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I am broadly interested in the ecology, evolution, and behavior of reptiles and amphibians - lizards in particular! I study how spatial and temporal changes in ecology and environment influence lizard behavior, morphology, physiology, genetic variation, and biological diversity across biological scales, from genes to species.

What are the effects of urbanization on lizard thermoregulatory and anti-predator behavior? How will reptiles adapt to rapid human landscape modification and global climate change? Are cities better for some herptiles than others?


Research in the lab is integrative and focuses on how urbanization and global change affect herptile ecology and evolution. To understand the effects of these human-mediated processes, we use a combination of large-scale field studies, laboratory experiments, and phylogenetic comparative analyses on individuals, populations, and species of reptiles (Podarcis wall lizards in particular).



Measuring Podarcis erhardii color morphs from Naxos, Greece




Several species of Podarcis lizards have repeatedly established thriving populations from introductions of a few individuals all over North American cities from Boston and Cincinnati to San Diego and LA and all the way up to Vancouver, often displacing native species. In San Diego and LA, introduced Italian wall lizards, Podarcis siculus, have established expanding populations that actively exclude native Western fence lizards, Sceloporus occidentalis. How do a couple of lizards from Italy explode into huge populations that push out native species that have been evolving in these places for millennia? Does Podarcis' long evolutionary history with humans give them a leg up for co-existing in cities with us? Having an understanding of the genetic, environmental, behavioral, and ecological properties of successful species invasions in cities provides insights into urban evolution and how species deal with sudden environmental change. 


I am sampling Italian wall lizards at the San Pedro, LA introduction site where the population (that started with just 7 individuals) is expanding fast. So far, I've found dramatic differences in anti-predator behavior in space (from the core to the expanding edges of the population) and through time (lizards are becoming used to human presence)! We are quantifying the rate of behavior change since their introduction to LA, along with other traits. We're also using genetic, environmental, and ecological data collected at both the source (Taormina, Sicily, Italy) and introduction (San Pedro, Los Angeles, USA) sites to try to understand how wall lizards are so successful in novel environments.

This work is in collaboration with Dr. Greg Pauly at the LA County Museum of Natural History, Anusha Bishop at UC Berkeley, Dr. Bree Putman at Cal State San Bernardino, Dr. Eric Gangloff at Ohio Wesleyan University, and Dr. Peter Bednekoff at Eastern Michigan University.


Color polymorphism defies evolutionary expectations as striking, genetically-based phenotypic variation is maintained within a single species. Yet, color polymorphism is quite common lizards - it has evolved independently multiple times in distantly related species across the lizard family tree. Color polymorphisms offer a great opportunity to study how variation, both genetic and phenotypic, is generated, maintained, and lost in natural populations. Check out some of our published work on color morph trait differences in Aegean wall lizards (Podarcis erhardii). Our findings suggest that morph colors are related to differences in traits that are crucial for social interactions and competitive ability, illustrating the need to look beyond color when studying polymorphism evolution.​

Read our paper on morph color, size, bite force, and chemical signal profiles in PeerJ

Read our paper on differential body temperature preferences between P. erhardii color morphs on bioRxiv

Read our paper on morph differences in habitat use and ectoparasite loads in Amphibia-Reptilia

More interesting work on color morph traits (aggressive behavior, microbiomes, and morphology) in collaboration with undergraduate and master's students is underway ! 



male morphs.JPG

Space is a limited resource that many animals need to access for basic functions such as feeding and reproductive opportunities. Competition over access to space can induce aggressive behavior, which may result in differential access to crucial resources related to survival and fitness. The Aegean wall lizard, Podarcis erhardii, is a tri-color polymorphic lizard that eponymously inhabits dry stone walls where they display to other lizards, access food, safely thermoregulate, and shelter from predators. Adult male color morphs exhibit physical differences in size and chemical signaling profiles, but nothing is known about potential morph-specific differences in behavior. Here, we conducted the first behavioral experiments on P. erhardii color morphs to characterize morph differences in behaviors involved in accessing and occupying a limited space resource. We captured 60 adult male lizards (20 orange, 20 yellow, and 20 white) and constructed contests between individuals of each morph type. Experimental trials conducted in a neutral laboratory arena revealed that male color morphs exhibit different behaviors and abilities to access a small stone wall with a heat source. We think these alternative strategies are involved in the evolutionary maintenance of color polymorphism.

Read our paper on morph-correlated social behaviors and contest abilities in Animal Behaviour !

Read our paper on differences in color morph escape behavior in Behavioral Ecology & Sociobiology



Evolutionary theory predicts that color polymorphic species are ripe for speciation due to dramatic phenotypic variation between color morphs within and among populations. One could imagine, for example, that across an entire species' range, that color morphs might be better suited to certain environments and ecologies, or that due to random chance a color morph might be lost from a population. But how does population-level variation in color polymorphism change across an entire species distribution? And what affect does this geographic variation in morph presence have on the mode and tempo of divergence (both morphological and genetic) among populations?


TeamSavres 2017-2019 (Team Lizards in Greek) and I travelled to the very farthest reaches of the Aegean wall lizard's (Podarcis erhardii) distribution, exploring more than 40 Aegean islands by ferry, fishing boats, and kayaks, in search of wall lizards to describe and explain color morph and genetic diversity across the archipelago. So far, we've found that color morphs in P. erhardii have been completely lost from some populations, and the pattern of morph loss seems to be ordered and non-random! We are quantifying the rate of morph loss across populations. We're also using environmental and ecological data collected at each site to try to understand drivers of morph loss.

This work has been the thrill of my life, and none it would have been possible without companionship and help from all of the wonderful TeamSavres field assistants. Ευχαριστώ πολύ Adam Rosso, Indiana Madden, Cynthia Ramos, Chris Vetros, Robert Degen, Shelby Stadler, and Cole Ayton! Thanks also to lab assistants Jazmin Reyes, Mindy Fernandez, and Liz MacDonald for extracting big beautiful fragments of lizard DNA. This work is a long-term collaboration with Professor Panayiotis Pafilis at the National and Kapodistrian University of Athens. 


Herpin' ain't easy, but it's necessary. Collecting genetic, phenotypic, and environmental data from the field is critical to advance our understanding of nature and conserving reptiles (and all biodiversity!) in a rapidly changing world. Thanks very much to the organizations that have funded TeamSavres research over the years.

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