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Ventral Coloration and Body Condition Do Not Affect Territorial Behavior in Two Sceloporus Lizards

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Ventral Coloration and Body Condition Do Not Affect Territorial Behavior in Two Sceloporus Lizards Ventral coloration and body condition do not affect territorial behavior in two Sceloporus lizards 1Marina Kelada, 1Courtney Moulton, 2Casey Nguyen, 3Griselda Robles Olague 1University of California, Riverside; 2University of California, Irvine; 3University of California, Santa Barbara Sceloporus lizards are known to be defensive of their territories, which provide shelter, food, water, and mates. They often exhibit territorial behavior through visual displays of aggression, specifically with pushups. Along with aggression, this display can also expose the blue coloration on their ventral side, serving as an intraspecific communication between lizards that reveals information about their dominance, sex, and species membership. In this study, we compared how blue coloration and body condition affected territorial behavior in two Sceloporus lizards: the well-studied western fence (Sceloporus occidentalis) and the understudied sagebrush (Sceloporus graciosus), two closely related lizards with overlapping ranges in the San Jacinto Mountains. We also addressed whether these species would exhibit character displacement in their ventral coloration as a result of their sympatry. ImageJ and Adobe Photoshop were used for photo analyses to quantify the proportion and intensity of blue on each lizard’s ventral side. Territoriality was measured through number of pushups displayed and distance traveled. We found that both ventral coloration and body condition had no effect on the territoriality of either species, suggesting that territorial behavior is dependent on factors other than their ventral coloration. Keywords: western fence lizard, sagebrush lizard, territory, blue pigmentation, Sceloporus lizard, pushups, body condition, territorial behavior INTRODUCTION Spiny lizards (Sceloporus) are no exception to this definition. This genus is one of the Territorial behavior is a crucial aspect of most diverse and species-rich clades of animal ecology, as it can influence the fitness reptiles in North America, comprising and reproductive success of many animals upwards of 90 species (Leache 2010). Some (Harris & Siefferman 2014). Although spiny lizards exhibit strong fidelity to specific animals manifest territorial behavior in localities, where they have established their various ways, the definition of territoriality home ranges, and it is to defend these provided by Sheldahl and Martins is territories that spiny lizards display especially applicable to lizards: it is the aggressive behavior (Davis & Ford 1983). relationship between aggression and the The aggression exhibited by spiny lizards consistent use of a particular area (2000). can also impact how large their territory is CEC Research | https://doi.org/10.21973/N34T0T Spring 2021 Vol. 5 Issue 4 1/8 (Davis & Ford, 1983). Having larger more territoriality than males with less blue territories yields an increase in the coloration (1987). Another study abundance of resources that animals have demonstrated that in wall lizards (Podarcis access to, like water and food availability and muralis), the blue pigmentation of the lizard access to mates. However, an animal can abdominal surface affected its bite force only defend so much area; as its home range (Lanuza et al. 2014). This study used bite expands, the territory will become more force as a proxy for aggression. Although difficult to defend (Davis & Ford 1983). This quantifying aggression in this way did trade-off is a challenge that many spiny demonstrate how differences in ventral lizards cope with, and their territory size coloration affect lizard aggression, the study must be above a certain threshold in order did not look at other aspects of territorial to maximize their fitness (Davis & Ford behavior in wall lizards like pushups (Lanuza 1983). In establishing and defending their et al. 2014). Along with this knowledge gap, territories, spiny lizards display a visual form research on the ventral coloration of of aggression: pushups, which are normally sagebrush lizards lacks, even though it has directed at another lizard (Sheldahl & ventral color patterns similar to that of the Martins 2000). However, pushups can also western fence lizard. be used to expose the unique color Among the social signals conveyed by the variations on the abdomen of male western ventral coloration of spiny lizards is species fence lizards (Sceloporus occidentalis) and identification (Cooper & Burns 1987). The male sagebrush lizards (Sceloporus ranges of the western fence lizard and the graciosus) as a mode of social signaling. sagebrush lizard are sympatric (overlapping) In several fence lizards, including the in several areas along the west coast, so it is western fence lizard and the eastern fence possible that the ventral coloration of these lizard (Sceloporus undulatas), the vibrant species would be different from each other patches of blue that males have on their in sympatric regions as a result of character throats and their abdomens are