oklahoma, 730 Van Vleet Oval, Norman, OK 73019, USA. joshuayork@ou.edu Received: May 15, 2015 Initial acceptance: June 6, 2015 Final acceptance: October 12, 2015 (L. Ebensperger) doi: /eth Keywords: antipredator behavior, antipredator display, lizard tail displays, predator avoidance behavior, predator deflection, predator deterrence Abstract Antipredator behavioral tactics have evolved in phylogenetically diverse animal clades and often involve prey initiating conspicuous display patterns when encountering potential predators. Using detailed behavioral observations in the field, we report the first description of conspicuous tail displays in juvenile collared lizards (Crotaphytus collaris), a species that does not have tail autotomy. When approached in the field, lizards gave four stereotypical displays involving the tail, suggesting that these might function as antipredator signals. To test this hypothesis, we compared the frequency of tail displays given by juveniles during approach experiments relative to control trials involving observation from afar. We then further manipulated the intensity (low threat, high threat) with which we threatened lizards by varying our angle of approach relative to the body-axis orientation of lizards resting on a perch sloping away from the observer. Our results show that juveniles consistently performed all four types of tail displays in response to our approaches, but subjects never displayed during non-threat control trials. Moreover, lizards were more likely to remain emergent and give tail displays when we approached them head-on with their bodies sloping away (low threat), but were more likely to take refuge when approached from the side and behind (high threat). The positive relationship between the frequency of display and risk level suggests that tail displays function to signal that juvenile lizards have detected potential predators, which may deter further pursuit. Together, our results provide the first account of visual displays involving the tail in collared lizards and suggest that these displays function to signal potential predators. Introduction A wide variety of behavioral traits are shaped by natural selection for predator avoidance, including those involved in escaping or minimizing attack (Greene 1988; Caro 2005; Stankowich & Blumstein 2005). Prey sometimes simply flee and/or take refuge when they encounter potential predators (Broom & Ruxton 2005). Although fleeing and hiding may decrease the likelihood of predation, these responses also may compromise the ability of prey to engage in other activities that promote fitness, such as remaining motionless for sit-and-wait foraging (Martın & Lopez 1999; Amo et al. 2007), as well as opportunities to mate or thermoregulate. The apparent selective tradeoff between predator avoidance and activities that require exposure to attack may promote the evolution of antipredator behavioral tactics that balance vulnerability against other essential demands of fitness (Ydenberg & Dill 1986; Ruxton et al. 2004). Such tactics have evolved independently in phylogenetically diverse animal clades, and often involve prey directing displays to approaching predators even though they appear to render prey more conspicuous and apparently more vulnerable (Cooper et al. 2004; Langkilde et al. 2004). For example, several species of Ethology 122 (2016) Blackwell Verlag GmbH 37

2 Antipredator Tail Displays in a Lizard J. R. York & T. A. Baird fish, mammals, and insects are known to advertise colorful/ornamented body parts to potential predators, which may deter further predatory pursuit (Caro 2005; Stankowich & Blumstein 2005). Similarly, several birds and amphibians produce acoustic signals in response to predatory threat to alert predators that they have been detected (Caro 2005; Stankowich & Blumstein 2005). Lizards have emerged as excellent models for studying the adaptive significance of antipredator behavioral signals. Many lizard species give highly conspicuous displays, most often involving the tail, when threatened by predators in both natural habitats and laboratory settings (Greene 1988; Hasson et al. 1989; Telemeco et al. 2011). Three primary mechanisms have been hypothesized to explain the evolution of conspicuous, apparently risk-prone tail displays that are given by lizards. The simplest explanation is that tail displays evolved to deflect attacks to a non-essential body part (Bateman & Fleming 2009; Cooper & Vitt 2010). Tail displays are particularly likely to evolve through deflection in lizards having highly ornamented tails that autotomize when grasped affording prey sufficient time to escape while the predator consumes the tail (Cooper 1998; Telemeco et al. 2011). Alternatively, especially in lizards that lack tail autotomy, tail displays may communicate more detailed content that functions to deter pursuit by predators (Hasson et al. 1989; Cooper et al. 2004). For example, prey may signal that they have detected approaching predators (perception advertisement hypothesis, sensu Caro 1995), causing predators to abandon further pursuit. Tail displays may also signal that prey are capable of evading an impending attack because nearby refuges are abundant, or from their current position relative to the predator, prey are physically capable of escape by fleeing (relative escape ability hypothesis, sensu Caro 1995). During routine mark recapture experiments, we observed that juvenile collared lizards (Crotaphytus collaris) sometimes moved their tails in a stereotypical manner when we approached them, suggesting that these displays may function as antipredator signals. Because these displays have not been previously reported in this species, we first give a detailed description of the display patterns involving the tail. We then experimentally tested the hypothesis that these stereotypical tail displays function as antipredator signals by (1) approaching free-ranging juveniles to simulate a predatory threat (Cooper 2003) and recording tail display activity and (2) manipulating the intensity with which we threatened lizards by varying the angle of approach. Materials and Methods Study Site and Population Collared lizards are diurnal medium-sized Iguanians that occur throughout the central and southwestern USA. Natural habitat for collared lizards consists of exposed rock outcroppings and washes, but they have also colonized human constructed habitats composed of hard substrates such as flood control spillways associated with dams (Baird 2013a,b; Husak 2006a, b). Collared lizards use elevated rock perches to bask and scan for arthropod prey, and they take refuge from potential predators and excessive heat in crevices beneath or adjacent to these perches (Baird & Sloan 2003; Baird et al. 2003; Baird 2013a). This study was conducted at the Arcadia Lake Dam located 9.6 km east of Edmond, OK, USA, which is well suited for observation of collared lizard behavior because human access is restricted and the lizards are undisturbed (Baird 2013a). We conducted experiments on two habitat patches ( m and m) composed of contiguous pieces of broken concrete slab. These patches are optimal for observation of lizard responses to manipulations of simulated predation threat because they are narrow, unobstructed, and bordered on both sides by grass. Because the broken pieces of concrete ( m) rest at variable angles, they provided abundant slanted perches and allowed us to approach lizards from carefully controlled angles which varied the degree to which lizards were exposed to the observer. Grounddwelling predators of collared lizards at our study site are coachwhips (Masticophis flagellum), roadrunners (Geococcyx californianus), and coyotes (Canis latrans), whereas raptors (Ictinia mississippiensis, Buteo jamaicensis) attack lizards from above. Experimental Subjects and Protocol From mid-july through late October 2011 and 2012, we surveyed the entire study site daily to capture (by noose), mark, and measure newly emerged hatchlings, and then recapture when they molted. At each capture, we recorded snout-to-vent length (1 mm) and total body mass (0.1 g). Lizards were marked permanently using unique toe-clips and identified from a distance by applying non-toxic paint spots to the dorsum. Newly emerged hatchlings in mid-july and early August were mm SVL and had approximately doubled in size ( mm SVL) by the end of October. For our experiments, we used 38 Ethology 122 (2016) Blackwell Verlag GmbH

