A comparison of display behavior between two divergent anolis carolinensis populations

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(1)LRH: N. BLOCH AND D.J. IRSCHICK RRH: DISPLAY BEHAVIOR AND DIVERGENT ANOLIS POPULATIONS. A COMPARISON OF DISPLAY BEHAVIOR BETWEEN TWO DIVERGENT ANOLIS CAROLINENSIS POPULATIONS.. Natasha Bloch1* and Duncan J. Irschick2. 1. Department of Biological Science. Carrera 1 # 18A-10 Universidad de los Andes Bogota, Colombia.. 2. Department of Ecology and Evolutionary Biology. 310 Dinwiddie Hall Tulane University New Orleans, LA 70118. *Corresponding author: Phone: (504) 862-8295, FAX: (504) 862 8706, e-mail: [email protected].

(2) TABLE OF CONTENTS. INTRODUCTION...........................................................................................................5 MATERIALS AND METHODS ...................................................................................10 RESULTS .....................................................................................................................13 DISCUSSION ...............................................................................................................18 REFERENCES..............................................................................................................25 ACKNOWLEDGMENTS .............................................................................................33 APPENDIX...................................................................................................................34. 2.

(3) LIST OF FIGURES. Figure 1: (A) Tulane University (TU) habitat and (B) Exxon (EX) habitat. ......................8 Figure 2: Anolis carolinensis lizard displays exposing its dewlap. ...................................9 Figure 3: Distribution of A, B and C displays for the Tulane University (TU) population (. ), and for the Good Hope Field (GH) population ( ). ...............................................15 Figure 4: Distribution of the number of displays per volley for the Tulane University (TU) population ( ), and for the Good Hope Field (GH) population ( ). ......................16. 3.

(4) LIST OF TABLES. TABLE 1: Loadings from a principal components analysis for various display variables. Substantial loadings are in bold. ....................................................................................13 TABLE 2: Summary of the Analyses of Variance (MANOVA) comparing Good Hope Field and Tulane anole populations for the principal components analyses factors.........14 TABLE 3: Mean (+ 1 SE) values for various display variables in the Tulane and Good Hope Field populations..................................................................................................17. 4.

(5) INTRODUCTION. The way sexual selection shapes the social behavior and mating systems in many vertebrates is strongly influenced by habitat (Anderson, 1994; Wade, 1995; Baird et al., 2003). Many features of the habitat can potentially affect social behavior and mating systems, including habitat visibility, predation risk, the availability of food, temperature, and many other factors (McCoy et al. 1995). Display behavior is an important aspect of the social behavior of many animal species. Indeed, the ability of both males and females of many different species to effectively display to conspecifics, other species, and predators, is critically important for survival and reproduction. Animals signal in many different ways, but some well-studied systems include bird song (Marten & Marler, 1977; Marten et al., 1977; Wiley and Richards, 1982), vocalizations in frogs (Ryan, 1985; Ryan 2003a; Ryan, 2003b; Wilczynski et al., 2001; Bosch et al., 2002), and visual displays in lizards (Whiting et al. 2003). How habitat affects the display behavior of animals is often complex. One key factor is habitat visibility (how far can a visual signal be detected by receivers), which should have a strong impact on both the form and intensity of visual signals, and could also affect how much time animals spend transmitting such signals (Luyten & Liley, 1991). For example, in an open habitat where animals can easily see another, males may invest significantly more time displaying compared to a more cluttered habitat in which densities are similar.. Another factor is whether the available habitat is distributed. continuously, or rather arrayed in discrete clumps that are separated from one another (Zamudio and Sinervo 2003). Such different habitat arrangements can profoundly affect. 5.

(6) the potential distribution of mates and resources, and consequently, the ability of males to monopolize those resources (Emlen & Oring, 1997; Baird et al., 1997). Indeed, discrete clumps of habitat may be especially pre-disposed towards highly territorial (“despotic”, see Zamudio and Sinervo 2003) males, because of their ability to effectively monopolize resources (Orrell & Jenssen, 2003; Zamudio and Sinervo 2003). Of course, such factors are not necessarily independent of one another, as small, discrete clumps of habitat (e.g., a large rock) may have higher visibility compared to a larger, more cluttered habitat. A third important factor on the evolution of display behavior is the relative risk of predation. For example, classic studies with tungara frogs and bats show that male frogs are forced to compromise between calling to enhance the probability of reproduction and not calling to avoid being consumed (Endler 1983; Ryan et al., 1982; Ryan 1985). Therefore, comparisons of display behavior in different populations of the same species that occupy habitats varying in such characteristics might shed light on these issues. Many lizard species possess colorful expanded throatfans (i.e., dewlaps), and badges on their ventral surfaces that they use to signal towards conspecifics and/or predators (Whiting et al. 2003). Further, lizard species and populations vary dramatically in the size and color or their dewlaps, and each species has a distinct pattern of elaborate visual displays (i.e., head-bobbing, throat extensions). Because lizard populations within the same species often differ in habitat use, they form an excellent model system for understanding the influence of habitat use on the evolution of visual displays. Here, we examined two populations of the well-studied green anole lizard (Anolis carolinensis) to understand the influence of habitat structure on display behavior.. 6.

(7) Green anoles are highly sexually dimorphic (adult males average about 1.5 times the body size of adult females), and adult males will actively defend territories. Green anoles have highly stereotyped headbob display forms: both males and females have a repertoire of displays termed A, B and C patterns that are highly conserved within populations (Jenssen, 1977; Lovern et al. 1999; Lovern & Jenssen, 2001; Orrell & Jenssen, 2003; Lovern & Jenssen, 2003).. These displays consist of stereotyped. sequences of head bobbing (vertical displacements of the head), which are generally accompanied by dewlap extensions. Other display modifications that enhance visibility can also accompany these displays (Jenssen, 1977). These displays are used to defend territories, attract females, and signal to predators (Ruby, 1984; Jenssen et al. 1995). Previous studies of green anole display behaviour have focused primarily on interpopulation differences in display structure (Jenssen & Gladson, 1984; Lovern et al., 1999; Macedonia & Clark, 2003), or in dewlap size (Losos & Chu, 1998), but we are aware of no studies that have investigated how inter-populational variation in habitat use affects overall display behavior. We compared two divergent populations of green anole lizards in southern Louisiana with the purpose of comparing their display behaviour. These two populations are highly divergent in habitat use, morphology, performance, and escape behavior (Irschick et al, in review), but no study has examined whether these two populations also differ in important aspects of their display behaviour. One site (Good Hope Field) is a freshwater swamp composed of a dense and continuous array of relatively tall bushes and trees (Figure 1A). This site contains a healthy Southern Louisiana freshwater swamp community, including large numbers of potential predators, such as herons and snakes.. 7.

