Town and Country Reptiles: A Review of Reptilian Responses to Urbanization

Town and Country Reptiles: A Review of Reptilian Responses to Urbanization Abstract The majority of the world population is now inhabiting urban areas, and with staggering population growth, urbanization is also increasing. While the work studying the effects of changing landscapes and specific urban pressures on wildlife is beginning to amass, the majority of this work focuses on avian or mammalian species. However, the effects of urbanization likely vary substantially across taxonomic groups due to differences in habitat requirements and life history. The current article aims first to broaden the review of urban effects across reptilian species; second, to summarize the responses of reptilian fauna to specific urban features; and third, to assess the directionality of individual and population level responses to urbanization in reptile species. Based on our findings, urban research in reptilian taxa is lacking in the following areas: (1) investigating interactive or additive urban factors, (2) measuring multiple morphological, behavioral, and physiological endpoints within an animal, (3) linking individual to population-level responses, and (4) testing genetic/genomic differences across an urban environment as evidence for selective pressures. Overview Increasing human population growth necessitates the development and expansion of urban areas. The urban environment poses novel and diverse challenges for species that inhabit the landscape. Abiotic factors such as noise, artificial light, hydrology, and temperature changes (e.g., urban heat island) can cause stress, alter timing of life history events, and affect behavior and basic physiological functioning. Human-built structures can also provide habitat options for reptile species. Biotic factors, such as invasive species, can also alter community and trophic interactions as well as pathogen exposure. Thus, the potential number of urban effectors for native species is large. While the general consensus is that urbanization reduces species richness, the mechanisms for that reduction are unclear (McKinney 2008). Studies have documented responses of wildlife to specific urban areas (e.g., changes within one municipality over time), or specific species to different urban areas (e.g., passerines in different cities across the United States and Europe), but the results are often conflicting. This discrepancy is due in part to the heterogeneity of urban landscapes, to the large number of interacting and coinciding stressors in an urban landscape, but perhaps most significantly to the fact that species responses vary considerably. While most work has focused on avian and mammalian responses to urbanization, there are a growing number of studies investigating other taxonomic groups (Mitchell et al. 2008), although much of the urban work in amphibians and reptiles is investigating the abundance and spread of invasive species (Gibbon et al. 2000). Assessing the impacts on multiple taxonomic groups and a diversity of ecosystems is critical due to differences in dispersal, habitat, ecology, physiology, and life history of species inhabiting urban landscapes as well as those that are unable to inhabit these areas (McKinney 2008; Allen et al. 2017). The current review is poised to accomplish three main aims. First, is to broaden the review of urban effects across reptilian species. It is critical to assess a diversity of taxonomic groups and not solely focus on a few models species for the health of the overall ecosystem. Second, it is critical to summarize the responses of reptilian fauna to specific urban features. This is necessary because the majority of the research on urban reptiles to date assesses presence or abundance, which is important, but understanding how animals respond to specific features (e.g., contaminants, invasive species, light, noise, etc.) will allow for better management moving forward. Third, it is essential for researchers to assess the directionality of behavioral, physiological, and population level responses to urbanization in reptiles. By doing so, we will gain a better understanding of the directionality of responses, both individual and population level, providing mechanisms for the effects of urban features on wildlife. This is critical as there is surmounting evidence that reptiles are declining worldwide for a number of reasons including urbanization (Gibbon et al. 2000; Mitchell et al. 2008; Todd et al. 2010). Responses to specific urban features Largescale landscape changes and habitat fragmentation The majority of research available regarding the effects of urbanization on reptiles focuses primarily on species richness or presence/absence data (Table 1A). While most work demonstrates a decrease in population size or species richness with urbanization, a few studies instead find the opposite relationship. Rodda and Tyrrell (2008) review life history characteristics of invasive, urban, and pet herpetofauna and found that many invasive species thrive in urban settings. However, some native species such as snakes also persist and even thrive in urban environments (Schlauch 1978). Moreno-Rueda and Pizarro (2007) found that reptile species richness is positively correlated with human populations, although their focus was primarily on agricultural landscapes associated with urban environments. Similarly, Barrett and Guyer (2008) found that unlike amphibian species, reptile species significantly increased in urban watersheds in western Georgia USA, likely because of changes in canopy cover. Ackley et al. (2015b) have taken this one step further and looked at microhabitat differences in Phoenix, AZ, USA. The authors did find a negative impact of building cover, and also found that affluent areas including patches of desert remnants still retained relatively high lizard diversity and abundance (Ackley et al. 2015b). Whereas, when identifying species factors of impact on herpetofauna in northern Italy, litter and direct disturbance are negatively related to species richness (Ficetola et al. 2007). Table 1 Reptilian responses to urban features and general outcomes General response measured Species Urban factor measured Outcome Reference (A) Abundance and diversity Green anole (A. carolinensis) Cats Cats ate a lot of lizards Loyd et al. (2013) Crested anole (A. cristatellus) Urbanization Decreased presence and abundance Kolbe et al. (2016b) Lesser Antillean iguana (I. delicatissima) Urbanization Decreased abundance and local extirpations, similar density Knapp and Perez-Heydrich (2012) Dunes sagebrush lizard (S. arenicolus) Urbanization Decreased abundance Smolensky and Fitzgerald (2011) Desert grassland whiptail (Aspidoscelis uniparens) and Lesser earless lizard (Holbrookia maculata) Urbanization Decreased abundance coincided with increased roadrunner abundance Audsley et al. (2006) Fence lizard (S. occidentalis) Urbanization More frequent tonic immobility and lower sprint speeds; Significantly shorter limbs in females. Sparkman et al. (2018) Common chuckwalla (S. ater) Urbanization Similar abundance Sullivan and Sullivan (2008) Common chuckwalla (S. ater) Urbanization Similar abundance Sullivan and Williams (2010) Northern watersnake (N. sipedon) Urbanization Altered habitat use and abundance Pattishall and Cundall (2009) Eastern long-necked turtle (C. longicollis) Urbanization during dry period Increased abundance Roe et al. (2011) Eastern long-necked turtle (C. longicollis) Urbanization Similar abundance