3 J. R. York & T. A. Baird Antipredator Tail Displays in a Lizard juveniles that were approximately mm SVL. We only used juveniles that had full tails, avoiding those having part of their tails bitten off by predators (collared lizards lack tail autotomy septa). All experiments were conducted from 0900 to 1300 h when the substrate temperature was C. Over the 8 C temperature range that we ran experiments, collared lizard activity, including escape behavior, is not dependent on surface temperature (Baird et al. 2001; Braun et al. 2010). To control for possible effects of lizard habituation to observers, all subjects were previously captured, marked, and measured no more than twice prior to experiments. For all trials, we first identified subjects perched on elevated rocks from a distance m using their unique color codes. The observer (JRY) approached subjects holding a noose pole, using one of three techniques to vary threat level categorically (non-threat, low threat, high threat; described below). For all treatments, we recorded the number of seconds that lizards spent giving tail displays for 10 min, and calculated the percent time per trial that lizards displayed using their tails. To test the hypothesis that tail displays function as antipredator signals, we first compared all tail activity by subjects (n = 12) during non-threat versus lowthreat conditions. The non-threat treatment involved the observer sighting lizards from afar and, once the observer had approached within 10 m of the subject, stopped and observed subjects without further approach (Fig. 1). For the low-threat treatment, we chose subjects that were resting on an elevated rock perch that sloped away from the observer, such that the lizard s median axis was oriented parallel with the observer s line of approach (Fig. 1), and only the head (and elevated tail) was exposed to the simulated predator. Subjects were approached using a constant pace (1.5 m/s; Cooper 2003). When subjects first moved, the observer paused and recorded whether lizards immediately took refuge in an adjacent crevice without giving tail displays, or responded by displaying using the tail while remaining emergent. If lizards gave tail displays, we characterized them (see Characterization of tail displays) and recorded the duration of each (1.0 s). When lizards remained emergent in response to approach and ceased displaying, the observer resumed approach until subjects either displayed again, or took refuge in a crevice and did not re-emerge. Each subject lizard was used in both the non- and low-threat treatments in random order. Trials on the same subjects were conducted no sooner than 1 d following their previous trial. Fig. 1: Experimental protocol for non-threat (open arrow), low-threat (hatched arrow), and high-threat (solid arrows) trials. All trials began with the observer 10 m away from a subject perched on an elevated rock. For both non-threat and low-threat trials, the observer was facing the lizard s head, parallel with the subject s midline. High-threat trials began behind and to the side of subjects degrees relative to the direction of the subject lizard s head (solid arrows). We further tested our hypothesis that tail displays by juvenile collared lizards function as antipredator signals by pooling the initial 12 low-threat trials with an additional 43 low-threat trials (total low-threat trials, n = 55 using the same protocol) for comparison with trials involving high threat (n = 38). High-threat trials involved the observer approaching perched lizards degrees to either side of the direction that the subject s head was facing (Fig. 1). Following Cooper (2008), we reasoned that these angled approaches from behind and laterally posed an increased threat because the entire body of the lizard was exposed to the simulated predator relative to only the head being exposed during the low-threat treatment, and the ability of lizards to visually evaluate the speed and distance of the predator when approached from behind and laterally would be compromised. Although the initial 12 low-threat trials were conducted before high-threat trials, over the entire sample of low- and high-threat trials, the order of treatments was random. Statistical Analyses For all analyses, we used generalized linear mixed models (GLMM; package lme4) in the program R v (R Development Core Team 2013). We report effect sizes for our categorical (Cohen s d) predictor variables, and considered effects Ethology 122 (2016) Blackwell Verlag GmbH 39

4 Antipredator Tail Displays in a Lizard J. R. York & T. A. Baird statistically significant when the associated 95% confidence intervals (CI) did not include zero (Colegrave & Ruxton 2003; Nakagawa & Cuthill 2007). We interpreted the relative strength of effect sizes from resulting models according to Cohen (1992) (small effect = below 0.2; moderate effect = ; large effect =>0.50). To test whether tail displays in juvenile collared lizards function as antipredator signals, we used threat level (non-threat versus low threat) during experiments as the categorical predictor variable in a paireddesign GLMM (Gaussian error structure, identity link function), with lizard identity as the random effect to control for multiple observations on some individuals. We used a GLMM (Gaussian error structure, identity link function) with threat level (low threat versus high threat) as the categorical predictor variable to test whether or not variation in the intensity of threat influenced the cumulative percent time per trial that juveniles gave all types of tail displays, and included lizard identity as a random effect. Finally, we used a GLMM (binomial error structure, log link function) to compare the probability that lizards stayed emergent and gave tail displays versus seeking refuge without displaying in low-threat versus high-threat trials. For this model, we used threat level (high threat, low threat) as the categorical predictor variable and lizard response (remain emergent and display or seek refuge immediately without displaying) as the dependent variable, with lizard identity as a random effect. We limited our analyses to the cumulative percent time displaying for all types of displays pooled, because there was no obvious pattern with which lizards gave the four display types in response to different threat treatment levels. and Use Committee at the University of Central Oklahoma (permit number 13009) and the Oklahoma Department of Wildlife (permit number 5553). We have conducted 25 consecutive seasons of longitudinal studies on collared lizard behavior, growth, and survival in this population. These have involved clipping the terminal phalanges of three digits, application of non-toxic acrylic paint to the dorsum, repetitive capture by noosing, and making body size measurements. Monitoring the behavior and survival of lizards has confirmed that these techniques have had no adverse effects on their health (Baird 2013a; York et al. 2014; York & Baird 2015). Results Characterization of Tail Displays Juvenile collared lizards performed four variations of displays using their tails. The most common display involved conspicuous elevation of the straightened tail above the substrate and moving it back and forth laterally (=lateral wave, display just beginning in Fig. 2a, lateral movement indicated by the arrows). Lizards also frequently displayed by elevating the tail and moving it using a stereotypical slow and deliberate sinusoidal motion (=sinusoidal wave, Fig. 2b). Less frequently, lizards raised and held the tail stiffly in two different positions. Tail curl (shown just beginning in Fig. 2c) involved slowly raising and curling the distal portion of the tail until it was directly above the head and pointed anteriorly. Tail raise involved rapidly elevating the tail and holding it stiffly above the substrate (same as in Fig. 2a, but without lateral movement). Ethical Note All procedures performed on live lizards were conducted with approval of the Institutional Animal Care Tail Display under Different Threat Levels Juvenile collared lizards did not initiate tail displays during non-threat control trials (n = 12), but they (a) (b) (c) Fig. 2: Tail displays given by juvenile collared lizards. (a) lateral wave (lateral motion of the tail indicated by arrows) or tail raise while it is held straight and stiff (without arrows); (b) sinusoidal wave; (c) tail curl just beginning in the photograph, but from here will ultimately be raised over the head. Photography by Teresa D. Baird. 40 Ethology 122 (2016) Blackwell Verlag GmbH

5 J. R. York & T. A. Baird Antipredator Tail Displays in a Lizard gave displays frequently (all four displays pooled) during 10 of 12 (83%) low-threat trials (t 2, 22 = 5.43, Cohen s d = 2.22, 95% CI = , Fig. 3). The average percent time spent giving tail displays during low-threat trials was 3.9 times higher (t 2, 32 = 3.17, Cohen s d, 95% CI = 0.63, ) than that during high-threat trials (Fig. 4). Moreover, in highthreat trials lizards were over 3.5 times more likely to immediately seek refuge without displaying in one of the many rock crevices adjacent to (<0.5 m) to their perches (t 2, 32 = 4.48, Cohen s d, 95% CI = 0.81, ) compared with lizards in lowthreat trials that more frequently remained emergent and displayed. Fig. 3: The percentage of time spent tail displaying during non-threat versus low-threat trials. Data are means 1.0 SE. Asterisk indicates a statistically significant effect (95% CI did not included zero). Fig. 4: The percentage of time spent tail displaying during low-threat versus high-threat trials. Data are means 1.0 SE. Asterisk indicates a statistically significant effect (95% CI did not included zero). Discussion Juvenile collared lizards consistently gave tail displays in response to approach by a simulated predator, but never gave these displays during non-threat trials. Moreover, during low-threat trials, subjects increased the frequency of tail display, whereas they usually sought refuge without displaying when approached from angles of increased predatory threat. When subjects were exposed to lower threat because they were more hidden and less able to judge speed and distance of the simulated predator, the costs of signaling to potential predators were likely reduced (Cooper 2008). By contrast, increased exposure during highthreat trials apparently made the cost of displaying prohibitively high, prompting lizards to instead take refuge without displaying. The selective trade-off between threat level and the decision to display is likely acute in species with highly developed vision such as collared lizards (Fox & Baird 1992), which can likely use visual cues of the rate and directionality of approach by predators to assess the relative intensity of threats (Burger & Gochfeld 1981; Braun et al. 2010). Nevertheless, it remains possible that other variables that we could not control during our experiments on free-ranging lizards (e.g., proximity to refuges, conspicuousness/vulnerability to predators) may have influenced perceptions of threat intensity, and hence decisions to display or take refuge. Our results support the hypothesis that tail displays given by juvenile collared lizards function as antipredator signals, similar to displays given by other lizards when threatened (Cooper et al. 2004; Font et al. 2012). Based on our experiments, we cannot unequivocally accept or reject any of the three hypothesized mechanisms proposed to explain how tail displays function in predator avoidance. One possibility is that tail displays deflect attacks to a nonessential body part. In the majority of lizards and other taxa in which displays function for deflection, the non-essential body parts are conspicuously colored and these species possess autotomy septa that suddenly sever the tail when it is struck affording prey the opportunity to escape (Hill & Vaca 2004; Telemeco et al. 2011). Collared lizards lack both conspicuous caudal coloration and autotomy septa. Nevertheless, the tail remains an expendable body part because it is not essential for immediate survival. Although the absence of both conspicuous caudal coloration and autotomy makes it unlikely that the deflection hypothesis explains the evolution of tail displays in juvenile collared lizards, we cannot rule out this possibility entirely. Ethology 122 (2016) Blackwell Verlag GmbH 41