(8) By contrast, the second site (Tulane University Campus) is composed of discrete clumps of simple, broad leaves (Figure 1B). Because the Tulane site is present within the center of the city of New Orleans, the site lacks the numerous potential predators, such as herons and snakes. Because these sites are fairly close to one another (within 40 km), and thus are quite closely related, they offer an excellent opportunity to compare how variation in habitat structure affects display behavior.. Figure 1: (A) Tulane University (TU) habitat and (B) Exxon (EX) habitat.. A. B. We predicted that, based on the highly clumped nature of the Tulane University campus habitat compared to the Good Hope Field habitat, intraspecific competition would be more extreme in the former habitat. Consequently, because strong intrasexual selection often results in higher display rates (Shine, 1979; Stamps, 1983; Hews 1990) 8.

(9) enhanced courtship behavior (Halliday, 1983), and enhanced territorial defense (Arak 1983; Stamps 1994) one might predict overall higher display rates in the Tulane University campus. Further, males in the two populations may also differ in their relative use of different kinds of displays (A, B, or C) in the two habitats, although the exact manner by which they would differ is difficult to predict. Therefore, we videotaped displays of adult males from both populations during the peak of the breeding season (April and May) in both habitats and asked whether these populations differ in basic aspects of display behavior (i.e., percent time displaying, percent use of display types, percent use of display modifiers, etc.).. Figure 2: Anolis carolinensis lizard displays exposing its dewlap.. 9.

(10) MATERIALS AND METHODS. Field sites and videotaping: We compared populations from two different sites in Louisiana: one from an urban setting at Tulane University in New Orleans, and the other from Good Hope Field in St Charles Parish, Southeastern Louisiana (both described in Irschick et al. in review). The habitat structure of the Tulane campus consisted of isolated clumps of vegetation separated in space from other clumps by 2-10 m along a 380 m road (Figure 1A). Tulane habitat is artificially maintained by Tulane workers (e.g., regular watering via a sprinkler system, occasional removal of dying plants), although lizards are generally undisturbed (Irschick, pers. obs.). Our second site is a 775 m transect that consists of a rarely used dirt access road bordered on either side by relatively thin continuous strips of vegetation (about 3-4 m on each side). On either side of each strip of vegetation is an open-water swamp. The vegetation at Good Hope Field is a mixture of low-lying shrubs and grass interspersed with larger trees and bushes (Figure 1B). To compare these populations, we gathered video focal data for 30 and 26 large adult males (> 60 mm SVL approximately) from the Tulane Campus and the Good Hope Field site, respectively. We videotaped individual males under free-ranging conditions for a minimum of 5 minutes and a maximum of 40 minutes (average duration 15 minutes ± 1.32). We used a Sony Handicam camera with mini-DV digital cassettes and a tripod to film the lizards. We maintained a distance of at least 2 m during filming and remained as still as possible to avoid startling the lizard. Only one investigator (Bloch) videotaped and analyzed all the video focals to ensure consistency. The study was conducted from April 1st 2004 to May 20th 2004 when males are actively attempting to acquire and defend. 10.

(11) territories. We only filmed lizards during warm (i.e., shaded ambient temperatures > 26° C) and non-rainy weather.. Video analyses: We analyzed the focal videos by reviewing the videotapes frame by frame (30 frames/s) to identify specific behaviors. Based on previous descriptions of Anolis carolinensis displays (DeCourcy & Jenssen, 1994. Jenssen et al.,2000; Lovern et al., 1999) we quantified different standardized behaviors from each videotape: (1) Bobbings, or vertical head bobbings movements of high amplitude while extending both front limbs; (2) quick bobs, or. shallow simple and double bobs, also called. “shudderbobs” in previous studies (Tinkle, 1967; Orrell & Jenssen, 2003); (3) The number of times the lizard extended its dewlap. We counted the number of times each of these behaviours was repeated in tandem and the duration of each tandem (by noting the exact moment [minute, second and frame] the first repetition started and the last one ended). We also recorded the display type according to the three stereotyped types of display A, B and C (Jenssen 1977; DeCourcy & Jenssen, 1994) . Lizards can alter stereotyped displays with modifiers that previous researchers (Lovern et al. 1999; Jenssen, 1977) believe are used to enhance visibility. For the present study, display modifiers are defined as quick bobs after the core component of the display (see further description in Jenssen 1977) and as “pushups” which are a full extension of all four limbs during the display. We then compared the frequency of display modifiers in the two habitats. Finally we recorded if each display was part of a volley or done singly. 11.

(12) (Displays in the same volley had to be less than 2 seconds apart following techniques used by Orrell and Jenssen, 2003). We compared the dewlap areas of adult males from both populations. To do this we took digital pictures of 20 adult males (>65mm SVL) for each site, pulling their dewlaps out with forceps and then digitizing the pictures using tpsdig version 1.40 to calculate the dewlap area. Data analysis: We calculated five primary variables for each focal video (1) Percentage of time displaying (2) Percentage of time exposing the dewlap, (3) average duration of each display, (4) total bobbings duration, (5) total quick bobs duration. We used a Principal Components Analyses to reduce the dimensionality of these variables and then performed a multivariate ANOVA (Analyses of Variance) using populations as a primary factor to compare the result between the two populations. We then calculated the relative proportions of the A, B and C displays for each study site (we first estimated these proportions for each individual and then evaluated the mean value for each study population); we then compared the two populations using a chisquare test. We used a Kolmogorov -Smirnov test to compare the volley length distributions in both populations. We also evaluated the percentage of displays done in volleys and finally calculated the percentage of displays performed with either kind of visibility enhancement modifier and compared them between our two study site. We used a non-parametric Kruskal-Wallis test to compare the percentages stated above between populations. All data analyses were performed using SYSTAT (version 10, SPSS 2000) and all errors are show as Standard errors.. 12.