Stokeld et al. (2014) Reptile species in southeastern Spain Population density Increased species richness Moreno-Rueda and Pizarro (2007) Reptile species in western Georgia, USA Watershed development Increased species richness Barrett and Guyer (2008) Reptile species in Oxford, UK Urbanization and habitat fragmentation Species richness decreased with increasing distance from permanent water and increased with patch size. Dickman (1987) Reptile species in Australia Urbanization Vegetation remnant size correlated positively with species number How and Dell (2000) Reptile species in South Bulgaria Urbanization Reptile species richness was highest in rural zone, 2nd highest was the urban zone, and last was the suburban zone. Mollov et al. (2009) Riparian reptile species Damming Decreased species richness and occupancy Hunt et al. (2013) Reptile species in Brisbane, Australia Urbanization Landscape structure and local scale habitat were most important for species assemblages Garden et al. (2010) Lizard species in Phoenix, AZ, USA Socioeconomic status and land cover Building cover negatively affected diversity Ackley et al. (2015b) Reptile species in Indianapolis, IN, USA Altered waterways Turtles disappeared and snakes decreased Minton (1968) Reptile species in Melbourne, Australia Urbanization Decreased presence Hamer and McDonnell (2009) Snake species in NJ, USA Urbanization Species-dependent effects Zappalorti and Mitchell (2008) (B) Diet Coastal horned lizard (P. coronatum) Invasive ants Altered prey selection in areas invaded by non-native ants Suarez et al. (2000) Dugite (P. affinis) Urbanization Smaller in mass and less likely to have food in stomach Wolfe et al. (2017) (C) Behavior Side-blotched lizard (U. stansburiana) and ornate tree lizard (U. ornatus) Temperature Urban vegetation allowed for extended lizard activity Ackley et al. (2015a) Puerto Rican crested anole (A. cristatellus) Predation Increased tail autonomy and regrowth Tyler et al. (2016) Anoles (A. sagrei and A. cristatellus) Urbanization Increased body size, longer latency to feeding when offered food, and lower overall response rates Chejanovski et al. (2017) Anoles (A. cristatellus and A. stratulus) Artifical substrates Artificial substrates slowed running speed but were still used frequently Kolbe et al. (2016a) Brown anole (A. sagrei) Urbanization More tolerant to humans, less aggressive, and spent more time exploring new habitat Lapiedra et al. (2017) Common wall lizard (P. muralis) Urbanization More asymmetric traits Lazić et al. (2015) Aegean wall lizard (P. erhardii) Human built structures Switched foraging mode in new environment Donihue (2016) Dalmatian wall lizard (P. melisellensis) Urbanization Similar risk-taking and neophobia of foraging behavior De Meester et al. (2018) Delicate skink (L. delicata) Urbanization No differences in learning metrics Kang et al. (2018) Delicate skink (L. delicata) Urbanization Similar activity, exploratory, and foraging behaviors Moule et al. (2016) Garden skink (L. guichenoti) Urbanization Greater flight initiation distance, approach distance, and sprint speed Prosser et al. (2006) Side-blotched lizard (U. stansburiana) Human presence Shorter flight initiation Keehn and Feldman (2018) South Indian rock agama (P. dorsalis) Urbanization Better body condition, less diverse diet, and altered hunting strategies Balakrishna et al. (2016) Peninsular rock agama (P. dorsalis) Urbanization Shorter flight initiation distance Batabyal et al. (2017) Gila monster (H. suspectum) Urbanization No difference in home range size and movement parameters; Population sex ratio was female-biased. Kwiatkowski et al. (2008) Blue-tongued skinks (Tiliqua spp.) Noise (decible frequency) Altered movement behavior Alarcon and Fabiola (2016) Sleepy lizard (T. rugosa) Human presence and handling Increases stride frequency for up to an hour Kerr et al. (2004) Liolaemus lizards Human presence Shorter approach distance Labra and Leonard (1999) Snake species in TN, USA Urbanization and habitat fragmentation Higher fecal parasite counts Davis et al. (2012) Australian freshwater turtle (C. longicollis) Urbanization and drought Less aestivation due to increased water Rees et al. (2009) (D) Physiology Eastern fence lizard (S. undulatus) Cadmium tire byproduct Acute mortality and altered thyroid hormone Brasfield et al. (2004) Ornate tree lizard (U. ornatus) Urbanization Lower baseline and stress-induced CORT levels and altered leukocyte counts French et al. (2008) Side-blotched lizard (U. stansburiana) Urbanization Higher CORT response, reproductive investment, and oxidative stress; lower survival, innate immunity, and antioxidants Lucas and French (2012) Anoles (A. sagrei and A. cristatellus) Temperature Higher urban temperatures accelerated development of non-native anole embryos Tiatragul et al. (2017) Eurpoean wall lizard (P. muralis) Urbanization Increased parasite loads and reduced body condition Lazić et al. (2017) Lesser Antillean iguana (I. delicatissima) Urbanization No change in growth rate or body condition; asymptotic body condition Knapp and Perez-Heydrich (2012) Galápagos marine iguana (A. cristatus) Human development and tourism (including urban) Higher oxidative stress, lower immunity, and sex-dependent responses in CORT and sex hormones French et al. 2017) Common gartersnake (T. sirtalis) Indoxocarb pesticide Acute increase in CORT and immunity Neuman-Lee et al. (2016) Western terrestrial gartersnake (T. elegans) Polybrominated diphenyl ethers (PBDEs) flame retardants Altered thyroid morphology and increased body size of reproductive females and offspring Neuman-Lee et al. (2015) Copperhead (A. contortrix) Roads and traffic Reduced CORT response and no difference in baseline CORT Owen et al. 2014) Eastern long-necked turtle (C. longicollis) Urbanization during wet period Similar reproductive output and growth rate Ferronato et al. (2017) Eastern long-necked turtle (C. longicollis) Urbanization during dry period Increased growth rate Roe et al. (2011) Yellow-bellied slider (T. scripta scripta) Trace elements of coal combustion (cadmium, copper, and arsenic) Increased bactericidal ability and no change in PHA response or parasitism Haskins et al. (2017) Painted turtle (C. picta) Roads and traffic No differences in baseline or stress-induced CORT levels Polich (2016) Painted turtle (C. picta) Roads and traffic No differences in baseline or stress-induced CORT levels Baxter-Gilbert et al. (2014) Common snapping turtles (C. serpentina) Mercury exposure Reduced hatching success Hopkins et al. (2013) American Alligator (A. mississippiensis) Chemicals from pristine and contaminated lakes Thyroid and sex steroid hormone abnormalities in contaminated lakes; Smaller phallus sizes Crain et al. (1998); Guillette et al. (1999) (E) Survival Common blue-tongued skink (T. scincoides) Domestic pets and habitat loss Increased injury and mortality with increased domestic pets and habitat loss Koenig et al. (2002) Green anole (A. carolinensis) Interaction of temperature and pyrethrin pesticide Temperature and dose of pesticide interact to affect mortality Talent (2005) Texas horned lizard (P. cornutum) Urbanization Increased survival Endriss et al. (2007) Texas horned Lizard (P. cornutum) Urbanization Decreased survival Wolf et al. (2013) Salt marsh snake (N. clarkii compressicauda) Herbicide Decreased survival Ackley and Meylan (2010) Spiny softshell turtle (A. spinifera) Urbanization Decreased survival Plummer and Mills (2008) Eastern long-necked turtle (C. longicollis) Urbanization