6 Antipredator Tail Displays in a Lizard J. R. York & T. A. Baird It also seems unlikely that tail displays advertise relative escape ability in our population. Lizard escape ability is often estimated by measuring sprint speed in the laboratory on flat straight tracks. Lizards are elicited to run by prodding them, and maximum sprint speed is measured as that attained at the end of the track (Husak 2006a, b; Husak & Fox 2006; Telemeco et al. 2011). Because we did not measure sprint speed, we cannot rule out a possible relationship between sprinting ability and tail display behavior, but this seems unlikely because it is not necessary for collared lizards at our site to run to access refuges. Because crevices are very abundant throughout our study site (Baird & Sloan 2003; York & Baird 2015), study subjects escaped by darting into refuges that were immediately adjacent (within 0.5 m) on at least two sides of chosen perches, a distance so short that maximal sprint speed is probably not even possible to attain. Other performance traits such as reaction time and the ability to move over very short distances into a crevice may promote escape behavior in collared lizards. However, we are not aware of any published data on such traits in lizards or any other vertebrates. Taken together, our current results suggest that the most likely adaptive explanation of increased tail display frequency in response to low levels of threat is to signal predators that they have been detected (perception advertisement, sensu Caro 1995). Traits that evolved originally for one function are sometimes selectively co-opted for different functions (Bock 1958; Gould & Vrba 1982; Endler & Basolo 1998), and this may have occurred in the case of antipredator displays in some species. For example, tail displays that evolved initially to promote intraspecific communication may subsequently take on an antipredator signaling role (Johnson & Wade 2010). Intraspecific selection associated with sexual behavior is not likely in reproductively immature juvenile collared lizards. However, observations of juvenile collared lizards performing sinusoidal tail movements while scanning the surrounding grass for insect prey and tail curling displays when juveniles were stalking grasshoppers both suggest that tail displays may also play a role in foraging (Braun & Baird unpublished data). Effective sit-and-wait foraging requires that lizards remain exposed on elevated perches for extended periods, whereas departing good foraging sites to take refuge disrupts opportunities to ambush prey (Scarratt & Godin 1992; Perez-Tris et al. 2004). Interruption of foraging activities would be especially costly for juvenile lizards because of their high energetic demands to support rapid growth and to build energy stores to survive the winter (Baird 2008; Braun et al. 2010). Tail displays that involve deliberate tail movements and postures may visually distract prey just long enough to give lizards a momentary advantage when they strike (see similarly, Baird 2008). Because ambush foraging requires sitting exposed on perches such that lizards are vulnerable, it is entirely possible that tail displays that evolved originally to deter predators may have been co-opted later to improve foraging efficiency through distraction of prey, or vice versa. Acknowledgments We thank Bill Parkerson of the US Army Corps of Engineers for permission to conduct field studies at the Arcadia Lake site. The Office of Research and Grants at the University of Central Oklahoma provided funding to JRY and TAB. We also thank Teresa Baird, Sharon LaFave, Will Unsell, and Emily York for providing technical assistance. Permission to conduct this research was provided by the Oklahoma Department of Wildlife, and all procedures were approved by the Institutional Animal Care and Use Committee of the University of Central Oklahoma. Conflict of Interest The authors declare no conflict of interest. Literature Cited Amo, L., Lopez, P. & Martın, J. 2007: Refuge use: a conflict between avoiding predation and using mass in lizards. Physiol. Behav. 22, Baird, T. A. 2008: A growth cost of experimentally induced conspicuous coloration in first-year collared lizard males. Behav. Ecol. 19, Baird, T. A. 2013a: Social life on the rocks: behavioral diversity and sexual selection in collared lizards. In: Reptiles in Research: Investigations of Ecology, Physiology, and Behavior From Desert to Sea. (Lutterschmidt, W. I., ed). Nova Biomedical Press, New York, NY, pp Baird, T. A. 2013b: Lizards and other reptiles as model systems for the study of aggressive contest behaviour. In: Animal Contests. (Hardy, I. C. W. & Briffa, M., eds). Cambridge Univ. Press, Cambridge, pp Baird, T. A. & Sloan, C. L. 2003: Interpopulation variation in the social organization of female collared lizards, Crotaphytus collaris. Ethology 109, Baird, T. A., Sloan, C. L. & Timanus, D. K. 2001: Intra-and inter-seasonal variation in the socio-spatial behavior of 42 Ethology 122 (2016) Blackwell Verlag GmbH

7 J. R. York & T. A. Baird Antipredator Tail Displays in a Lizard adult male collared lizards, Crotaphytus collaris (Reptilia, Crotaphytidae). Ethology 107, Baird, T. A., Timanus, D. K. & Sloan, C. L. 2003: Intra- and intersexual variation in social behavior: effects of ontogeny, phenotype, resources, and season. In: Lizard Social Behavior. (Fox, S. F., McCoy, J. K. & Baird, T. A., eds). Johns Hopkins Univ. Press, Baltimore, MD, pp Bateman, P. W. & Fleming, P. A. 2009: To cut a long tail short: a review of lizard caudal autotomy studies carried out over the last 20 years. J. Zool. 277, Bock, W. J. 1958: Preadaptation and multiple evolutionary pathways. Evolution 13, Braun, C. A., Baird, T. A. & LeBeau, J. K. 2010: Influence of substrate temperature and directness of approach on the escape responses of juvenile collared lizards. Herpetologica 66, Broom, M. & Ruxton, G. D. 2005: You can run, or you can hide: optimal strategies for cryptic prey. Behav. Ecol. 16, Burger, J. & Gochfeld, M. 1981: Discrimination of the threat of direct versus tangential approach to the nest by incubating herring and great black-backed gulls. J. Comp. Physiol. Psychol. 95, Caro, T. M. 1995: Pursuit-deterrence revisited. Trends Ecol. Evol. 10, Caro, T. M. 2005: Antipredator Defences in Birds and Mammals, 1st edn. Univ. Chicago Press, Chicago, IL. Cohen, J. 1992: A power primer. Psychol. Bull. 112, 155. Colegrave, N. & Ruxton, G. D. 2003: Confidence intervals are a more useful complement to nonsignificant tests than are power calculations. Behav. Ecol. 14, Cooper, W. E. Jr 1998: Reactive and anticipatory display to deflect predatory attack to an autotomous lizard tail. Can. J. Zool. 76, Cooper, W. E. Jr 2003: Effect of risk on aspects of escape behavior by a lizard, Holbrookia propinqua, in relation to optimal escape theory. Ethology 109, Cooper, W. E. Jr 2008: Visual monitoring of predators: occurrence, cost and benefit for escape. Anim. Behav. 76, Cooper, W. E. Jr & Vitt, L. J. 2010: Blue tails and autotomy: enhancement of predation avoidance in juvenile skinks. Ethology 70, Cooper, W. E. Jr, Perez-Mellado, V., Baird, T. A., Caldwell, J. P. & Vitt, L. J. 2004: Pursuit deterrent signalling by the Bonaire whiptail lizard Cnemidophorus murinus. Behaviour 141, Endler, J. A. & Basolo, A. L. 1998: Sensory ecology, receiver biases and sexual selection. Trends Ecol. Evol. 13, Font, E., Carazo, P., Perez, G. & Kramer, M. 2012: Predator-elicited foot shakes in wall lizards (Podarcis muralis): evidence for a pursuit-deterrent function. J. Comp. Psychol. 126, Fox, S. F. & Baird, T. A. 1992: The dear enemy phenomenon in the collared lizard, Crotaphytus collaris, with a cautionary note on experimental methodology. Anim. Behav. 44, Gould, S. J. & Vrba, E. S. 1982: Exaptation a missing term in the science of form. Paleobiology 8, Greene, H. W. 1988: Antipredator mechanisms in reptiles. In: Biology of the Reptilia, Vol. 16. (Gans, C. & Huey, R. B., eds). John Wiley & Sons, New York, NY, pp Hasson, O., Hibbard, R. & Ceballos, G. 1989: The pursuit deterrent function of tail-wagging in the zebra-tailed lizard (Callisaurus draconoides). Can. J. Zool. 67, Hill, R. I. & Vaca, J. F. 2004: Differential wing strength in Pierella butterflies (Nymphalidae, Satyrinae) supports the deflection hypothesis. Biotropica 36, Husak, J. F. 2006a: Does speed help you survive? A test with collared lizards of different ages. Funct. Ecol. 20, Husak, J. F. 2006b: Does survival depend on how fast you can run or how fast you do run? Funct. Ecol. 20, Husak, J. F. & Fox, S. F. 2006: Field use of maximal sprint speed by collared lizards (Crotaphytus collaris): compensation and sexual selection. Evolution 60, Johnson, M. A. & Wade, J. 2010: Behavioural display systems across nine Anolis lizard species: sexual dimorphisms in structure and function. Proc. R. Soc. Lond. B Biol. Sci. 277, Langkilde, T., Schwarzkopf, L. & Alford, R. A. 2004: The function of tail displays in male rainbow skinks (Carlia jarnoldae). J. Herpetol. 37, Martın, J. & Lopez, P. 1999: An experimental test of the costs of antipredatory refuge use in the wall lizard, Podarcis muralis. Oikos 84, Nakagawa, S. & Cuthill, I. C. 2007: Effect size, confidence interval and statistical significance: a practical guide for biologists. Biol. Rev. (Camb). 82, Perez-Tris, J., Di az, J. A. & Telleri a, J. L. 2004: Loss of body mass under predation risk: cost of antipredatory behaviour or adaptive fit-for-escape? Anim. Behav. 67, R Development Core Team. 2013: R: A language and environment for statistical computing. Vienna, Austria: R Foundation for Statistical Computing. Ruxton, G. D., Sherratt, T. N. & Speed, M. P. 2004: Avoiding Attack: The Evolutionary Ecology of Crypsis, Warning Signals and Mimicry, 1st edn. Oxford Univ. Press, Oxford, UK. Scarratt, A. M. & Godin, J. G. J. 1992: Foraging and antipredator decisions in the hermit crab Pagurus Ethology 122 (2016) Blackwell Verlag GmbH 43