(13) RESULTS The average dewlap areas 2.21 ± 0.096 cm2 and 2.29 ± 0.097 cm2 for TU and GH respectively, do not differ statistically after conducting a one-way ANOVA (P>0.5). PC 1 showed high positive loadings for the percentage of displaying, percentage of bobbing and percentage of quick bobs. PC 1. accounts for a variance of 56.8 %. (Eigenvalue = 2.840). PC 2 has very strong and positive loadings for the percentage of time dewlaping and the average duration of each display. PC 2 explains 36.3 % of the variance (Eigenvalue = 1.815).. Therefore, lizards with longer displays expose their. dewlaps for longer.. TABLE 1: Loadings from a principal components analysis for various display variables. Substantial loadings are in bold. Variables. PC1. PC2. % Display. 0.962. 0.247. % Dewlap. 0.135. 0.931. % Bobbings. 0.967. 0.124. % Quick bobs. 0.941. 0.270. Average display duration (s). 0.272. 0.894. Eigenvalue. 2.840. 1.815. The multivariate ANOVA for the Principal components shows that the populations differ significantly in the percentage of time displaying (F-value= 13.490; DF=1,54; P < 0.001).. 13.

(14) Lizards from the TU population spend almost twice as much the time displaying compared to GH lizards (9.31 ± 0.84 compared to 5.19 ± 0.83. Both study populations did not differ in the average duration of each display (F-value= 0.019; DF=1,54; P> 0.5).. TABLE 2: Summary of the Analyses of Variance (MANOVA) comparing Good Hope Field and Tulane anole populations for the principal components analyses factors. Variable. F-value. DF. p-value. PC1. 13.490. 1,54. 0.001. PC2. 0.019. 1,54. 0.890. Wilks' Lambda. 6.64. 2,53. 0.003. We also compared the display ratios (relative proportions of A, B and C) between both populations (Fig.3). We obtained a ratio of 2:5:18 for TU and of 1:3:16 for GH. A chisquare test showed significant differences between both of these ratios (P<0.001; DF=2; X2= 26.46), lizards from GH seem to be using more B displays than expected.. 14.

(15) Figure 3: Distribution of A, B and C displays for the Tulane University (TU) population (. ), and for the Good Hope Field (GH) population ( ).. Lizards from both populations displayed in volleys of 2 to 8 displays and the average volley lengths were 1.95 and 2.00 for TU and GH population respectively. When we compared the volley length distribution (Fig.2) for our study populations, we obtained significant differences between them (dmax=0.174; df=9; P<0.001). This KS tests means that there is a significant tendency of Good Hope field anoles to perform longer volleys.. 15.

(16) Figure 4: Distribution of the number of displays per volley for the Tulane University (TU) population ( ), and for the Good Hope Field (GH) population ( ).. By estimating the percentage of displays done in volleys (as opposed to singly) we observed that lizards perform 79.7% and 76.4% of the displays in volleys at TU and GH respectively. The Kruskal-Wallis test showed no difference between both these percentages (Mann-Whitney U test statistic = 257.0; P=0.782; df=1). However when we compared the percentage of displays with visibility modifiers we observed a significant difference between both populations (P=0.001; df=1): Lizards from TU use these kinds of modifiers 8.98% of the time while lizards from GH only use them 3.15% of the time.. 16.

(17) TABLE 3: Mean (+ 1 SE) values for various display variables in the Tulane and Good Hope Field populations. Variable. Tulane. Good Hope Field. % time displaying. 9.31 ± 0.84. 5.19 ± 0.83. % time dewlaping. 61.60 ± 2.07. 56.34 ± 3.59. % time headbobs. 3.26 ± 0.32. 1.91 ± 0.29. % time quickbobs. 5.93 ± 0.59. 3.09 ± 0.52. Average display duration (s). 4.34 ± 0.18. 3.39 ± 0.22. 17.

(18) DISCUSSION. By comparing two Anolis carolinensis populations with very distinct habitats we found that (1) Dewlap areas do not differ, thus indicating that these two populations have diverged in display behavior, but not in the size of the display structure. (2) Tulane and Good Hope field anoles differ greatly in the percentage of time they spend displaying: Tulane anoles display almost twice as often as Good Hope field anoles even if the average duration of a display is similar for both these populations. (3) Both study populations differ in the relative frequency of display types (4) Lizards from TU use display modifiers significantly more often than the lizards from GH (5) Both populations also differ in volley length. Previous studies (Irschick et al., in review) have already shown that these two populations differ significantly in habitat structure, and here we show that they also differ in their display behaviour. The amount of time spent displaying is often viewed as a good indicator of the investment that males contribute towards territorial defense (ref). Previous research shows that highly polygynous anoles, like many trunk-ground anoles, display more than 5 % of their time, whereas largely non-polygynous species, spend only about 1% or less (Irschick & Losos, 1996).. For the particular case of Good Hope field anoles the. percentage of display, 5.19 % ± 0.83, is similar to the one reported by Irschick & Losos (1996) for trunk-ground and trunk-crown anoles (5.7±1.2 and 6.0±1.4 respectively) but much higher than the ones reported for twig anoles (1.3 ± 0.002) or trunk anoles (2.0 ± 0.8). Even if the data we found for GH is much lower than the one for TU, 9.31 % ± 0.84, this value is still high according to the data available for the display rates of anoles. The. 18.

(19) percentage of time displaying found for Tulane anoles is extremely high, almost double as the ones reported for trunk anoles, which are themselves the highest ones among anoles (Irschick & Losos, 1996). The difference in the amount of time displaying we observed between our study populations (9.31 % ± 0.84 for TU and 5.19 % ± 0.83 for GH) is considerably high and spans approximately the same range as polygynous and non-polygynous species. The capacity for a male of any species to monopolize females might determine the intensity with which male’s signals, assuming all the costs this carries. The tremendous difference in percent time displaying between green anole populations can be explained by understanding the difference in the potential for sexual selection in the two environments (concept used by Baird et al., 1997). At Tulane University the lizard density is very high: demographic studies have found that within a 3x2 m patch, over 30 animals were captured, including about 10 adult males (Irschick, in review). The high densities in combination with the clumped nature of the habitat make for an especially stressful situation for males, which are exposed to stronger intrasexual competition (McCoy et al., 2003; Baird & Sloan, 2003; Baird et al., 1997; Stamps & Losos, 1997). Social behaviour is usually affected by population densities as seen in previous studies on bats, which change the frequency, timing and intensity of their calls when they are in groups to maintain contact and obtain information about each other (Fenton, 2003). Metzner (1999) observed the same phenomenon in electrolocating fish. Furthermore in his study of collared lizards, Baird (1997) found that as male competition intensifies, the amount of sexual signals increase too. But, besides population densities, the environment’s potential for polygyny (Anderson, 1994; Wade, 1995) could also. 19.