during wet period Similar abundance and survival Ferronato et al. (2017) Semi-aquatic turtle species in NC, USA Urbanization Species dependent survival Eskew et al. (2010) Reptile species in the Southwestern USA Urbanization and altered riparian habitat Species dependent extinctions and declines Lowe (1985) Lacertid lizard species in Poland Cats Cats killed more lizards in rural areas Krauze-Gryz et al. (2017) Skink species in Australia Habitat fragmentation Increased bird predation on the edge of fragmented remnants Anderson and Burgin (2008) Painted turtles (C. picta) and common snapping turtles (C. serpentine) Roads Sex-dependent mortality on roads changes population structure to male biased Steen and Gibbs (2004); Steen et al. (2006) (F) Genetic responses Crested anole (A. cristatellus) Urbanization Phenotypic shifts Winchell et al. (2016) Common wall lizard (P. muralis) Urbanization Decreased gene flow Beninde et al. (2016) West coast laterite ctenotus (C. fallens) Urbanization Decreased gene flow, similar genetic diversity, and no genetic differentiation Krawiec et al. (2015) Florida sand skink (Plestiodon reynoldsi) Urbanization Decreased gene flow and similar genetic diversity Richmond et al. (2009) California legless lizard (Anniella pulchra) Urbanization Similar genetic diversity Parham and Papenfuss (2009) Reticulated velvet gecko (Hesperoedura reticulata) Tree Dtella (Gehyra variegata) Urbanization Increased genetic differentiation Hoehn et al. 2007) Alameda striped racer (Coluber lateralis euryxanthus) Urbanization Decreased gene flow Richmond et al. (2016) Mexican dusky rattlesnake (Crotalus triseriatus) Urbanization Increased genetic differentiation and normal gene flow Sunny et al. (2015) Blanding's turtle (Emydoidea blandingii) Urbanization Decreased gene flow and genetic diversity Rubin et al. (2001) Ornate box turtle (Terrapene ornata) Urbanization Decreased gene flow, similar genetic diversity, and no genetic differentiation Cureton et al. (2014) Tuatara (Sphenodon punctatus) Urbanization Increased genetic differentiation Moore et al. (2008) Lizard species in Southern California, USA Urbanization Decreased gene flow, decreased genetic diversity, and increased genetic differentiation Delaney et al. (2010) Lizard species in Southern California, USA Urbanization Decreased gene flow and similar genetic diversity Thomassen et al. (2018) General response measured Species Urban factor measured Outcome Reference (A) Abundance and diversity Green anole (A. carolinensis) Cats Cats ate a lot of lizards Loyd et al. (2013) Crested anole (A. cristatellus) Urbanization Decreased presence and abundance Kolbe et al. (2016b) Lesser Antillean iguana (I. delicatissima) Urbanization Decreased abundance and local extirpations, similar density Knapp and Perez-Heydrich (2012) Dunes sagebrush lizard (S. arenicolus) Urbanization Decreased abundance Smolensky and Fitzgerald (2011) Desert grassland whiptail (Aspidoscelis uniparens) and Lesser earless lizard (Holbrookia maculata) Urbanization Decreased abundance coincided with increased roadrunner abundance Audsley et al. (2006) Fence lizard (S. occidentalis) Urbanization More frequent tonic immobility and lower sprint speeds; Significantly shorter limbs in females. Sparkman et al. (2018) Common chuckwalla (S. ater) Urbanization Similar abundance Sullivan and Sullivan (2008) Common chuckwalla (S. ater) Urbanization Similar abundance Sullivan and Williams (2010) Northern watersnake (N. sipedon) Urbanization Altered habitat use and abundance Pattishall and Cundall (2009) Eastern long-necked turtle (C. longicollis) Urbanization during dry period Increased abundance Roe et al. (2011) Eastern long-necked turtle (C. longicollis) Urbanization Similar abundance Stokeld et al. (2014) Reptile species in southeastern Spain Population density Increased species richness Moreno-Rueda and Pizarro (2007) Reptile species in western Georgia, USA Watershed development Increased species richness Barrett and Guyer (2008) Reptile species in Oxford, UK Urbanization and habitat fragmentation Species richness decreased with increasing distance from permanent water and increased with patch size. Dickman (1987) Reptile species in Australia Urbanization Vegetation remnant size correlated positively with species number How and Dell (2000) Reptile species in South Bulgaria Urbanization Reptile species richness was highest in rural zone, 2nd highest was the urban zone, and last was the suburban zone. Mollov et al. (2009) Riparian reptile species Damming Decreased species richness and occupancy Hunt et al. (2013) Reptile species in Brisbane, Australia Urbanization Landscape structure and local scale habitat were most important for species assemblages Garden et al. (2010) Lizard species in Phoenix, AZ, USA Socioeconomic status and land cover Building cover negatively affected diversity Ackley et al. (2015b) Reptile species in Indianapolis, IN, USA Altered waterways Turtles disappeared and snakes decreased Minton (1968) Reptile species in Melbourne, Australia Urbanization Decreased presence Hamer and McDonnell (2009) Snake species in NJ, USA Urbanization Species-dependent effects Zappalorti and Mitchell (2008) (B) Diet Coastal horned lizard (P. coronatum) Invasive ants Altered prey selection in areas invaded by non-native ants Suarez et al. (2000) Dugite (P. affinis) Urbanization Smaller in mass and less likely to have food in stomach Wolfe et al. (2017) (C) Behavior Side-blotched lizard (U. stansburiana) and ornate tree lizard (U. ornatus) Temperature Urban vegetation allowed for extended lizard activity Ackley et al. (2015a) Puerto Rican crested anole (A. cristatellus) Predation Increased tail autonomy and regrowth Tyler et al. (2016) Anoles (A. sagrei and A. cristatellus) Urbanization Increased body size, longer latency to feeding when offered food, and lower overall response rates Chejanovski et al. (2017) Anoles (A. cristatellus and A. stratulus) Artifical substrates Artificial substrates slowed running speed but were still used frequently Kolbe et al. (2016a) Brown anole (A. sagrei) Urbanization More tolerant to humans, less aggressive, and spent more time exploring new habitat Lapiedra et al. (2017) Common wall lizard (P. muralis) Urbanization More asymmetric traits Lazić et al. (2015) Aegean wall lizard (P. erhardii) Human built structures Switched foraging mode in new environment Donihue (2016) Dalmatian wall lizard (P. melisellensis) Urbanization Similar risk-taking and neophobia of foraging behavior De Meester et al. (2018) Delicate skink (L. delicata) Urbanization No differences in learning metrics Kang et al. (2018) Delicate skink (L. delicata) Urbanization Similar activity, exploratory, and foraging behaviors Moule et al. (2016) Garden skink (L. guichenoti) Urbanization Greater flight initiation distance, approach distance, and sprint speed Prosser et al. (2006) Side-blotched lizard (U. stansburiana) Human presence Shorter flight initiation Keehn and Feldman (2018) South Indian rock agama (P. dorsalis) Urbanization Better body condition, less diverse diet, and altered hunting strategies Balakrishna et al. (2016) Peninsular rock agama (P. dorsalis) Urbanization Shorter flight initiation distance Batabyal et al. (2017) Gila monster (H. suspectum) Urbanization No difference in home range size and movement parameters; Population sex ratio was female-biased. Kwiatkowski et al. (2008) Blue-tongued skinks (Tiliqua spp.) Noise (decible frequency) Altered movement behavior Alarcon and Fabiola (2016) Sleepy lizard (T. rugosa) Human