8 Antipredator Tail Displays in a Lizard J. R. York & T. A. Baird acadianus (Benedict). J. Exp. Mar. Biol. Ecol. 156, Stankowich, T. & Blumstein, D. T. 2005: Fear in animals: a meta-analysis and review of risk assessment. Proc. R. Soc. Lond. B Biol. Sci. 272, Telemeco, R. S., Baird, T. A. & Shine, R. 2011: Tail waving in a lizard (Bassiana duperreyi) functions to deflect attacks rather than as a pursuit-deterrent signal. Anim. Behav. 82, Ydenberg, R. C. & Dill, L. M. 1986: The economics of fleeing from predators. Adv. Study Behav. 16, York, J. R. & Baird, T. A. 2015: Testing the adaptive significance of sex-specific mating tactics in collared lizards (Crotaphytus collaris). Biol. J. Linn. Soc. 115, York, J. R., Baird, T. A. & Haynie, M. L. 2014: Unexpected high fitness payoff of subordinate social tactics in male collared lizards. Anim. Behav. 91, Ethology 122 (2016) Blackwell Verlag GmbH

DOES EXPERIMENTALLY INDUCED CONSPICUOUS COLORATION INCREASE RISK OF PREDATION AND CONSPECIFIC AGGRESSION IN FIRST-YEAR COLLARED LIZARD MALES?

DOES EXPERIMENTALLY INDUCED CONSPICUOUS COLORATION INCREASE RISK OF PREDATION AND CONSPECIFIC AGGRESSION IN FIRST-YEAR COLLARED LIZARD MALES? Herpetologica, 65(1), 2009, 31 38 E 2009 by The Herpetologists League, Inc. DOES EXPERIMENTALLY INDUCED CONSPICUOUS COLORATION INCREASE RISK OF PREDATION AND CONSPECIFIC AGGRESSION IN FIRST-YEAR COLLARED

More information

The role of visual cues in learning escape behaviour in the little brown skink (Scincella lateralis)

The role of visual cues in learning escape behaviour in the little brown skink (Scincella lateralis) Behaviour 151 (2014) 2015 2028 brill.com/beh The role of visual cues in learning escape behaviour in the little brown skink (Scincella lateralis) Mark A. Paulissen Department of Natural Sciences, Northeastern

More information

Uncertainty about future predation risk modulates monitoring behavior from refuges in lizards

Uncertainty about future predation risk modulates monitoring behavior from refuges in lizards Behavioral Ecology doi:10.1093/beheco/arq065 Advance Access publication 13 January 2011 Original Article Uncertainty about future predation risk modulates monitoring behavior from refuges in lizards Vicente

More information

Animal Behaviour 78 (2009) Contents lists available at ScienceDirect. Animal Behaviour. journal homepage:

Animal Behaviour 78 (2009) Contents lists available at ScienceDirect. Animal Behaviour. journal homepage: Animal Behaviour 78 (2009) 1011 1018 Contents lists available at ScienceDirect Animal Behaviour journal homepage: www.elsevier.com/locate/anbehav When to run from an ambush predator: balancing crypsis

More information

From ethology to sexual selection: trends in animal behavior research. Animal behavior then & now

From ethology to sexual selection: trends in animal behavior research. Animal behavior then & now From ethology to sexual selection: trends in animal behavior research Terry J. Ord, Emília P. Martins Department of Biology, Indiana University Sidharth Thakur Computer Science Department, Indiana University

More information

Ciccaba virgata (Mottled Owl)

Ciccaba virgata (Mottled Owl) Ciccaba virgata (Mottled Owl) Family: Strigidae (Typical Owls) Order: Strigiformes (Owls) Class: Aves (Birds) Fig. 1. Mottled owl, Ciccaba virgata. [http://www.owling.com/mottled13.htm, downloaded 12 November

More information

Sex differences in antipredator tail-waving displays of the diurnal yellow-headed gecko Gonatodes albogularis from tropical forests of Colombia

Sex differences in antipredator tail-waving displays of the diurnal yellow-headed gecko Gonatodes albogularis from tropical forests of Colombia J Ethol (2010) 28:305 311 DOI 10.1007/s10164-009-0186-4 ARTICLE Sex differences in antipredator tail-waving displays of the diurnal yellow-headed gecko Gonatodes albogularis from tropical forests of Colombia

More information

Contrasting Response to Predator and Brood Parasite Signals in the Song Sparrow (melospiza melodia)

Contrasting Response to Predator and Brood Parasite Signals in the Song Sparrow (melospiza melodia) Luke Campillo and Aaron Claus IBS Animal Behavior Prof. Wisenden 6/25/2009 Contrasting Response to Predator and Brood Parasite Signals in the Song Sparrow (melospiza melodia) Abstract: The Song Sparrow

More information

Plestiodon (=Eumeces) fasciatus Family Scincidae

Plestiodon (=Eumeces) fasciatus Family Scincidae Plestiodon (=Eumeces) fasciatus Family Scincidae Living specimens: - Five distinct longitudinal light lines on dorsum - Juveniles have bright blue tail - Head of male reddish during breeding season - Old

More information

Crotophaga major (Greater Ani)

Crotophaga major (Greater Ani) Crotophaga major (Greater Ani) Family: Cuculidae (Cuckoos and Anis) Order: Cuculiformes (Cuckoos, Anis and Turacos) Class: Aves (Birds) Fig. 1. Greater ani, Crotophaga major. [http://www.birdforum.net/opus/greater_ani,

More information

Is it better to be bigger? Featured scientists: Aaron Reedy and Robert Cox from the University of Virginia Co-written by Matt Kustra

Is it better to be bigger? Featured scientists: Aaron Reedy and Robert Cox from the University of Virginia Co-written by Matt Kustra Is it better to be bigger? Featured scientists: Aaron Reedy and Robert Cox from the University of Virginia Co-written by Matt Kustra Research Background: When Charles Darwin talked about the struggle for

More information

Red-Tailed Hawk Buteo jamaicensis

Red-Tailed Hawk Buteo jamaicensis Red-Tailed Hawk Buteo jamaicensis This large, dark headed, broad-shouldered hawk is one of the most common and widespread hawks in North America. The Red-tailed hawk belongs to the genus (family) Buteo,

More information

Analysis of Sampling Technique Used to Investigate Matching of Dorsal Coloration of Pacific Tree Frogs Hyla regilla with Substrate Color

Analysis of Sampling Technique Used to Investigate Matching of Dorsal Coloration of Pacific Tree Frogs Hyla regilla with Substrate Color Analysis of Sampling Technique Used to Investigate Matching of Dorsal Coloration of Pacific Tree Frogs Hyla regilla with Substrate Color Madeleine van der Heyden, Kimberly Debriansky, and Randall Clarke

More information

J.K. McCoy CURRICULUM VITAE. J. Kelly McCoy. Department of Biology Angelo State University San Angelo, TX

J.K. McCoy CURRICULUM VITAE. J. Kelly McCoy. Department of Biology Angelo State University San Angelo, TX CURRICULUM VITAE J. Kelly McCoy Department of Biology Angelo State University San Angelo, TX 76909 325-486-6646 Kelly.McCoy@angelo.edu Education: B.S. 1990 Zoology Oklahoma State University Ph.D. 1995

More information

Habitats and Field Methods. Friday May 12th 2017

Habitats and Field Methods. Friday May 12th 2017 Habitats and Field Methods Friday May 12th 2017 Announcements Project consultations available today after class Project Proposal due today at 5pm Follow guidelines posted for lecture 4 Field notebooks

More information

Migration. Migration = a form of dispersal which involves movement away from and subsequent return to the same location, typically on an annual basis.