(20) contribute to the differences observed in the display rates of both populations. The concept of environment’s potential for polygyny refers to the potential for a male to monopolize many females due to habitat structure, as in a polygynous system. In their study of sac-winged bats, Saccopteryx bilineata,Voigt and Helversen (1999), found that males hovering display behaviour varies greatly with males capacity to monopolize females. In this species of bats males defend harem territories where females gather and increase the number of hovering displays as the harem size gets bigger. As a consequence of this, males S. bilineata spend a substantial part of their daily energy budget to defend a harem territory. The small clumps of vegetation at Tulane are likely to make it easier for an adult male to monopolize many females compared to GH, therefore increasing the effects of polygyny and intrasexual selection (according to McCoy’s (1995) conclusions from the study of Crotaphytus collaris collaris. On the other hand, we can expect that TU females concentrate on the clumps of available habitat making it possible for males to monopolize more females therefore increasing the habitat’s potential for polygyny (Ruby, 1984; Jenssen et al., 1995; Zamudio & Sinervo, 2003). However an alternative hypothesis is that continuous habitats favor female’s capacity to assess more than one male and consequently are expected to have stronger intrasexual contests (Baird et al., 1997). According to this theory lizards at GH should display more frequently than lizards at TU. Nevertheless display behaviour is a result of various selective mechanisms that could be pulling in different directions (McCoy et al., 2003; Arnold, 1983). Display behaviour in other animals, as well as for anoles, is a compromise between attracting mates, defending territories, and avoiding predators. The Túngara frog, studied by Mike Ryan (1982, 1985), is an excellent example: males must call, but. 20.

(21) also risks being eaten by bats that also respond to the call. Predation risk may therefore act as a counterselective mechanism by increasing the costs of signaling, and thus imposing a dilemma in all animal species from birds to snails (Heindl & Winkler, 2003; Kilpimaa, 2003; Lewis, 2001; Semler, 1971; Endler, 1980, 1983; Stoner and Breden, 1988; Stoddard, 1999; Zuk, 1998). Because Good Hope field is a swamp population, the abundance and diversity of predators is probably higher compared to Tulane, which is an urban population. Large numbers of bird, invertebrate and snake potential predators for green anoles have been observed at GH while absent at TU (Irschick pers. obs.) Our results are consistent with this observation since GH anoles showed significant lower display rates than urban TU anoles. Nevertheless we have to be very cautious when using differences in predation pressures between TU and GH to justify the observed differences in display rates; Further studies evaluating the density and diversity of predators in both sites are still necessary to confirm our view on this matter. When different populations of the same species have to adapt to different environments they can either diverge in their morphology or their behaviour. Previous studies proved that both possibilities are observed in nature: Plovers vary in the melanization of their plumage according with habitat light conditions (Bokony et al., 2003) and Bowerbirds diverge in their crest areas (Madden et al., 2004). On the other hand animals, like emballonurid bats (Ibanez et al., 2004), show differences in their display behaviour when living in habitats with different structures. We found no difference in the dewlap areas of TU and GH anoles, thus signal size is conserved across populations, but the signal behaviour has evolved. These findings are surprising considering that Irschick (in press). 21.

(22) found significant morphological differences in limb length and toepad area between anoles from TU and GH as a response to different habitats. The Anolis carolinensis repertoire is highly conserved across populations from different states in North America and Hawaii (Lovern et al., 1999). Here we found that green anoles from both Louisiana populations also used the previously described A, B and C display types in different proportions. GH anoles used proportionally less A and B displays and more C displays than TU anoles. Our results are consistent with previous studies on the display behaviour of green anoles that showed than even if the structure of the display itself is highly conserved, the display use is variable among populations and social contexts (DeCourcy & Jenssen, 1994; Jenssen et al., 2000; Lovern & Jenssen 2003). DeCourcy & Jenssen (1994) recommended that display patterns should not be associated with specific social contexts after males from their study were shown to use all display types for every social context. Only a few generalizations have proved to be reliable. First, with regard to the frequency of use of A, B and C displays is that it differs between sexes, that males use predominantly C displays whereas females used predominantly A and B (Orrell & Jenssen, 2003). And second that males tend to use more C displays when signaling towards individuals at a distance but increase the proportion of As and Bs during encounters at a shorter distance (Orrell & Jenssen, 2003). This could explain that TU anoles, where resources are clumped and individuals exist in higher densities, use larger proportion of A and B displays. However more studies are necessary to be certain of any explanation for the differences in the ratios of As, Bs and Cs use between TU and GH population.. 22.

(23) Tulane anoles use modifiers to enhance the visibility of the display 8.98% of the time, which is significantly more than GH anoles, which only use them 3.15% of the time. This could also be a response to lower predation risk at a urban location. It could also be an adaptation to the structure of the Tulane habitat. As noted earlier, Tulane habitat is formed almost exclusively of plants (Aspidistra elatior) that do not grow very high above the ground (Irschick et al. in review). Shorter plants may give lizards a better visibility, which would make the use of modifiers profitable for males. Even if all the previously reported green anole displays were observed in our study populations, 4 out of the 30 adult male lizards videotaped at the TU site appeared to be doing new variants of the C display, which does not fit into any of the previously described patterns. Lovern & Jenssen (2003) described a type of display exclusive to juveniles that he called X display. This deviant behaviour could be the result of some juvenile display that was not abandoned and fixed as a form of “baby talk” or could be a completely different display variant. Because previous studies on the green anole repertoire were laboratory experiments, further studies should be conducted in the field to reveal the function and context of this new display variant. Finally we found that even if anoles in both populations have similar percentages of displays in volleys, GH anoles tend to perform longer volleys more frequently. Previous studies show that, during male-male signaling, the larger the distance between displaying males the longer the volleys and that males tend to perform singly or in shorter volleys when closer to each other (DeCourcy & Jenssen, 1994; Orrell & Jenssen, 2003). This might be a result of the risk increase, as competitors get closer to each other. Since males. 23.

(24) at TU, are present in higher densities and hence, are closer to each other, which is consistent with the above-mentioned observations. As an alternative explanation for the use of longer volley by GH anoles we could concentrate on GH field habitat structure. This habitat is composed of thin strips of dense vegetation considerably high that could block visual signals and therefore decrease the habitat’s visibility. Good Hope field of trees and higher bushes could diminish the distance at which a signal can be perceived. If the male’s purpose for performing in longer volleys were to make sure that the signal reaches its receivers (Fleishmann, 1988), it could be al alternative explanation for the use of longer volleys by GH males.. 24.