presence and handling Increases stride frequency for up to an hour Kerr et al. (2004) Liolaemus lizards Human presence Shorter approach distance Labra and Leonard (1999) Snake species in TN, USA Urbanization and habitat fragmentation Higher fecal parasite counts Davis et al. (2012) Australian freshwater turtle (C. longicollis) Urbanization and drought Less aestivation due to increased water Rees et al. (2009) (D) Physiology Eastern fence lizard (S. undulatus) Cadmium tire byproduct Acute mortality and altered thyroid hormone Brasfield et al. (2004) Ornate tree lizard (U. ornatus) Urbanization Lower baseline and stress-induced CORT levels and altered leukocyte counts French et al. (2008) Side-blotched lizard (U. stansburiana) Urbanization Higher CORT response, reproductive investment, and oxidative stress; lower survival, innate immunity, and antioxidants Lucas and French (2012) Anoles (A. sagrei and A. cristatellus) Temperature Higher urban temperatures accelerated development of non-native anole embryos Tiatragul et al. (2017) Eurpoean wall lizard (P. muralis) Urbanization Increased parasite loads and reduced body condition Lazić et al. (2017) Lesser Antillean iguana (I. delicatissima) Urbanization No change in growth rate or body condition; asymptotic body condition Knapp and Perez-Heydrich (2012) Galápagos marine iguana (A. cristatus) Human development and tourism (including urban) Higher oxidative stress, lower immunity, and sex-dependent responses in CORT and sex hormones French et al. 2017) Common gartersnake (T. sirtalis) Indoxocarb pesticide Acute increase in CORT and immunity Neuman-Lee et al. (2016) Western terrestrial gartersnake (T. elegans) Polybrominated diphenyl ethers (PBDEs) flame retardants Altered thyroid morphology and increased body size of reproductive females and offspring Neuman-Lee et al. (2015) Copperhead (A. contortrix) Roads and traffic Reduced CORT response and no difference in baseline CORT Owen et al. 2014) Eastern long-necked turtle (C. longicollis) Urbanization during wet period Similar reproductive output and growth rate Ferronato et al. (2017) Eastern long-necked turtle (C. longicollis) Urbanization during dry period Increased growth rate Roe et al. (2011) Yellow-bellied slider (T. scripta scripta) Trace elements of coal combustion (cadmium, copper, and arsenic) Increased bactericidal ability and no change in PHA response or parasitism Haskins et al. (2017) Painted turtle (C. picta) Roads and traffic No differences in baseline or stress-induced CORT levels Polich (2016) Painted turtle (C. picta) Roads and traffic No differences in baseline or stress-induced CORT levels Baxter-Gilbert et al. (2014) Common snapping turtles (C. serpentina) Mercury exposure Reduced hatching success Hopkins et al. (2013) American Alligator (A. mississippiensis) Chemicals from pristine and contaminated lakes Thyroid and sex steroid hormone abnormalities in contaminated lakes; Smaller phallus sizes Crain et al. (1998); Guillette et al. (1999) (E) Survival Common blue-tongued skink (T. scincoides) Domestic pets and habitat loss Increased injury and mortality with increased domestic pets and habitat loss Koenig et al. (2002) Green anole (A. carolinensis) Interaction of temperature and pyrethrin pesticide Temperature and dose of pesticide interact to affect mortality Talent (2005) Texas horned lizard (P. cornutum) Urbanization Increased survival Endriss et al. (2007) Texas horned Lizard (P. cornutum) Urbanization Decreased survival Wolf et al. (2013) Salt marsh snake (N. clarkii compressicauda) Herbicide Decreased survival Ackley and Meylan (2010) Spiny softshell turtle (A. spinifera) Urbanization Decreased survival Plummer and Mills (2008) Eastern long-necked turtle (C. longicollis) Urbanization during wet period Similar abundance and survival Ferronato et al. (2017) Semi-aquatic turtle species in NC, USA Urbanization Species dependent survival Eskew et al. (2010) Reptile species in the Southwestern USA Urbanization and altered riparian habitat Species dependent extinctions and declines Lowe (1985) Lacertid lizard species in Poland Cats Cats killed more lizards in rural areas Krauze-Gryz et al. (2017) Skink species in Australia Habitat fragmentation Increased bird predation on the edge of fragmented remnants Anderson and Burgin (2008) Painted turtles (C. picta) and common snapping turtles (C. serpentine) Roads Sex-dependent mortality on roads changes population structure to male biased Steen and Gibbs (2004); Steen et al. (2006) (F) Genetic responses Crested anole (A. cristatellus) Urbanization Phenotypic shifts Winchell et al. (2016) Common wall lizard (P. muralis) Urbanization Decreased gene flow Beninde et al. (2016) West coast laterite ctenotus (C. fallens) Urbanization Decreased gene flow, similar genetic diversity, and no genetic differentiation Krawiec et al. (2015) Florida sand skink (Plestiodon reynoldsi) Urbanization Decreased gene flow and similar genetic diversity Richmond et al. (2009) California legless lizard (Anniella pulchra) Urbanization Similar genetic diversity Parham and Papenfuss (2009) Reticulated velvet gecko (Hesperoedura reticulata) Tree Dtella (Gehyra variegata) Urbanization Increased genetic differentiation Hoehn et al. 2007) Alameda striped racer (Coluber lateralis euryxanthus) Urbanization Decreased gene flow Richmond et al. (2016) Mexican dusky rattlesnake (Crotalus triseriatus) Urbanization Increased genetic differentiation and normal gene flow Sunny et al. (2015) Blanding's turtle (Emydoidea blandingii) Urbanization Decreased gene flow and genetic diversity Rubin et al. (2001) Ornate box turtle (Terrapene ornata) Urbanization Decreased gene flow, similar genetic diversity, and no genetic differentiation Cureton et al. (2014) Tuatara (Sphenodon punctatus) Urbanization Increased genetic differentiation Moore et al. (2008) Lizard species in Southern California, USA Urbanization Decreased gene flow, decreased genetic diversity, and increased genetic differentiation Delaney et al. (2010) Lizard species in Southern California, USA Urbanization Decreased gene flow and similar genetic diversity Thomassen et al. (2018) Notes: An organized look at original research investigating reptilian responses to urban features organized by (A) abundance and diversity, (B) diet, (C) behavior, (D) physiology, (E) survival, and (F) genetic. Within each subsection of the table, studies are ordered by taxonomic group and then followed by multispecies studies. Table 1 Reptilian responses to urban features and general outcomes General response measured Species Urban factor measured Outcome Reference (A) Abundance and diversity Green anole (A. carolinensis) Cats Cats ate a lot of lizards Loyd et al. (2013) Crested anole (A. cristatellus) Urbanization Decreased presence and abundance Kolbe et al. (2016b) Lesser Antillean iguana (I. delicatissima) Urbanization Decreased abundance and local extirpations, similar density Knapp and Perez-Heydrich (2012) Dunes sagebrush lizard (S. arenicolus) Urbanization Decreased abundance Smolensky and Fitzgerald (2011) Desert grassland whiptail (Aspidoscelis uniparens) and Lesser earless lizard (Holbrookia maculata) Urbanization Decreased abundance coincided with increased roadrunner abundance Audsley et al. (2006) Fence lizard (S. occidentalis) Urbanization More frequent tonic immobility and lower sprint speeds; Significantly shorter limbs in females. Sparkman et al. (2018) Common chuckwalla (S. ater) Urbanization Similar abundance Sullivan and Sullivan (2008) Common chuckwalla (S. ater) Urbanization