Migration. Migration = a form of dispersal which involves movement away from and subsequent return to the same location, typically on an annual basis. Migration Migration = a form of dispersal which involves movement away from and subsequent return to the same location, typically on an annual basis. To migrate long distance animals must navigate through

More information

Seasonal Shifts in Reproductive Investment of Female Northern Grass Lizards ( Takydromus septentrionalis

Seasonal Shifts in Reproductive Investment of Female Northern Grass Lizards ( Takydromus septentrionalis Seasonal Shifts in Reproductive Investment of Female Northern Grass Lizards (Takydromus septentrionalis) from a Field Population on Beiji Island, China Author(s): Wei-Guo Du and Lu Shou Source: Journal

More information

Effects of prey availability and climate across a decade for a desert-dwelling, ectothermic mesopredator. R. Anderson Western Washington University

Effects of prey availability and climate across a decade for a desert-dwelling, ectothermic mesopredator. R. Anderson Western Washington University Effects of prey availability and climate across a decade for a desert-dwelling, ectothermic mesopredator R. Anderson Western Washington University Trophic interactions in desert systems are presumed to

More information

Rubber Boas in Radium Hot Springs: Habitat, Inventory, and Management Strategies

Rubber Boas in Radium Hot Springs: Habitat, Inventory, and Management Strategies : Habitat, Inventory, and Management Strategies ROBERT C. ST. CLAIR 1 AND ALAN DIBB 2 1 9809 92 Avenue, Edmonton, AB, T6E 2V4, Canada, email rstclair@telusplanet.net 2 Parks Canada, Box 220, Radium Hot

More information

Intraspecific relationships extra questions and answers (Extension material for Level 3 Biology Study Guide, ISBN , page 153)

Intraspecific relationships extra questions and answers (Extension material for Level 3 Biology Study Guide, ISBN , page 153) i Intraspecific relationships extra questions and answers (Extension material for Level 3 Biology Study Guide, ISBN 978-1-927194-58-4, page 153) Activity 9: Intraspecific relationships extra questions

More information

Like mother, like daughter: inheritance of nest-site

Like mother, like daughter: inheritance of nest-site Like mother, like daughter: inheritance of nest-site location in snakes Gregory P. Brown and Richard Shine* School of Biological Sciences A0, University of Sydney, NSW 00, Australia *Author for correspondence

More information

Escape Behaviors and Flight Initiation Distance in the Common Water Snake Nerodia sipedon

Escape Behaviors and Flight Initiation Distance in the Common Water Snake Nerodia sipedon Journal of Herpetology, Vol. 42, No. 3, pp. 493 500, 2008 Copyright 2008 Society for the Study of Amphibians and Reptiles Escape Behaviors and Flight Initiation Distance in the Common Water Snake Nerodia

More information

Predation Cost of Conspicuous Male Coloration in Collared Lizards (Crotaphytus collaris): An Experimental Test Using Clay-Covered Model Lizards

Predation Cost of Conspicuous Male Coloration in Collared Lizards (Crotaphytus collaris): An Experimental Test Using Clay-Covered Model Lizards Ethology Predation Cost of Conspicuous Male Coloration in Collared Lizards (Crotaphytus collaris): An Experimental Test Using Clay-Covered Model Lizards Jerry F. Husak*, Joseph M. Macedonia, Stanley F.

More information

Signalling displays during predator prey interactions in a Puerto Rican anole, Anolis cristatellus

Signalling displays during predator prey interactions in a Puerto Rican anole, Anolis cristatellus Anim. Behav., 1997, 54, 1147 1154 Signalling displays during predator prey interactions in a Puerto Rican anole, Anolis cristatellus MANUEL LEAL* & JAVIER A. RODRIuGUEZ-ROBLES *Department of Biology, Washington

More information

Escaping predators on vertical surfaces: Lacerta perspicillata in limestone quarries of Lithaca

Escaping predators on vertical surfaces: Lacerta perspicillata in limestone quarries of Lithaca Escaping predators on vertical surfaces: Lacerta perspicillata in limestone quarries of Lithaca Laurie J. Vitt, William E. Cooper, Jr., Anna Perera, and Valentín Pérez-Mellado 1803 Abstract: Escape behavior

More information

A.13 BLAINVILLE S HORNED LIZARD (PHRYNOSOMA BLAINVILLII)

A.13 BLAINVILLE S HORNED LIZARD (PHRYNOSOMA BLAINVILLII) A. BLAINVILLE S HORNED LIZARD (PHRYNOSOMA BLAINVILLII) A.. Legal and Other Status Blainville s horned lizard is designated as a Department of Fish and Game (DFG) Species of Concern. A.. Species Distribution

More information

Animal Adaptations. Structure and Function

Animal Adaptations. Structure and Function Name period date assigned date due date returned 1. What is a variation 2. What is an adaptation omplete the chart with the examples from the power point. List adaptations that help animals do the following:

More information

Prey or predator? Body size of an approaching animal affects decisions to attack or escape

Prey or predator? Body size of an approaching animal affects decisions to attack or escape Behavioral Ecology doi:10.1093/beheco/arq142 Advance Access publication 21 September 2010 Prey or predator? Body size of an approaching animal affects decisions to attack or escape William E. Cooper Jr

More information

Introduction. Lizards: very diverse colour patterns intra- and interspecific differences in colour

Introduction. Lizards: very diverse colour patterns intra- and interspecific differences in colour Jessica Vroonen Introduction Lizards: very diverse colour patterns intra- and interspecific differences in colour Introduction Lizards intra- and interspecific differences in colour Introduction Lizards

More information

4B: The Pheasant Case: Handout. Case Three Ring-Necked Pheasants. Case materials: Case assignment

4B: The Pheasant Case: Handout. Case Three Ring-Necked Pheasants. Case materials: Case assignment 4B: The Pheasant Case: Handout Case Three Ring-Necked Pheasants As you can see, the male ring-necked pheasant is brightly colored. The white ring at the base of the red and green head stand out against

More information

Flexibility in antipredatory behavior allows wall lizards to cope with multiple types of predators

Flexibility in antipredatory behavior allows wall lizards to cope with multiple types of predators Ann. Zool. Fennici 42: 109 121 ISSN 0003-455X Helsinki 26 April 2005 Finnish Zoological and Botanical Publishing Board 2005 Flexibility in antipredatory behavior allows wall lizards to cope with multiple

More information

Adjustments In Parental Care By The European Starling (Sturnus Vulgaris): The Effect Of Female Condition

Adjustments In Parental Care By The European Starling (Sturnus Vulgaris): The Effect Of Female Condition Proceedings of The National Conference on Undergraduate Research (NCUR) 2003 University of Utah, Salt Lake City, Utah March 13-15, 2003 Adjustments In Parental Care By The European Starling (Sturnus Vulgaris):

More information

ethology Ethology RESEARCH PAPER

ethology Ethology RESEARCH PAPER international journal of behavioural biology ethology Ethology RESEARCH PAPER The Effect of Human Presence and Human Activity on Risk Assessment and Flight Initiation Distance in Skinks Marilyn M. McGowan,

More information

08 alberts part2 7/23/03 9:10 AM Page 95 PART TWO. Behavior and Ecology

08 alberts part2 7/23/03 9:10 AM Page 95 PART TWO. Behavior and Ecology 08 alberts part2 7/23/03 9:10 AM Page 95 PART TWO Behavior and Ecology 08 alberts part2 7/23/03 9:10 AM Page 96 08 alberts part2 7/23/03 9:10 AM Page 97 Introduction Emília P. Martins Iguanas have long

More information

8/19/2013. Topic 14: Body support & locomotion. What structures are used for locomotion? What structures are used for locomotion?