employed as displacement. Character displacement is the a means of intraspecific communication phenomenon in which the differences (Cooper & Burns 1987; Langkilde et al. 2012). among closely related species are more The ventral coloration can indicate individual pronounced where those species co-occur. quality traits to other lizards, such as social The differences are less pronounced where dominance (Langkilde et al. 2012) and those species do not overlap. Previous agonistic prowess (Cooper & Burns 1987). studies have discovered this phenomenon in Although the chromatic signaling of fence the sizes of closely related spiny lizards lizards has been well studied, it remains (Colwell & Gatz 1993). Furthermore, unclear how this signaling impacts their character displacement in the ventral territoriality. coloration of sympatric western fence lizards A previous study by Cooper and Burns and sagebrush lizards would make sense, demonstrated the importance of abdominal given it would reduce the chance of coloration in social responses of the western hybridization between the species (Ferguson fence lizard, but it did not address whether 1973). If there is character displacement in males with more blue coloration exhibited the ventral coloration of these lizards where CEC Research | https://doi.org/10.21973/N34T0T Spring 2021 Vol. 5 Issue 4 2/8 their ranges overlap, then it is possible that western fence lizard and the sagebrush one species is more aggressive than the lizard overlap. The reserve hosts unique rock other and, thus, is more territorial. formations and various plant communities We expected that the western fence lizard such as riparian forests, oak woodlands, and would have more ventral blue coloration montane chaparral. Spiny lizards prefer than the sagebrush lizard based on our initial locations with high-intensity sunlight, and observations. If this held true, we also they can be found basking atop of rocks and expected that the western fence lizard tree logs with consistent sun exposure. On would be more aggressive and would display the reserve, the western fence and more territorial behavior than the sagebrush sagebrush lizards were some of the most lizard. We also predicted that an increase in common species of lizards and would be the body condition of a lizard would frequently found within close proximity of correspond to an increase in its territoriality, each other. Both species were caught in since healthier lizards are expected to have three different locations on the reserve: the more blue and thus, should display more Flux road, the cabin site, and the solar panels pushups and maintain larger territory sizes. (Fig. 1). The Flux road and solar panels had a mix of forest and chaparral plant METHODS communities along with some rocks and human development. The vegetation 2.1 Study System included shrubs, trees, and logs/wood piles, which all provided shelter and basking areas This study was conducted at the James San for lizards. The cabin site consisted of forest Jacinto Mountains Reserve (33.80° N, and development, and lizards were found on 116.77° W), a 30-acre reserve located in top of trees, logs, cement walls, and fences. Riverside County, where the ranges of the Figure 1. Location of capturing sites. The three locations where western fence and sagebrush lizards were caught in the James San Jacinto Reserve in Riverside, CA. Each point represents one lizard that was captured in that area. CEC Research | https://doi.org/10.21973/N34T0T Spring 2021 Vol. 5 Issue 4 3/8 2.2 Capturing was continued until a total of four flags were placed for each lizard, including the original Fieldwork took place from May 5–9, 2021. flag from the time of capture. The distances We captured a total of 24 spiny lizards: 15 from each flag to the original flag were western fence (10 male, 5 female) and nine measured and averaged to acquire the sagebrush (6 male, 3 female). Both species distance (cm) a lizard traveled. This of lizards were captured using a 5.5-foot measurement was established as a proxy for fishing pole with a noose made of dental territory size as lizards rarely travel outside floss tied to the end of it. Each lizard was their home range. handled by its foreleg, and the researcher would use her forefinger and thumb to 2.4 Image Analysis gently hold the individual. While being held, species and sex was identified. Each lizard Images were analyzed using ImageJ was then photographed (iPhone XR) under software, v1.44 (SAS Institute, 2019). Each the shadow of one researcher, and a ruler photo contained a ruler and the ventral side and blue notecard were placed in the of a lizard. Processing every photo followed background of the picture. The weight of the the same four steps (Fig. 2). First, using the lizard was measured to the nearest “straight line” option, we drew a 1 cm line on hundredth of a gram. The lizards were then the ruler in the photo and set the scale to an marked using white-out and a colored fabric increment of one centimeter. Second, we paint before being released back to their drew a line from the tip of the lizard’s mouth catch site. Finally, a flag was placed where to its vent to obtain its snout-vent length in each lizard was originally seen.
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