Similar abundance Sullivan and Williams (2010) Northern watersnake (N. sipedon) Urbanization Altered habitat use and abundance Pattishall and Cundall (2009) Eastern long-necked turtle (C. longicollis) Urbanization during dry period Increased abundance Roe et al. (2011) Eastern long-necked turtle (C. longicollis) Urbanization Similar abundance Stokeld et al. (2014) Reptile species in southeastern Spain Population density Increased species richness Moreno-Rueda and Pizarro (2007) Reptile species in western Georgia, USA Watershed development Increased species richness Barrett and Guyer (2008) Reptile species in Oxford, UK Urbanization and habitat fragmentation Species richness decreased with increasing distance from permanent water and increased with patch size. Dickman (1987) Reptile species in Australia Urbanization Vegetation remnant size correlated positively with species number How and Dell (2000) Reptile species in South Bulgaria Urbanization Reptile species richness was highest in rural zone, 2nd highest was the urban zone, and last was the suburban zone. Mollov et al. (2009) Riparian reptile species Damming Decreased species richness and occupancy Hunt et al. (2013) Reptile species in Brisbane, Australia Urbanization Landscape structure and local scale habitat were most important for species assemblages Garden et al. (2010) Lizard species in Phoenix, AZ, USA Socioeconomic status and land cover Building cover negatively affected diversity Ackley et al. (2015b) Reptile species in Indianapolis, IN, USA Altered waterways Turtles disappeared and snakes decreased Minton (1968) Reptile species in Melbourne, Australia Urbanization Decreased presence Hamer and McDonnell (2009) Snake species in NJ, USA Urbanization Species-dependent effects Zappalorti and Mitchell (2008) (B) Diet Coastal horned lizard (P. coronatum) Invasive ants Altered prey selection in areas invaded by non-native ants Suarez et al. (2000) Dugite (P. affinis) Urbanization Smaller in mass and less likely to have food in stomach Wolfe et al. (2017) (C) Behavior Side-blotched lizard (U. stansburiana) and ornate tree lizard (U. ornatus) Temperature Urban vegetation allowed for extended lizard activity Ackley et al. (2015a) Puerto Rican crested anole (A. cristatellus) Predation Increased tail autonomy and regrowth Tyler et al. (2016) Anoles (A. sagrei and A. cristatellus) Urbanization Increased body size, longer latency to feeding when offered food, and lower overall response rates Chejanovski et al. (2017) Anoles (A. cristatellus and A. stratulus) Artifical substrates Artificial substrates slowed running speed but were still used frequently Kolbe et al. (2016a) Brown anole (A. sagrei) Urbanization More tolerant to humans, less aggressive, and spent more time exploring new habitat Lapiedra et al. (2017) Common wall lizard (P. muralis) Urbanization More asymmetric traits Lazić et al. (2015) Aegean wall lizard (P. erhardii) Human built structures Switched foraging mode in new environment Donihue (2016) Dalmatian wall lizard (P. melisellensis) Urbanization Similar risk-taking and neophobia of foraging behavior De Meester et al. (2018) Delicate skink (L. delicata) Urbanization No differences in learning metrics Kang et al. (2018) Delicate skink (L. delicata) Urbanization Similar activity, exploratory, and foraging behaviors Moule et al. (2016) Garden skink (L. guichenoti) Urbanization Greater flight initiation distance, approach distance, and sprint speed Prosser et al. (2006) Side-blotched lizard (U. stansburiana) Human presence Shorter flight initiation Keehn and Feldman (2018) South Indian rock agama (P. dorsalis) Urbanization Better body condition, less diverse diet, and altered hunting strategies Balakrishna et al. (2016) Peninsular rock agama (P. dorsalis) Urbanization Shorter flight initiation distance Batabyal et al. (2017) Gila monster (H. suspectum) Urbanization No difference in home range size and movement parameters; Population sex ratio was female-biased. Kwiatkowski et al. (2008) Blue-tongued skinks (Tiliqua spp.) Noise (decible frequency) Altered movement behavior Alarcon and Fabiola (2016) Sleepy lizard (T. rugosa) Human presence and handling Increases stride frequency for up to an hour Kerr et al. (2004) Liolaemus lizards Human presence Shorter approach distance Labra and Leonard (1999) Snake species in TN, USA Urbanization and habitat fragmentation Higher fecal parasite counts Davis et al. (2012) Australian freshwater turtle (C. longicollis) Urbanization and drought Less aestivation due to increased water Rees et al. (2009) (D) Physiology Eastern fence lizard (S. undulatus) Cadmium tire byproduct Acute mortality and altered thyroid hormone Brasfield et al. (2004) Ornate tree lizard (U. ornatus) Urbanization Lower baseline and stress-induced CORT levels and altered leukocyte counts French et al. (2008) Side-blotched lizard (U. stansburiana) Urbanization Higher CORT response, reproductive investment, and oxidative stress; lower survival, innate immunity, and antioxidants Lucas and French (2012) Anoles (A. sagrei and A. cristatellus) Temperature Higher urban temperatures accelerated development of non-native anole embryos Tiatragul et al. (2017) Eurpoean wall lizard (P. muralis) Urbanization Increased parasite loads and reduced body condition Lazić et al. (2017) Lesser Antillean iguana (I. delicatissima) Urbanization No change in growth rate or body condition; asymptotic body condition Knapp and Perez-Heydrich (2012) Galápagos marine iguana (A. cristatus) Human development and tourism (including urban) Higher oxidative stress, lower immunity, and sex-dependent responses in CORT and sex hormones French et al. 2017) Common gartersnake (T. sirtalis) Indoxocarb pesticide Acute increase in CORT and immunity Neuman-Lee et al. (2016) Western terrestrial gartersnake (T. elegans) Polybrominated diphenyl ethers (PBDEs) flame retardants Altered thyroid morphology and increased body size of reproductive females and offspring Neuman-Lee et al. (2015) Copperhead (A. contortrix) Roads and traffic Reduced CORT response and no difference in baseline CORT Owen et al. 2014) Eastern long-necked turtle (C. longicollis) Urbanization during wet period Similar reproductive output and growth rate Ferronato et al. (2017) Eastern long-necked turtle (C. longicollis) Urbanization during dry period Increased growth rate Roe et al. (2011) Yellow-bellied slider (T. scripta scripta) Trace elements of coal combustion (cadmium, copper, and arsenic) Increased bactericidal ability and no change in PHA response or parasitism Haskins et al. (2017) Painted turtle (C. picta) Roads and traffic No differences in baseline or stress-induced CORT levels Polich (2016) Painted turtle (C. picta) Roads and traffic No differences in baseline or stress-induced CORT levels Baxter-Gilbert et al. (2014) Common snapping turtles (C. serpentina) Mercury exposure Reduced hatching success Hopkins et al. (2013) American Alligator (A. mississippiensis) Chemicals from pristine and contaminated lakes Thyroid and sex steroid hormone abnormalities in contaminated lakes; Smaller phallus sizes Crain et al. (1998); Guillette et al. (1999) (E) Survival Common blue-tongued skink (T. scincoides) Domestic pets and habitat loss Increased injury and mortality with increased domestic pets and habitat loss Koenig et al. (2002) Green anole (A. carolinensis) Interaction of temperature and pyrethrin pesticide Temperature and dose of pesticide interact to affect mortality Talent (2005) Texas horned lizard (P. cornutum) Urbanization Increased survival Endriss et