8/19/2013. Topic 14: Body support & locomotion. What structures are used for locomotion? What structures are used for locomotion? Topic 4: Body support & locomotion What are components of locomotion? What structures are used for locomotion? How does locomotion happen? Forces Lever systems What is the difference between performance

More information

The S Files Success with Maria: Sunshine: Biting Reported by S.G. Friedman, PhD and L. McGuire

The S Files Success with Maria: Sunshine: Biting Reported by S.G. Friedman, PhD and L. McGuire The S Files Success with Maria: Sunshine: Biting Reported by S.G. Friedman, PhD and L. McGuire In Press, Good Bird Magazine Volume x(x), pp-pp The S Files are real case studies of behavior challenges faced

More information

Tail Autotomy Does Not Increase Locomotor Costs in the Oriental Leaf-toed Gecko Hemidactylus bowringii

Tail Autotomy Does Not Increase Locomotor Costs in the Oriental Leaf-toed Gecko Hemidactylus bowringii Asian Herpetological Research 2012, 3(2): 141 146 DOI: 10.3724/SP.J.1245.2012.00141 Tail Autotomy Does Not Increase Locomotor Costs in the Oriental Leaf-toed Gecko Hemidactylus bowringii Guohua DING, Tianbao

More information

PORTRAIT OF THE AMERICAN BALD EAGLE

PORTRAIT OF THE AMERICAN BALD EAGLE PORTRAIT OF THE AMERICAN BALD EAGLE Objectives: To know the history of the bald eagle and the cause of it's decline. To understand what has been done to improve Bald Eagle habitat. To know the characteristics

More information

Requirements for the employment as helper in phase C

Requirements for the employment as helper in phase C GSSCC - Raino Fluegge, President Page 1 of 8 A.) Requirements for the employment as helper in phase C 1. The guidelines and regulations of the trial regulations regarding helper work must be followed.

More information

Use of Agent Based Modeling in an Ecological Conservation Context

Use of Agent Based Modeling in an Ecological Conservation Context 28 RIThink, 2012, Vol. 2 From: http://photos.turksandcaicostourism.com/nature/images/tctb_horz_033.jpg Use of Agent Based Modeling in an Ecological Conservation Context Scott B. WOLCOTT 1 *, Michael E.

More information

Egyptian vulture (Neophron percnopterus) research & monitoring Breeding Season Report- Beypazarı, Turkey

Egyptian vulture (Neophron percnopterus) research & monitoring Breeding Season Report- Beypazarı, Turkey Egyptian vulture (Neophron percnopterus) research & monitoring - 2011 Breeding Season Report- Beypazarı, Turkey October 2011 1 Cover photograph: Egyptian vulture landing in Beypazarı dump site, photographed

More information

Motuora island reptile monitoring report for common & Pacific gecko 2016

Motuora island reptile monitoring report for common & Pacific gecko 2016 Motuora island reptile monitoring report for common & Pacific gecko 6 Prepared by Su Sinclair August 7 Work on this monitoring project was carried out under a Wildlife Act Authority issued by the Department

More information

A tail of two scorpions Featured scientists: Ashlee Rowe and Matt Rowe from University of Oklahoma

A tail of two scorpions Featured scientists: Ashlee Rowe and Matt Rowe from University of Oklahoma A tail of two scorpions Featured scientists: Ashlee Rowe and Matt Rowe from University of Oklahoma Animals have evolved many ways to defend themselves against predators. Many species use camouflage to

More information

A Population Analysis of the Common Wall Lizard Podarcis muralis in Southwestern France

A Population Analysis of the Common Wall Lizard Podarcis muralis in Southwestern France - 513 - Studies in Herpetology, Rocek Z. (ed.) pp. 513-518 Prague 1986 A Population Analysis of the Common Wall Lizard Podarcis muralis in Southwestern France R. BARBAULT and Y. P. MOU Laboratoire d'ecologie

More information

King penguin brooding and defending a sub-antarctic skua chick

King penguin brooding and defending a sub-antarctic skua chick King penguin brooding and defending a sub-antarctic skua chick W. Chris Oosthuizen 1 and P. J. Nico de Bruyn 1 (1) Department of Zoology and Entomology, Mammal Research Institute, University of Pretoria,

More information

BLACK OYSTERCATCHER NEST MONITORING PROTOCOL

BLACK OYSTERCATCHER NEST MONITORING PROTOCOL BLACK OYSTERCATCHER NEST MONITORING PROTOCOL In addition to the mid-late May population survey (see Black Oystercatcher abundance survey protocol) we will attempt to continue monitoring at least 25 nests

More information

Mental stim ulation it s not just for dogs!! By Danielle Middleton- Beck BSc hons, PGDip CABC

Mental stim ulation it s not just for dogs!! By Danielle Middleton- Beck BSc hons, PGDip CABC Milo, Congo African Grey by Elaine Henley Mental stim ulation it s not just for dogs!! By Danielle Middleton- Beck BSc hons, PGDip CABC Dexter, Green Iguana by Danielle Middleton-Beck Exotic pets include

More information

Supporting Online Material for

Supporting Online Material for www.sciencemag.org/cgi/content/full/314/5802/1111/dc1 Supporting Online Material for Rapid Temporal Reversal in Predator-Driven Natural Selection Jonathan B. Losos,* Thomas W. Schoener, R. Brian Langerhans,

More information

Short-term Water Potential Fluctuations and Eggs of the Red-eared Slider Turtle (Trachemys scripta elegans)

Short-term Water Potential Fluctuations and Eggs of the Red-eared Slider Turtle (Trachemys scripta elegans) Zoology and Genetics Publications Zoology and Genetics 2001 Short-term Water Potential Fluctuations and Eggs of the Red-eared Slider Turtle (Trachemys scripta elegans) John K. Tucker Illinois Natural History

More information

Ardea herodias (Great Blue Heron)

Ardea herodias (Great Blue Heron) Ardea herodias (Great Blue Heron) Family: Ardeidae (Herons and Egrets) Order: Ciconiiformes (Storks, Herons and Ibises) Class: Aves (Birds) Fig.1. Great blue heron, Ardea herodias. [http://birdingbec.blogspot.com,

More information

Iguana aggression. A relaxed green iguana. Defensive aggression

Iguana aggression. A relaxed green iguana. Defensive aggression Iguana aggression Iguanas are still wild animals, they are not domestic animals, and they have just been tamed to enable them to fit into a human lifestyle. Ideally iguanas should be housed in a large

More information

Shooting the poop Featured scientist: Martha Weiss from Georgetown University

Shooting the poop Featured scientist: Martha Weiss from Georgetown University Research Background: Shooting the poop Featured scientist: Martha Weiss from Georgetown University Imagine walking through a forest in the middle of summer. You can hear birds chirping, a slight breeze

More information

New Mexico Avian Protection (NMAP) Feather Identification Guide

New Mexico Avian Protection (NMAP) Feather Identification Guide New Mexico Avian Protection (NMAP) Feather Identification Guide It is very common to find only feathers as remains beneath a power line due to predation, length of elapsed time since the mortality, weather,

More information

BEHAVIOUR OF DOGS DURING OLFACTORY TRACKING

BEHAVIOUR OF DOGS DURING OLFACTORY TRACKING J. exp. Biol. 180, 247-251 (1993) Printed in Great Britain The Company of Biologists Limited 1993 247 BEHAVIOUR OF DOGS DURING OLFACTORY TRACKING AUD THESEN, JOHAN B. STEEN* and KJELL B. DØVING Division

More information

Everglades Invasive Reptile and Amphibian Monitoring Program 1

Everglades Invasive Reptile and Amphibian Monitoring Program 1 WEC386 Everglades Invasive Reptile and Amphibian Monitoring Program 1 Rebecca G. Harvey, Mike Rochford, Jennifer Ketterlin, Edward Metzger III, Jennifer Nestler, and Frank J. Mazzotti 2 Introduction South

More information

Departamento de Biologia Animal, Universidad de Salamanca, Salamanca, Spain. INTRODUCTION

Departamento de Biologia Animal, Universidad de Salamanca, Salamanca, Spain. INTRODUCTION Acta Herpetologica 6(2): 247-259, 2011 Escape by the Balearic Lizard (Podarcis lilfordi) is affected by elevation of an approaching predator, but not by some other potential predation risk factors William

More information

AN APPLIED CASE STUDY of the complexity of ecological systems and process: Why has Lyme disease become an epidemic in the northeastern U.S.