(25) REFERENCES. Anderson, M. 1994. Sexual selection. Princeton University press, Princeton, N.J. Arak, A.1983. Male-male competition and mate choice in anuran amphibians. In Mate choice, edited by P. Bateson, 3-32. Cambridge University Press, Cambridge, U.K. Arnold, S.J. 1983. Sexual selection: the interface of theory and empiricism. In Mate choice, edited by P. Bateson, 67-107. Cambridge University Press, Cambridge, U.K. Baird, T.A & Sloan, C.L. 2003. Interpopulation Variation in Social Organization of Female Collared lizards, Crotaphytus collaris. Ethology 109:879-894. Baird, T.A., Timanus, D.K. & Sloan, C.L. 2003. Intra- and intersexual Variation in Social Behavior. Effects on Ontogeny, Phenotype, Resources, and Season. In Lizard Social Behavior. Edited by Stanley F. Fox , J. Kelly McCoy and Troy A Baird. The Johns Hopkins University press. Baltimore. 7-46. Baird, T.A., Fox, S.F. & McCoy, J.K., 1997. Population differences in the roles of size and coloration in intra- and intersexual selection in the collared lizard, Crotaphytus collaris: influence of habitat and social organization. Behavioral Ecology 8(5):506517. Bosch, J., Rand, A.S.& Ryan, M.J. 2002. Response to variation in chuck frequency by male and female Túngara frogs. HERPETOLOGICA, 58 (1): 95-103 Bokony, V., Liker, A., Szekely, T. and Kis, J. 2003. Melanin-based plumage coloration and flight displaysin plovers and allies. Proceedings of the Royal Society of London B. 270:2491-2497.. 25.

(26) Davies, N.B. 1992. Dunnock behavior and social evolution. Oxford: Oxford University Press. DeCourcy, K.R. & Jenssen, T.A. 1994. Structure and use of male territorial headbob signals by the lizard Anolis carolinensis. Animal Behavior. 47:251-262. Emlen, S.T & Oring, L.W. 1997. Ecology, sexual selection and the evolution of mating systems. Science. 197:215-223. Endler, J.A.1980. Natural Selection on color patterns in Poecilia reticulate. Evolution 34:76-91. Endler, J.A. 1983.Natural and sexual seleccion on color patterns in poeciliid fishes. Environ. Biol Fish. 9:173-190. Fenton, M.B. 2003. Eavesdropping on the echolocation and social calls of bats. Mammal. Review 33(3):193-204. Fleishmann, L.J. 1988. Sensory influence in physical design of a visual display. Animal Behavior. 36:1420-1424. Fleishmann, L.J. 1992. The influence of the sensory system and the environment on motion patterns in the visual displays of anoline lizards and other vertebrates. American Naturalist 139:S36-S61. Grant, J.W.A. 1993. Whether or not to defend? The influence of resource distribution. Mar. Behav. Physiology. 23:137-153. Goldberg, J.L., Grant, J.W.A. & Lefebvre, L. 2001. Effect of the temporal predictability and spatial clumping of food on the intensity of competitive aggression in the Zanaida dove. Behavioral Ecology. 12:490-495.. 26.

(27) Gross, M.R. 1996. Alternative reproductive tactics: diversity within the sexes. Trends Ecol. Evol., 11:92-98. Halliday,T.R. 1983. The story of mate choice. In Mate choice, edited by P. Bateson, 3-32. Cambridge University Press, Cambridge, U.K. Hews, D.K. 1990. Examining hypothesis generated by field measures of sexual selection on male lizards, Uta palmeri. Animal Behavior 46:279-291. Heindl, M. & Winkler, H. 2003. Interacting effects of ambient light and plumage color patterns in displaying Wire-tailed Manakins (Aves, Pipridae). Behavioral Ecology and Sociobiology. 53:153-162. Ibañez, C., Juste, J., López-Wilchis, R. & Núñez-Garduño, A. 2004. Habitat variation and jamming avoidance in echolocation calls of the sac-winged bat (Balantiopteryx plicata). Journal of Mammalogy 85(1):38-42. Irschick, D.J. & Losos, J.B. 1996. Morphology, ecology, and behavior of the Twig Anole, Anolis angusticeps. Jenssen, T.A. 1977. Evolution of Anoline Lizard Display Behavior. American Zoologist 17:203-215. Jenssen, T.A. & Gladson, N.L. 1984. A comparative display analysis of the Anolis brevirostris complex in Haiti. Journal of Herpetology 18:217-230. Jenssen, T.A. & Nuñez, S.C. 1998. Spatial and breeding relationships of the lizard Anolis carolinensis: evidence of intrasexual selection. Behaviuor 135:981-1003. Jenssen, T.A., Greenberg, N. & Hovde, K.A. 1995. Behavioral profile of free-ranging male lizards, Anolis carolinensis, across breeding and non breeding seasons. Herpetological Monographs 9:41-62.. 27.

(28) Jenssen, T.A., Orrell, K.S. & Lovern, M.B. 2000. Sexual dimorphism in aggressive signal structure and the use by a polygynous lizard, Anolis carolinensis. Copeia. 2000:140149. Kwiatkowski, M.A. & Sullivan, B.K. 2002. Mating system structure and population density in a polygynous lizard, Sauromalus obesus. Behavioral ecology. 13:201208. Kilpimaa, J., Alatalo, R.V. and Siitari, H. 2004. Trade-offs between sexual advertisement and immunefunction in the pied flycatcher ( Ficedula hypoleuca). Proceedings of the Royal Society of London B. 271:245-250. Lott, D. 1991. Intraspecific variation in the social systems of wild vertebrates. Cambridge: Cambridge University Press. Losos, J.B. & Chu, L-R. 1998. Examination of factors Potentially Affecting Dewlap size in Caribbean Anoles. Copeia 1998 (2):430-438. Luyten, P.H. & Liley, N.R. 1991. Sexual selection and competitive mating success of male guppies (Poecilia reticulate) from four Trinidad populations. Behavioral ecology and Sociobiology. 29:133-138. Lovern M.B., Jenssen T.A., Orrell K.S., Tuchak T. 1999 Comparisons of temporal display structure across contexts and populations in male Anolis carolinensis: Signal stability or lability? Herpetologica55 (2): 222-234 Lovern M.B. & Jenssen T.A. 2001. The effects of context, sex and body size on staged social interactions in juvenile mele and female green anoles (Anolis carolinensis). Behaviour 138:1117-1135.. 28.