al. (2007) Texas horned Lizard (P. cornutum) Urbanization Decreased survival Wolf et al. (2013) Salt marsh snake (N. clarkii compressicauda) Herbicide Decreased survival Ackley and Meylan (2010) Spiny softshell turtle (A. spinifera) Urbanization Decreased survival Plummer and Mills (2008) Eastern long-necked turtle (C. longicollis) Urbanization during wet period Similar abundance and survival Ferronato et al. (2017) Semi-aquatic turtle species in NC, USA Urbanization Species dependent survival Eskew et al. (2010) Reptile species in the Southwestern USA Urbanization and altered riparian habitat Species dependent extinctions and declines Lowe (1985) Lacertid lizard species in Poland Cats Cats killed more lizards in rural areas Krauze-Gryz et al. (2017) Skink species in Australia Habitat fragmentation Increased bird predation on the edge of fragmented remnants Anderson and Burgin (2008) Painted turtles (C. picta) and common snapping turtles (C. serpentine) Roads Sex-dependent mortality on roads changes population structure to male biased Steen and Gibbs (2004); Steen et al. (2006) (F) Genetic responses Crested anole (A. cristatellus) Urbanization Phenotypic shifts Winchell et al. (2016) Common wall lizard (P. muralis) Urbanization Decreased gene flow Beninde et al. (2016) West coast laterite ctenotus (C. fallens) Urbanization Decreased gene flow, similar genetic diversity, and no genetic differentiation Krawiec et al. (2015) Florida sand skink (Plestiodon reynoldsi) Urbanization Decreased gene flow and similar genetic diversity Richmond et al. (2009) California legless lizard (Anniella pulchra) Urbanization Similar genetic diversity Parham and Papenfuss (2009) Reticulated velvet gecko (Hesperoedura reticulata) Tree Dtella (Gehyra variegata) Urbanization Increased genetic differentiation Hoehn et al. 2007) Alameda striped racer (Coluber lateralis euryxanthus) Urbanization Decreased gene flow Richmond et al. (2016) Mexican dusky rattlesnake (Crotalus triseriatus) Urbanization Increased genetic differentiation and normal gene flow Sunny et al. (2015) Blanding's turtle (Emydoidea blandingii) Urbanization Decreased gene flow and genetic diversity Rubin et al. (2001) Ornate box turtle (Terrapene ornata) Urbanization Decreased gene flow, similar genetic diversity, and no genetic differentiation Cureton et al. (2014) Tuatara (Sphenodon punctatus) Urbanization Increased genetic differentiation Moore et al. (2008) Lizard species in Southern California, USA Urbanization Decreased gene flow, decreased genetic diversity, and increased genetic differentiation Delaney et al. (2010) Lizard species in Southern California, USA Urbanization Decreased gene flow and similar genetic diversity Thomassen et al. (2018) General response measured Species Urban factor measured Outcome Reference (A) Abundance and diversity Green anole (A. carolinensis) Cats Cats ate a lot of lizards Loyd et al. (2013) Crested anole (A. cristatellus) Urbanization Decreased presence and abundance Kolbe et al. (2016b) Lesser Antillean iguana (I. delicatissima) Urbanization Decreased abundance and local extirpations, similar density Knapp and Perez-Heydrich (2012) Dunes sagebrush lizard (S. arenicolus) Urbanization Decreased abundance Smolensky and Fitzgerald (2011) Desert grassland whiptail (Aspidoscelis uniparens) and Lesser earless lizard (Holbrookia maculata) Urbanization Decreased abundance coincided with increased roadrunner abundance Audsley et al. (2006) Fence lizard (S. occidentalis) Urbanization More frequent tonic immobility and lower sprint speeds; Significantly shorter limbs in females. Sparkman et al. (2018) Common chuckwalla (S. ater) Urbanization Similar abundance Sullivan and Sullivan (2008) Common chuckwalla (S. ater) Urbanization Similar abundance Sullivan and Williams (2010) Northern watersnake (N. sipedon) Urbanization Altered habitat use and abundance Pattishall and Cundall (2009) Eastern long-necked turtle (C. longicollis) Urbanization during dry period Increased abundance Roe et al. (2011) Eastern long-necked turtle (C. longicollis) Urbanization Similar abundance Stokeld et al. (2014) Reptile species in southeastern Spain Population density Increased species richness Moreno-Rueda and Pizarro (2007) Reptile species in western Georgia, USA Watershed development Increased species richness Barrett and Guyer (2008) Reptile species in Oxford, UK Urbanization and habitat fragmentation Species richness decreased with increasing distance from permanent water and increased with patch size. Dickman (1987) Reptile species in Australia Urbanization Vegetation remnant size correlated positively with species number How and Dell (2000) Reptile species in South Bulgaria Urbanization Reptile species richness was highest in rural zone, 2nd highest was the urban zone, and last was the suburban zone. Mollov et al. (2009) Riparian reptile species Damming Decreased species richness and occupancy Hunt et al. (2013) Reptile species in Brisbane, Australia Urbanization Landscape structure and local scale habitat were most important for species assemblages Garden et al. (2010) Lizard species in Phoenix, AZ, USA Socioeconomic status and land cover Building cover negatively affected diversity Ackley et al. (2015b) Reptile species in Indianapolis, IN, USA Altered waterways Turtles disappeared and snakes decreased Minton (1968) Reptile species in Melbourne, Australia Urbanization Decreased presence Hamer and McDonnell (2009) Snake species in NJ, USA Urbanization Species-dependent effects Zappalorti and Mitchell (2008) (B) Diet Coastal horned lizard (P. coronatum) Invasive ants Altered prey selection in areas invaded by non-native ants Suarez et al. (2000) Dugite (P. affinis) Urbanization Smaller in mass and less likely to have food in stomach Wolfe et al. (2017) (C) Behavior Side-blotched lizard (U. stansburiana) and ornate tree lizard (U. ornatus) Temperature Urban vegetation allowed for extended lizard activity Ackley et al. (2015a) Puerto Rican crested anole (A. cristatellus) Predation Increased tail autonomy and regrowth Tyler et al. (2016) Anoles (A. sagrei and A. cristatellus) Urbanization Increased body size, longer latency to feeding when offered food, and lower overall response rates Chejanovski et al. (2017) Anoles (A. cristatellus and A. stratulus) Artifical substrates Artificial substrates slowed running speed but were still used frequently Kolbe et al. (2016a) Brown anole (A. sagrei) Urbanization More tolerant to humans, less aggressive, and spent more time exploring new habitat Lapiedra et al. (2017) Common wall lizard (P. muralis) Urbanization More asymmetric traits Lazić et al. (2015) Aegean wall lizard (P. erhardii) Human built structures Switched foraging mode in new environment Donihue (2016) Dalmatian wall lizard (P. melisellensis) Urbanization Similar risk-taking and neophobia of foraging behavior De Meester et al. (2018) Delicate skink (L. delicata) Urbanization No differences in learning metrics Kang et al. (2018) Delicate skink (L. delicata) Urbanization Similar activity, exploratory, and foraging behaviors Moule et al. (2016) Garden skink (L. guichenoti) Urbanization Greater flight initiation distance, approach distance, and sprint speed Prosser et al. (2006) Side-blotched lizard (U. stansburiana) Human presence Shorter flight initiation Keehn and Feldman (2018) South Indian rock agama (P. dorsalis) Urbanization Better body condition, less diverse diet, and altered hunting strategies Balakrishna et al. (2016) Peninsular rock agama (P. dorsalis) Urbanization Shorter flight initiation