AN APPLIED CASE STUDY of the complexity of ecological systems and process: Why has Lyme disease become an epidemic in the northeastern U.S. AN APPLIED CASE STUDY of the complexity of ecological systems and process: Why has Lyme disease become an epidemic in the northeastern U.S. over the last few decades? What causes Lyme disease? 1 Frequency

More information

Biol 160: Lab 7. Modeling Evolution

Biol 160: Lab 7. Modeling Evolution Name: Modeling Evolution OBJECTIVES Help you develop an understanding of important factors that affect evolution of a species. Demonstrate important biological and environmental selection factors that

More information

8/19/2013. Who eats herps? Topic 20: Predators. Who eats herps? Who eats herps? Who eats herps? Who eats herps?

8/19/2013. Who eats herps? Topic 20: Predators. Who eats herps? Who eats herps? Who eats herps? Who eats herps? Topic 20: Predators Variation in predators across taxa Variation in predators through ontogeny How do herps avoid being eaten? Introduction to the diversity of anti-predator defenses Many animals Depends

More information

NOTES ON THE ECOLOGY AND NATURAL HISTORY OF CTENOPHORUS CAUDICINCTUS (AGAMIDAE) IN WESTERN AUSTRALIA

NOTES ON THE ECOLOGY AND NATURAL HISTORY OF CTENOPHORUS CAUDICINCTUS (AGAMIDAE) IN WESTERN AUSTRALIA NOTES ON THE ECOLOGY AND NATURAL HISTORY OF CTENOPHORUS CAUDICINCTUS (AGAMIDAE) IN WESTERN AUSTRALIA By ERIC R. PIANKA Integrative Biology University of Texas at Austin Austin, Texas 78712 USA Email: erp@austin.utexas.edu

More information

Objectives: Outline: Idaho Amphibians and Reptiles. Characteristics of Amphibians. Types and Numbers of Amphibians

Objectives: Outline: Idaho Amphibians and Reptiles. Characteristics of Amphibians. Types and Numbers of Amphibians Natural History of Idaho Amphibians and Reptiles Wildlife Ecology, University of Idaho Fall 2005 Charles R. Peterson Herpetology Laboratory Department of Biological Sciences, Idaho Museum of Natural History

More information

Behaviour and spatial ecology of Gilbert s dragon Lophognathus gilberti (Agamidae: Reptilia)

Behaviour and spatial ecology of Gilbert s dragon Lophognathus gilberti (Agamidae: Reptilia) Journal of the Royal Society of Western Australia, 84:153-158, 2001 Behaviour and spatial ecology of Gilbert s dragon Lophognathus gilberti (Agamidae: Reptilia) G G Thompson 1 & S A Thompson 2 1 Edith

More information

Survivorship. Demography and Populations. Avian life history patterns. Extremes of avian life history patterns

Survivorship. Demography and Populations. Avian life history patterns. Extremes of avian life history patterns Demography and Populations Survivorship Demography is the study of fecundity and survival Four critical variables Age of first breeding Number of young fledged each year Juvenile survival Adult survival

More information

Density, growth, and home range of the lizard Uta stansburiana stejnegeri in southern Dona Ana County, New Mexico

Density, growth, and home range of the lizard Uta stansburiana stejnegeri in southern Dona Ana County, New Mexico Great Basin Naturalist Volume 33 Number 2 Article 8 6-30-1973 Density, growth, and home range of the lizard Uta stansburiana stejnegeri in southern Dona Ana County, New Mexico Richard D. Worthington University

More information

Avian Ecology: Life History, Breeding Seasons, & Territories

Avian Ecology: Life History, Breeding Seasons, & Territories Avian Ecology: Life History, Breeding Seasons, & Territories Life History Theory Why do some birds lay 1-2 eggs whereas others 12+? Why do some species begin reproducing at < 1 year whereas others not

More information

Northern Copperhead Updated: April 8, 2018

Northern Copperhead Updated: April 8, 2018 Interpretation Guide Northern Copperhead Updated: April 8, 2018 Status Danger Threats Population Distribution Habitat Diet Size Longevity Social Family Units Reproduction Our Animals Scientific Name Least

More information

Mate protection in pre-nesting Canada Geese Branta canadensis

Mate protection in pre-nesting Canada Geese Branta canadensis Mate protection in pre-nesting Canada Geese Branta canadensis I. P. JOHNSON and R. M. SIBLY Fourteen individually marked pairs o f Canada Geese were observedfrom January to April on their feeding grounds

More information

Puppy Development. Part One

Puppy Development. Part One Puppy Development Part One Periods of Development Neonatal from birth to two weeks - the puppy is totally dependant on its mother Transitional from two to three weeks- the beginning stages of independence

More information

Territoriality in a snake

Territoriality in a snake Territoriality in a snake Jonathan K. Webb, Mitchell L. Scott, Martin J. Whiting & Richard Shine Behavioral Ecology and Sociobiology ISSN 0340-5443 Volume 69 Number 10 Behav Ecol Sociobiol (2015) 69:1657-1661

More information

DO BROWN-HEADED COWBIRDS LAY THEIR EGGS AT RANDOM IN THE NESTS OF RED-WINGED BLACKBIRDS?

DO BROWN-HEADED COWBIRDS LAY THEIR EGGS AT RANDOM IN THE NESTS OF RED-WINGED BLACKBIRDS? Wilson Bull., 0(4), 989, pp. 599605 DO BROWNHEADED COWBIRDS LAY THEIR EGGS AT RANDOM IN THE NESTS OF REDWINGED BLACKBIRDS? GORDON H. ORTANS, EIVIN RDSKAPT, AND LES D. BELETSKY AssrnAcr.We tested the hypothesis

More information

MITOCW MIT9_20F13_lec17.mp3

MITOCW MIT9_20F13_lec17.mp3 MITOCW MIT9_20F13_lec17.mp3 The following content is provided under a Creative Commons license. Your support will help MIT OpenCourseWare continue to offer high quality educational resources for free.

More information

Evaluation of XXXXXXX mixed breed male dog

Evaluation of XXXXXXX mixed breed male dog Evaluation of XXXXXXX mixed breed male dog Evaluation at Paradise Pet 48 West Passaic Ave - Bloomfield, NJ on April 29, 2013 Conducted by Jeff Coltenback; assisted by Mike Trombetta Video by Diana Coltenback

More information

AnOn. Behav., 1971, 19,

AnOn. Behav., 1971, 19, AnOn. Behav., 1971, 19, 575-582 SHIFTS OF 'ATTENTION' IN CHICKS DURING FEEDING BY MARIAN DAWKINS Department of Zoology, University of Oxford Abstract. Feeding in 'runs' of and grains suggested the possibility

More information

Distribution, population dynamics, and habitat analyses of Collared Lizards

Distribution, population dynamics, and habitat analyses of Collared Lizards Distribution, population dynamics, and habitat analyses of Collared Lizards The proposed project focuses on the distribution and population structure of the eastern collared lizards (Crotaphytus collaris

More information

Supplementary Fig. 1: Comparison of chase parameters for focal pack (a-f, n=1119) and for 4 dogs from 3 other packs (g-m, n=107).

Supplementary Fig. 1: Comparison of chase parameters for focal pack (a-f, n=1119) and for 4 dogs from 3 other packs (g-m, n=107). Supplementary Fig. 1: Comparison of chase parameters for focal pack (a-f, n=1119) and for 4 dogs from 3 other packs (g-m, n=107). (a,g) Maximum stride speed, (b,h) maximum tangential acceleration, (c,i)

More information

AGGRESSION (CATS) DIAGNOSING AND TREATING

AGGRESSION (CATS) DIAGNOSING AND TREATING AGGRESSION (CATS) DIAGNOSING AND TREATING Aggression is a serious and dangerous behavior problem for cat owners. There are many different types of aggression. Making a diagnosis, determining the prognosis

More information

6 Month Progress Report. Cape vulture captive breeding and release programme Magaliesberg Mountains, South Africa. VulPro NPO

6 Month Progress Report. Cape vulture captive breeding and release programme Magaliesberg Mountains, South Africa. VulPro NPO 6 Month Progress Report Cape vulture captive breeding and release programme Magaliesberg Mountains, South Africa VulPro NPO Page Brooder and Incubator room construction 2 Cape Vulture captive bred chick

More information

Class Reptilia Testudines Squamata Crocodilia Sphenodontia

Class Reptilia Testudines Squamata Crocodilia Sphenodontia Class Reptilia Testudines (around 300 species Tortoises and Turtles) Squamata (around 7,900 species Snakes, Lizards and amphisbaenids) Crocodilia (around 23 species Alligators, Crocodiles, Caimans and

More information

Evolution of Birds. Summary:

Evolution of Birds. Summary: Oregon State Standards OR Science 7.1, 7.2, 7.3, 7.3S.1, 7.3S.2 8.1, 8.2, 8.2L.1, 8.3, 8.3S.1, 8.3S.2 H.1, H.2, H.2L.4, H.2L.5, H.3, H.3S.1, H.3S.2, H.3S.3 Summary: Students create phylogenetic trees to

More information

Agenda. Warm-up: Look in your notebook for your grades. Review Notes on Genetic Variation Rat Island. Retake: Monday- last day!!!