(29) Lovern M.B., McNabb, F.M.A., & Jenssen T.A. 2001. Developmental effects of testosterone on behavior in male and female green anoles (Anolis carolinensis). Hormones and Behavior 39:131-143. Lovern M.B., Jenssen T.A. 2003. Form Emergence and fixation of head bobbing displays in the green anole lizard (Anolis carolinensis): A reptilian model of signal ontogeny. Journal of comparative Psychology 117(2):133-141. Lewis, B.D. 2001. Trade-off between growth and survival: Response of freshwater snails to predacious crayfish. Ecology 82(3):758-765. Macedonia, J.M. & Clark, D.L. 2003. Headbob display structure in the naturalized Anolis lizards of Bermuda: Sex, context, and population effects. Journal of Herpetology 37(2):266-276. Madden, J.R., Endler, J.A. & Jury, F. 2004. Morphological signals of sex and status in spotted bowerbirds. Emu 104:21-30. Marten, K. & Marler, P. 1977. Sound transmission and its significance for animal vocalization. I. Temperate habitats. Behavioral Ecology and Sociobiology 2: 271– 290. Marten, K., Quine, D. & Marler, P. 1977. Sound transmission and its significance for animal vocalization. II. Tropical forest habitats. Behavioral Ecology and Sociobiology 2:291–302. McCoy, J.K., Fox, S.F. & Braid, T.A. 2003. Sexual Selection, Social Behavior, and the environmental Potential for Polygyny. In Lizard Social Behavior. Edited by Stanley F. Fox , J. Kelly McCoy and Troy A Baird. The Johns Hopkins University press. Baltimore. 149-171.. 29.

(30) McCoy, J.K. 1995. Mechanisms of selection for the evolution of sexual dimorphism in the collared lizard (Crotaphytus collaris). Ph.D dissertation, Oklahoma State University, Stillwater. Metzner, W. (1999) Neural circuitry for communication and jamming avoidance in gymnotiform electric fish. Journal of Experimental Biology, 202, 1365–1375. Moore, M.C. & Thompson, C.W. 1990. Field endocrinology of reptiles: Hormonal control of alternative male reproductive tactics. In Progress in comparative endocrinology, edited by A. Epple, C.G. Scanes, and M.H. Stetson. 685-690. Wiley-liss, New York. Orrell, K.S. & Jenssen. T.A. 2003. Heterosexual signaling by the lizard Anolis carolinensis, with intersexual comparisons across contexts. Behaviour 140:603:634. Ruby, D.E.1984. Male breeding success and differential access to females in Anolis carolinensis. Herpetologica 40(3):272-280. Ryan, M.J., Tuttle, M.J. & Rand, R.S. 1982. Bat predation and sexual advertisement in a neotropical anuran. American Naturalist 119:136-139. Ryan, M.J. 1985. The Túngara frog: a study of sexual selection and communication. University of Chicago Press, Chicago. Ryan, M.J.& Rand, A.S. 2003a. Sexual selection in female perceptual space: How female Túngara frogs perceive and respond to complex population variation in acoustic mating signals. EVOLUTION, 57 (11): 2608-2618 Ryan, M.J., Rand, W., Hurd, P.L., Phelps, S.M. & Rand, A.S. 2003. Generalization in response to mate recognition signals. American Naturalist 161(3):380-394.. 30.

(31) Shine, R. 1979. Sexual selection and sexual dimorphism in amphibians. Copeia 1979:297-306. Stamps, J.A. 1983. Sexual selection, sexual dimorphism, and territoriality. In Lizard ecology: Studies os a model organism, edited by R.B Huey, E.R. Pianka, and T.W. Schoener, 169-204. Harvard University Press, Cambridge, Mass. Stamps, J.A. 1994. Territorial behavior: testing the assumptions. In Advances in the study of behavior. Vol 23, edited by P.J.B. Slater, J.S. Rosenblatt, C.T. Snowden, and M. Milinski, 173-232. Academic, San Diego. Stamps, J.A., Losos, J.B. & Andrews, R.M. 1997. A comparative study of population density and sexual size dimorphism in lizards. American Naturalist, 149 (1): 6490. Stoner, G. & Breden, F. 1988. Phenotypic differentiation in female preference related to geographic variation in male predation risk in the Trinidad guppy (Poecilia eticulate). Behavioral Ecology and Sociobiology. 22:285-291. Semler, D.E. 1971. Some aspects of adaptation in a polymorphism for breeding colours in the threespine stickleback (Gasterosteus aculeatus). Journal of Zoology Lond. 165:291-302. Stoddard, P.K. 1999. Predation enhances complexity in the evolution of electric fish signals. Nature 400: 254-256. Tinkle, D.W. 1967. The life and demography of the side-blotched lizard, Uta stansburiana. Misc. Publ. Mus. Zool., University of Michigan. 132:1-182.. 31.

(32) Voigt, C.C & O. von Helversen. 1999. Storage and display of odour by male Saccopteryx bilineata(Chiroptera, Emballonuridae). Behavioral Ecology and Sociobiology. 47:29-40. Wade, M.J. 1995. The ecology of sexual selection: mean crowding of females and resource defence polygyny. Wilczynski, W., Rand, A.S.& Ryan, M.J. 2001.Evolution of calls and auditory tuning in the Physalaemus pustulosus species group. BRAIN BEHAVIOR AND EVOLUTION, 58 (3): 137-151 Whiting, M. J, Lailvaux, S. P, Reaney, L. T, and Wymann, M. 2003. To run or hide? Age-dependent escape behaviour in the common flat lizard (Platysaurus intermedius wilhelmi). Journal of Zoology, London. 260:123-128. Wiley, R. H. & Richards, D. G. 1982. Adaptations for acoustic communication in birds: sound transmission and signal detection. In: Acoustic Communication in Birds (Ed. by D. E. Kroodsma, E. H. Miller & H. Ouellet), pp. 131–181. New York: Academic Press. Zamudio, K.R. & Sinervo, B. 2003. Ecological and social contexts for the evolution of alternative mating strategies. In Lizard Social Behavior. Edited by Stanley F. Fox , J. Kelly McCoy and Troy A Baird. The Johns Hopkins University press. Baltimore. 7-46. Zuk, M. & Kolluru, G.R. 1998. Exploitation of sexual signals by predators and parasitoids. The Quarterly Review of Biology. 73(4):415-438. 32.

(33) ACKNOWLEDGMENTS. Dr. Duncan Irschick (Tulane University) Dr. Carlos Arturo Mejia Dr. Emilio Realpe Dr. Thomas A. Jenssen (Virginia Tech) Margarita Ramos y Esteban Toro Juan Carlos Fonnegra Elizabeth Morel. 33.

(34) APPENDIX. 34.