distance Batabyal et al. (2017) Gila monster (H. suspectum) Urbanization No difference in home range size and movement parameters; Population sex ratio was female-biased. Kwiatkowski et al. (2008) Blue-tongued skinks (Tiliqua spp.) Noise (decible frequency) Altered movement behavior Alarcon and Fabiola (2016) Sleepy lizard (T. rugosa) Human presence and handling Increases stride frequency for up to an hour Kerr et al. (2004) Liolaemus lizards Human presence Shorter approach distance Labra and Leonard (1999) Snake species in TN, USA Urbanization and habitat fragmentation Higher fecal parasite counts Davis et al. (2012) Australian freshwater turtle (C. longicollis) Urbanization and drought Less aestivation due to increased water Rees et al. (2009) (D) Physiology Eastern fence lizard (S. undulatus) Cadmium tire byproduct Acute mortality and altered thyroid hormone Brasfield et al. (2004) Ornate tree lizard (U. ornatus) Urbanization Lower baseline and stress-induced CORT levels and altered leukocyte counts French et al. (2008) Side-blotched lizard (U. stansburiana) Urbanization Higher CORT response, reproductive investment, and oxidative stress; lower survival, innate immunity, and antioxidants Lucas and French (2012) Anoles (A. sagrei and A. cristatellus) Temperature Higher urban temperatures accelerated development of non-native anole embryos Tiatragul et al. (2017) Eurpoean wall lizard (P. muralis) Urbanization Increased parasite loads and reduced body condition Lazić et al. (2017) Lesser Antillean iguana (I. delicatissima) Urbanization No change in growth rate or body condition; asymptotic body condition Knapp and Perez-Heydrich (2012) Galápagos marine iguana (A. cristatus) Human development and tourism (including urban) Higher oxidative stress, lower immunity, and sex-dependent responses in CORT and sex hormones French et al. 2017) Common gartersnake (T. sirtalis) Indoxocarb pesticide Acute increase in CORT and immunity Neuman-Lee et al. (2016) Western terrestrial gartersnake (T. elegans) Polybrominated diphenyl ethers (PBDEs) flame retardants Altered thyroid morphology and increased body size of reproductive females and offspring Neuman-Lee et al. (2015) Copperhead (A. contortrix) Roads and traffic Reduced CORT response and no difference in baseline CORT Owen et al. 2014) Eastern long-necked turtle (C. longicollis) Urbanization during wet period Similar reproductive output and growth rate Ferronato et al. (2017) Eastern long-necked turtle (C. longicollis) Urbanization during dry period Increased growth rate Roe et al. (2011) Yellow-bellied slider (T. scripta scripta) Trace elements of coal combustion (cadmium, copper, and arsenic) Increased bactericidal ability and no change in PHA response or parasitism Haskins et al. (2017) Painted turtle (C. picta) Roads and traffic No differences in baseline or stress-induced CORT levels Polich (2016) Painted turtle (C. picta) Roads and traffic No differences in baseline or stress-induced CORT levels Baxter-Gilbert et al. (2014) Common snapping turtles (C. serpentina) Mercury exposure Reduced hatching success Hopkins et al. (2013) American Alligator (A. mississippiensis) Chemicals from pristine and contaminated lakes Thyroid and sex steroid hormone abnormalities in contaminated lakes; Smaller phallus sizes Crain et al. (1998); Guillette et al. (1999) (E) Survival Common blue-tongued skink (T. scincoides) Domestic pets and habitat loss Increased injury and mortality with increased domestic pets and habitat loss Koenig et al. (2002) Green anole (A. carolinensis) Interaction of temperature and pyrethrin pesticide Temperature and dose of pesticide interact to affect mortality Talent (2005) Texas horned lizard (P. cornutum) Urbanization Increased survival Endriss et al. (2007) Texas horned Lizard (P. cornutum) Urbanization Decreased survival Wolf et al. (2013) Salt marsh snake (N. clarkii compressicauda) Herbicide Decreased survival Ackley and Meylan (2010) Spiny softshell turtle (A. spinifera) Urbanization Decreased survival Plummer and Mills (2008) Eastern long-necked turtle (C. longicollis) Urbanization during wet period Similar abundance and survival Ferronato et al. (2017) Semi-aquatic turtle species in NC, USA Urbanization Species dependent survival Eskew et al. (2010) Reptile species in the Southwestern USA Urbanization and altered riparian habitat Species dependent extinctions and declines Lowe (1985) Lacertid lizard species in Poland Cats Cats killed more lizards in rural areas Krauze-Gryz et al. (2017) Skink species in Australia Habitat fragmentation Increased bird predation on the edge of fragmented remnants Anderson and Burgin (2008) Painted turtles (C. picta) and common snapping turtles (C. serpentine) Roads Sex-dependent mortality on roads changes population structure to male biased Steen and Gibbs (2004); Steen et al. (2006) (F) Genetic responses Crested anole (A. cristatellus) Urbanization Phenotypic shifts Winchell et al. (2016) Common wall lizard (P. muralis) Urbanization Decreased gene flow Beninde et al. (2016) West coast laterite ctenotus (C. fallens) Urbanization Decreased gene flow, similar genetic diversity, and no genetic differentiation Krawiec et al. (2015) Florida sand skink (Plestiodon reynoldsi) Urbanization Decreased gene flow and similar genetic diversity Richmond et al. (2009) California legless lizard (Anniella pulchra) Urbanization Similar genetic diversity Parham and Papenfuss (2009) Reticulated velvet gecko (Hesperoedura reticulata) Tree Dtella (Gehyra variegata) Urbanization Increased genetic differentiation Hoehn et al. 2007) Alameda striped racer (Coluber lateralis euryxanthus) Urbanization Decreased gene flow Richmond et al. (2016) Mexican dusky rattlesnake (Crotalus triseriatus) Urbanization Increased genetic differentiation and normal gene flow Sunny et al. (2015) Blanding's turtle (Emydoidea blandingii) Urbanization Decreased gene flow and genetic diversity Rubin et al. (2001) Ornate box turtle (Terrapene ornata) Urbanization Decreased gene flow, similar genetic diversity, and no genetic differentiation Cureton et al. (2014) Tuatara (Sphenodon punctatus) Urbanization Increased genetic differentiation Moore et al. (2008) Lizard species in Southern California, USA Urbanization Decreased gene flow, decreased genetic diversity, and increased genetic differentiation Delaney et al. (2010) Lizard species in Southern California, USA Urbanization Decreased gene flow and similar genetic diversity Thomassen et al. (2018) Notes: An organized look at original research investigating reptilian responses to urban features organized by (A) abundance and diversity, (B) diet, (C) behavior, (D) physiology, (E) survival, and (F) genetic. Within each subsection of the table, studies are ordered by taxonomic group and then followed by multispecies studies. Habitat fragmentation also plays an important role in species richness of herpetofauna (Dickman 1987; Irwin et al. 2010). Patch size in particular has repeatedly been shown as an important predictor of reptile species richness in many different environments (Dickman 1987; Garden et al. 2007; 2010) and of species evenness (i.e., species diversity) in others (Sullivan et al. 2014). For example, remnants in metropolitan Perth, Australia were found to be an important predictor of species richness with bigger remnants having more reptile species (How and Dell 2000). Similarly, by using specialized habitats in NJ, USA (i.e., Pine Barrens) some species of snakes are able to thrive (Zappalorti and Mitchell 2008). Importantly, many of these studies