Agenda. Warm-up: Look in your notebook for your grades. Review Notes on Genetic Variation Rat Island. Retake: Monday- last day!!! Agenda Warm-up: Look in your notebook for your grades Were you missing any of the assignments? Review Notes on Genetic Variation Rat Island Retake: Monday- last day!!! Gene Pools 1.What makes a species?

More information

Title of Project: Distribution of the Collared Lizard, Crotophytus collaris, in the Arkansas River Valley and Ouachita Mountains

Title of Project: Distribution of the Collared Lizard, Crotophytus collaris, in the Arkansas River Valley and Ouachita Mountains Title of Project: Distribution of the Collared Lizard, Crotophytus collaris, in the Arkansas River Valley and Ouachita Mountains Project Summary: This project will seek to monitor the status of Collared

More information

Coccyzus minor (Mangrove Cuckoo)

Coccyzus minor (Mangrove Cuckoo) Coccyzus minor (Mangrove Cuckoo) Family: Cuculidae (Cuckoos and Anis) Order: Cuculiformes (Cuckoos, Anis and Turacos) Class: Aves (Birds) Fig. 1. Mangrove cuckoo, Coccyzus minor. [http://birds.audubon.org/birds/mangrove-cuckoo,

More information

Sprint speed capacity of two alpine skink species, Eulamprus kosciuskoi and Pseudemoia entrecasteauxii

Sprint speed capacity of two alpine skink species, Eulamprus kosciuskoi and Pseudemoia entrecasteauxii Sprint speed capacity of two alpine skink species, Eulamprus kosciuskoi and Pseudemoia entrecasteauxii Isabella Robinson, Bronte Sinclair, Holly Sargent, Xiaoyun Li Abstract As global average temperatures

More information

May Dear Blunt-nosed Leopard Lizard Surveyor,

May Dear Blunt-nosed Leopard Lizard Surveyor, May 2004 Dear Blunt-nosed Leopard Lizard Surveyor, Attached is the revised survey methodology for the blunt-nosed leopard lizard (Gambelia sila). The protocol was developed by the San Joaquin Valley Southern

More information

Breeding behavior of the boreal toad, Bufo boreas boreas (Baird and Girard), in western Montana

Breeding behavior of the boreal toad, Bufo boreas boreas (Baird and Girard), in western Montana Great Basin Naturalist Volume 31 Number 2 Article 13 6-30-1971 Breeding behavior of the boreal toad, Bufo boreas boreas (Baird and Girard), in western Montana Jeffrey Howard Black University of Oklahoma,

More information

This article is downloaded from.

This article is downloaded from. This article is downloaded from http://researchoutput.csu.edu.au It is the paper published as: Author: A. Wichman, L. Rogers and R. Freire Title: Visual lateralisation and development of spatial and social

More information

BREEDING ROBINS AND NEST PREDATORS: EFFECT OF PREDATOR TYPE AND DEFENSE STRATEGY ON INITIAL VOCALIZATION PATTERNS

BREEDING ROBINS AND NEST PREDATORS: EFFECT OF PREDATOR TYPE AND DEFENSE STRATEGY ON INITIAL VOCALIZATION PATTERNS Wilson Bull., 97(2), 1985, pp. 183-190 BREEDING ROBINS AND NEST PREDATORS: EFFECT OF PREDATOR TYPE AND DEFENSE STRATEGY ON INITIAL VOCALIZATION PATTERNS BRADLEY M. GOTTFRIED, KATHRYN ANDREWS, AND MICHAELA

More information

Texas Quail Index. Result Demonstration Report 2016

Texas Quail Index. Result Demonstration Report 2016 Texas Quail Index Result Demonstration Report 2016 Cooperators: Jerry Coplen, County Extension Agent for Knox County Amanda Gobeli, Extension Associate Dr. Dale Rollins, Statewide Coordinator Circle Bar

More information

ethology international journal of behavioural biology

ethology international journal of behavioural biology ethology international journal of behavioural biology Ethology Tail Autotomy Plays No Important Role in Influencing Locomotor Performance and Anti-Predator Behavior in a Cursorial Gecko Hong-Liang Lu*,

More information

Meal Size Effects on Antipredator Behavior of Hatchling Trinket Snakes, Elaphe helena

Meal Size Effects on Antipredator Behavior of Hatchling Trinket Snakes, Elaphe helena Ethology Meal Size Effects on Antipredator Behavior of Hatchling Trinket Snakes, Elaphe helena Rita S. Mehta Department of Biology, University of Texas, Tyler, TX, USA Correspondence Rita S. Mehta, Department

More information

Effects of Natural Selection

Effects of Natural Selection Effects of Natural Selection Lesson Plan for Secondary Science Teachers Created by Christine Taylor And Mark Urban University of Connecticut Department of Ecology and Evolutionary Biology Funded by the

More information

Diversity of Animals

Diversity of Animals Classifying Animals Diversity of Animals Animals can be classified and grouped based on similarities in their characteristics. Animals make up one of the major biological groups of classification. All

More information

MA41 Colour variability and the ecological use of colour in the chameleons and geckos of Mahamavo

MA41 Colour variability and the ecological use of colour in the chameleons and geckos of Mahamavo MA41 Colour variability and the ecological use of colour in the chameleons and geckos of Mahamavo Colour and the ability to change colour are some of the most striking features of lizards. Unlike birds

More information

Unit 3 Sustainability and interdependence Sub Topic 3.4: Animal welfare

Unit 3 Sustainability and interdependence Sub Topic 3.4: Animal welfare Unit 3 Sustainability and interdependence Sub Topic 3.4: Animal welfare Page 1 of 12 On completion of this topic I will be able to: Describe the costs, benefits and ethics of providing different levels

More information

ENVIRONMENT GROUP NEWS Issue 17 May 2014

ENVIRONMENT GROUP NEWS Issue 17 May 2014 EPIPHANY 2006 EARTHQUAKE Many of you may remember the large earthquake that took place during the day on the 8 th Jan 2006. The earthquake was of magnitude 6.9 and was felt over quite a wide distance and

More information

Lab 7. Evolution Lab. Name: General Introduction:

Lab 7. Evolution Lab. Name: General Introduction: Lab 7 Name: Evolution Lab OBJECTIVES: Help you develop an understanding of important factors that affect evolution of a species. Demonstrate important biological and environmental selection factors that

More information

ANS 490-A: Ewe Lamb stemperament and Effects on Maze Entry, Exit Order and Coping Styles When Exposed to Novel Stimulus

ANS 490-A: Ewe Lamb stemperament and Effects on Maze Entry, Exit Order and Coping Styles When Exposed to Novel Stimulus Animal Industry Report AS 663 ASL R3182 2017 ANS 490-A: Ewe Lamb stemperament and Effects on Maze Entry, Exit Order and Coping Styles When Exposed to Novel Stimulus Emily Strong Iowa State University Samaneh

More information

ESWDA. Police Service Test

ESWDA. Police Service Test ESWDA Police Service Test To obtain a Police Service Dog Certification the handler and dog (hereafter referred to as the K-9 team) will be tested in all phases of this test. The following areas to be tested

More information

Sheikh Muhammad Abdur Rashid Population ecology and management of Water Monitors, Varanus salvator (Laurenti 1768) at Sungei Buloh Wetland Reserve,

Sheikh Muhammad Abdur Rashid Population ecology and management of Water Monitors, Varanus salvator (Laurenti 1768) at Sungei Buloh Wetland Reserve, Author Title Institute Sheikh Muhammad Abdur Rashid Population ecology and management of Water Monitors, Varanus salvator (Laurenti 1768) at Sungei Buloh Wetland Reserve, Singapore Thesis (Ph.D.) National

More information

Connecticut Police Work Dog Association

Connecticut Police Work Dog Association Connecticut Police Work Dog Association Certification Test Standards The following test standards have been adopted by the Connecticut Police Work Dog Association, hereinafter referred to as the CPWDA.

More information