(35) SITE. ID. C C C C C C C C C C C C C C C C C C C C C C C C C C C C C C. C1 C2. FOCAL DURATION 98170 64035. TIME DISPAYING 3556 1332. # DISPLAYS 20 11. A 1 0. B 1 0. C 18 11. X 0 0. AV. DURAT OF DISPLAY (sec) 5.926666667 4.036363637. C3 C4. 36104 15515. 1434 690. 12 5. 3 0. 5 0. 4 5. 0 0. 3.983333333 4.6. C5 C6. 5418 42929. 390 2340. 3 22. 0 2. 0 6. 3 14. 0 0. 4.333333333 3.545454547. C7 C8 C9. 20743. 180. 4. 1. 2. 1. 0. 1.5. 65530 26279. 690 1800. 5 13. 0 3. 0 4. 5 6. 0 0. 4.6 4.615384617. C10 C11. 34322 44683. 3060 3150. 30 27. 2 2. 5 3. 14 22. 9 0. 3.4 3.88888889. C12 C13. 37530 38786. 4080 3900. 23 33. 2 5. 2 5. 19 23. 0 0. 5.913043477 3.93939394. C14 C15. 27881 36522. 2601 3781. 17 31. 1 6. 6 10. 8 14. 2 1. 5.1 4.065591397. C16 C17. 33480 33681. 2486 2627. 24 28. 2 1. 9 6. 12 21. 1 0. 3.452777777 3.127380952. C18 C19. 34307 19130. 5074 2808. 38 14. 1 1. 4 9. 33 4. 0 0. 4.450877193 6.685714287. C20 C21. 19965. 2906. 28. 4. 14. 10. 0. 3.45952381. 22100. 3836. 29. 2. 0. 26. 0. 4.409195403. C22 C23 C24. 22620 14041 24115. 1851 1576 3214. 13 9 25. 0 2 0. 1 2 7. 12 5 18. 0 0 0. 4.746153847 5.837037037 4.285333333. C25 C26. 13808 16947. 1738 1574. 13 14. 0 0. 1 0. 12 14. 0 0. 4.456410257 3.747619047. C27 C28. 27773 9560. 3895 1754. 29 12. 1 2. 4 2. 24 8. 0 0. 4.477011493 4.872222223. C29. 14118. 1492. 10. 0. 0. 10. 0. 4.973333333. C30. 23254. 2997. 26. 1. 4. 21. 0. 3.842307693. Appendix 1: Data obtained after video analyses of the 30 focal videos from Tulane University population. (X corresponds to the C display variant observed for Tulane lizards). 35.

(36) DISPLAY RATE 0.036222879 0.020801124 0.039718591 0.044473091 0.071982281 0.054508607 0.008677626 0.010529528 0.068495757 0.089155644 0.070496609 0.10871303 0.100551745 0.093289337 0.103526641 0.074253286 0.077996497 0.147899846 0.146785154 0.145554721 0.173574661 0.081830239 0.112242718 0.133278043 0.125869061 0.092877795 0.140244122 0.183472803 0.105680691. DEWLAP 2540 952 817 540 270 1230 30 480 1110 1170 1920 2640 2460 1644 2251 1454 1240 3324 1685 1544 2715 1253 1037 2013 1093 960 2405 1056 970. DEWLAP RATE 0.714285714 0.714714715 0.569735007 0.782608696 0.692307692 0.525641026 0.166666667 0.695652174 0.616666667 0.382352941 0.60952381 0.647058824 0.630769231 0.632064591 0.595345147 0.584875302 0.472021317 0.655104454 0.600071225 0.531314522 0.707768509 0.676931388 0.657994924 0.62632234 0.628883774 0.609911055 0.61745828 0.602052452 0.650134048. TIME B 767 391 463 125 85 1017 179 247 584 876 1165 1488 1290 957 1530 1032 1344 1701 743 1325 1458 397 436 1182 590 665 1329 579 522. TIME U 1881 640 787 405 235 1260 14 430 1259 1920 1951 2579 2488 1644 4132 1454 1240 3373 2065 1531 2334 1238 1140 2032 1093 909 2277 1155 970. B% 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0. U% 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0. DISPLAY % 3.622287868 2.080112439 3.971859074 4.447309056 7.198228128 5.450860724 0.867762619 1.052952846 6.849575707 8.915564361 7.049660945 10.87130296 10.05517455 9.328933682 10.35266415 7.425328554 7.799649654 14.78998455 14.67851542 14.55547208 17.35746606 8.183023873 11.22427178 13.32780427 12.58690614 9.287779548 14.0244122 18.34728033 10.56806913. DEWLAP % 71.42857143 71.47147147 56.9735007 78.26086957 69.23076923 52.56410256 16.66666667 69.56521739 61.66666667 38.23529412 60.95238095 64.70588235 63.07692308 63.20645905 59.53451468 58.48753017 47.20213171 65.51044541 60.00712251 53.13145217 70.77685089 67.69313884 65.79949239 62.63223398 62.88837745 60.99110546 61.74582798 60.20524515 65.01340483. P4 6 1 3 3 1 0 0 0 0 8 0 0 2 0 0 0 1 0 0 1 1 5 0 0 1 0 0 0 0. EXTRA U 0 0 0 0 0 0 0 0 2 0 0 2 0 0 0 0 0 1 3 0 0 2 1 1 0 0 3 2 0. 0.128881053. 1748. 0.583249917. 1205. 1748. 0. 0. 12.88810527. 58.32499166. 1. 0. Appendix 1: continuation…. 36.

(37) SITE. ID. FOCAL DURATION. TIME DISPAYING. # DISPLAYS. A. B. C. X. AV. DURAT OF DISPLAY (sec). E E. E1. 12308. 0. 0. 0. 0. 0. 0. 0. E2. 40648. 1507. 15. 0. 1. 14. 0. 3.34888889. E3. 74858. 4237. 33. 3. 3. 27. 0. 4.27979798. E4. 44956. 1586. 13. 1. 6. 6. 0. 4.066666667. E5. 37400. 1125. 9. 2. 3. 4. 0. 4.166666667. E6. 10899. 909. 8. 0. 0. 8. 0. 3.7875. E7. 19161. 1293. 13. 1. 2. 10. 0. 3.315384615. E8. 9293. 195. 2. 1. 1. 0. 0. 3.25. E9. 18549. 1103. 11. 1. 3. 7. 0. 3.342424243. E10. 13845. 1032. 9. 1. 2. 6. 0. 3.822222223. E11. 12495. 2370. 19. 0. 3. 16. 0. 4.157894737. E12. 14712. 0. 0. 0. 0. 0. 0. 0. E13. 8524. 341. 3. 0. 0. 3. 0. 3.78888889. E14. 38334. 2241. 22. 2. 5. 15. 0. 3.395454547. E15. 21393. 541. 4. 0. 0. 4. 0. 4.508333333. E16. 9108. 961. 6. 0. 0. 6. 0. 5.33888889. E17. 16002. 1368. 14. 0. 0. 14. 0. 3.257142857. E18. 11574. 1367. 15. 0. 0. 15. 0. 3.037777778. E19. 30997. 185. 2. 0. 1. 2. 0. 3.083333333. E20. 30881. 2631. 24. 0. 2. 22. 0. 3.654166667. E21. 28364. 329. 3. 1. 1. 1. 0. 3.655555557. E22. 34532. 951. 9. 0. 0. 9. 0. 3.522222223. E23. 9726. 286. 3. 0. 0. 3. 0. 3.177777778. E24. 11889. 411. 4. 0. 2. 2. 0. 3.425. E25. 13100. 226. 2. 0. 0. 2. 0. 3.766666667. E E E E E E E E E E E E E E E E E E E E E E E. Appendix 2: Data obtained after video analyses of the xx focal videos from Good Hope field population.. 37.

(38) DISPLAY RATE. DEWLAP. DEWLAP RATE. TIME B. TIME U. B%. U%. DISPLAY %. DEWLAP %. P4. EXTRA U. 0. 0. 0. 0. 0. 0. 0. 0. 0. 0. 0. 0.037074395. 876. 0.581287326. 631. 876. 1.552352. 2.155088. 3.70743948. 58.1287326. 0. 0. 0.056600497. 2533. 0.597828652. 1524. 2590. 2.035855. 3.459884. 5.66004969. 59.7828652. 1. 2. 0.035278939. 915. 0.576923077. 589. 997. 1.31017. 2.217724. 3.52789394. 57.6923077. 0. 1. 0.030080214. 772. 0.686222222. 399. 726. 1.066845. 1.941176. 3.00802139. 68.6222222. 0. 0. 0.083402147. 582. 0.640264026. 327. 582. 3.000275. 5.339939. 8.3402147. 64.0264026. 0. 0. 0.06748082. 737. 0.569992266. 556. 688. 2.901727. 3.590627. 6.74808204. 56.9992266. 0. 0. 0.020983536. 97. 0.497435897. 98. 97. 1.054557. 1.043796. 2.0983536. 49.7435897. 0. 0. 0.059464122. 620. 0.562103354. 483. 620. 2.603914. 3.342498. 5.94641221. 56.2103354. 0. 0. 0.074539545. 589. 0.570736434. 443. 540. 3.199711. 3.900325. 7.4539545. 57.0736434. 0. 0. 0.18967587. 1532. 0.646413502. 856. 1468. 6.85074. 11.7487. 18.967587. 64.6413502. 1. 0. 0. 0. 0. 0. 0. 0. 0. 0. 0. 0. 0. 0.040004693. 199. 0.583577713. 142. 199. 1.665885. 2.334585. 4.00046926. 58.3577713. 0. 0. 0.058459853. 1315. 0.586791611. 926. 1243. 2.41561. 3.242552. 5.84598529. 58.6791611. 0. 0. 0.025288646. 341. 0.630314233. 200. 341. 0.934885. 1.593979. 2.52886458. 63.0314233. 0. 0. 0.105511638. 676. 0.703433923. 285. 676. 3.129117. 7.422047. 10.5511638. 70.3433923. 0. 0. 0.085489314. 1084. 0.792397661. 524. 844. 3.274591. 5.274341. 8.54893138. 79.2397661. 0. 0. 0.118109556. 944. 0.690563277. 370. 763. 3.19682. 6.592362. 11.8109556. 69.0563277. 1. 0. 0.00596832. 88. 0.475675676. 97. 88. 0.312934. 0.283898. 0.59683195. 47.5675676. 0. 0. 0.085198018. 1702. 0.646902319. 933. 1658. 3.021275. 5.368997. 8.51980182. 64.6902319. 1. 0. 0.01159921. 178. 0.541033435. 151. 178. 0.532365. 0.627556. 1.15992103. 54.1033435. 0. 0. 0.027539673. 528. 0.555205047. 423. 528. 1.224951. 1.529017. 2.75396733. 55.5205047. 0. 0. 0.029405717. 157. 0.548951049. 129. 157. 1.326342. 1.61423. 2.94057166. 54.8951049. 0. 0. 0.03456977. 241. 0.586374696. 170. 241. 1.429893. 2.027084. 3.45697704. 58.6374696. 0. 0. 0.017251908. 130. 0.575221239. 96. 130. 0.732824. 0.992366. 1.72519084. 57.5221239. 0. 0. Appendix 2: continuation…. 38.

(39) SITE. SEX. SVL. DEWLAP AREA. campus. M. 67.0. 2.937. campus. M. 69.4. 2.521. campus. M. 64.9. 1.856. campus. M. 65.8. 2.534. campus. M. 69.6. 1.642. campus. M. 68.0. 2.87. campus. M. 66.7. 1.95. campus. M. 67.6. 2.56. campus. M. 66.4. 1.956. campus. M. 64.9. 1.656. campus. M. 69.5. 2.159. campus. M. 69.8. 2.173. campus. M. 66.5. 2.33. campus. M. 71.4. 2.4. campus. M. 66.3. 1.13. campus. M. 67.7. 2.231. campus. M. 66.6. 2.255. campus. M. 68.1. 2.305. campus. M. 71.0. 2.373. campus. M. 70.4. 2.424. exxon. M. 66. 2.2342. exxon. M. 67. 1.6552. exxon. M. 67. 2.2962. exxon. M. 67. 1.855. exxon. M. 70. 2.2389. exxon. M. 71. 2.4016. exxon. M. 69. 1.9943. exxon. M. 67. 2.06. exxon. M. 66. 2.4439. exxon. M. 66. 2.2187. exxon. M. 68. 1.7815. exxon. M. 67. 1.8055. exxon. M. 65. 1.5653. exxon. M. 66. 3.1098. exxon. M. 68. 2.8012. exxon. M. 70. 2.9606. exxon. M. 67. 2.3632. exxon. M. 67. 2.2887. exxon. M. 68. 2.3352. exxon. M. 67. 2.9909. exxon. M. 70. 2.8008. Appendix 3: Dewlap areas for lizards from TU and GH..

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