show that maintaining structural complexity and remnant habitat patches can help protect some species (Hamer and McDonnell 2008). The effects of patch size appear to be species dependent, whereby habitat fragmentation in the Midwestern United States adversely effects some species of turtles (red-eared sliders, Trachemys scripta elegans) more than other species (e.g., midland painted turtles, Chrysemys picta marginata, eastern spiny softshells, Apalone spinifera spinifera, common snapping turtles, Chelydra serpentina serpentina) (Rizkalla and Swihart 2006; Ryan et al. 2014). The vast majority of work, however, demonstrates a negative impact of urbanization on reptiles. For example, Lowe found that encroaching urbanization was associated with altered riparian habitat and the extinction of several reptiles (e.g., Thamnophis species) in the American southwest (1985). These effects are seen worldwide, as for example, reptile species richness in Bulgaria was found to be highest in rural zones and followed by urban and suburban zones (Mollov et al. 2009; Mollov 2011). Many other studies also document apparent adverse effects of urbanization on reptile abundance and species richness (Table 1; Ackley et al. 2009; Banville and Bateman 2012; Hunt et al. 2013; Sullivan et al. 2017), whereby development beyond moderate levels leads to decreased number of species and abundance for those that remain (Germaine and Wakeling 2001). Modification of aquatic habitat in IN, USA, also led to declines in both turtles and snakes from the area (Minton 1968). Finally, by comparing wildlife databases of herpetofauna, researchers demonstrated that reptiles are negatively impacted by urbanization (Hamer and McDonnell 2008). While distribution studies are a critical first step in determining what species are most impacted, an important next step is to begin investigating specific factors within the urban environment that may be influencing reptile species using a combination of field and controlled laboratory studies. Biotic factors Human presence and invasive species In general, urbanization tends to decrease native species richness but promotes diversity of exotic and/or non-native species (McKinney 2006; 2008). Because urbanization can promote the establishment of non-native and generalist species, there is the potential for urbanization to influence species interactions through the introduction of novel predators, prey, and parasites. The number of invasive species is significantly increasing and thought to be one of the predominate risks for native species (Pimentel et al. 2005). For example, the presence of native crested anoles (Anolis cristatellus) is negatively associated with the presence of a non-native competitor species, brown anoles (Anolis sagrei). Here, brown anoles not only reduced the abundance of crested anoles, but also caused a shift in their use of perches (Kolbe et al. 2016b). Humans are one of the predominant invasive species present in urban environments that have been shown to alter reptilian behavior and movement patterns. Lizards from areas of high human density have shorter approach distances before initiating flight compared to lizards from areas of low human density (Labra and Leonard 1999). Marine iguanas close to towns come in more frequent contact with people including tourists in the Galapagos Islands and also show behavioral and physiological responses to intensity of exposure, including elevated stress reactivity, oxidative stress, and suppressed immunity (French et al. 2010; 2017). Another study found that lizards which have been observed, briefly handled, or extensively handled all increased stride frequency following exposure to humans (Kerr et al. 2004). While allowing humans to get closer before running away could be a result of habituation to human presence, increased stride frequency would suggest an increased cost of movement associated with human presence. While humans are the leading factor altering environments for native species, the introduction of other invasive species can also alter the landscape and ecology, having significant implications for native species. There is evidence of increased predation for reptiles in urbanized areas primarily by invasive species and pets (Fischer et al. 2012). Household pets, such as domestic cats, are known to prey upon local fauna. For example, video monitoring of free-roaming cats in southeastern United States showed that 33% of prey recovered from outdoor cats were reptiles, specifically the green anoles (Anolis carolinensis) (Loyd et al. 2013). Similarly, Koenig et al. (2002) found that blue-tongued skinks (Tiliqua scincoides) were more likely to be attacked by household pets in suburban areas compared to urban areas. However, Krauze-Gryz et al. (2017) found that reptiles in Poland were more likely to be predated by cats in rural areas. With native predators, skinks were more likely to be predated by birds on the edge of fragmented remnants compared to the core (Anderson and Burgin 2008). Similarly, in southeast Arizona, terrestrial lizard abundance decreased in urban areas along with an increase in roadrunners abundance, a common natural predator (Audsley et al. 2006), suggesting a causal link. These results provide evidence to suggest that urbanization generally increases chances of predation from both native and non-native predators. Diet Urbanization can affect reptile diet composition and frequency of feeding by altering the availability of native food sources and introducing non-native prey species (Table 1B), which, in some instances, can be beneficial. In the case of the threatened Lake Erie Water Snake (Nerodia sipedon insularum), the invasive round goby (Neogobius melanostomus) is an important food source allowing for increased growth rates and body size which can reduce predation risk during vulnerable developmental stages (King et al. 2006). Similarly, Balakrishna et al. (2016) found that Indian rock agamas (Psammophilus dorsalis) in urban environments had better body condition, had a less diverse diet, and even altered their foraging strategy compared to rural conspecifics. Not all species perform better on urban diets or with novel diet sources. For example, a study comparing diet of road-killed and museum-collected specimens showed that dugites (Pseudonaja affinis) occupying urban areas in Australia were less likely to contain a meal and were smaller in mass compared to their rural counterparts (Wolfe et al. 2017). Suarez et al. (2000) found that invasive Argentine ants (Linepithema humile) originating from urban areas displaced native ant species and significantly altered the diet composition for coastal horned lizard (Phrynosoma coronatum). Furthermore, ontogenetic differences in diet suggest the need for a diverse ant community to sustain populations, and raise concern that the documented decline in native ant species and diversity through displacement by the Argentine ant could potentially affect survival and population persistence of many ant predator species. Feeding behaviors may also differ for lizards in urban versus forested environments. Anolis lizards in urban environments of Puerto Rico have been observed to be larger and to have longer latency to feeding when offered food (Chejanovski et al. 2017). Some lizards have even switched foraging modes in response to habitat changes occurring with human presence. Aegean wall lizards (Podarcis erhardii) which utilized rock walls were more sedentary, exhibited morphological changes, and ate less sedentary prey compared to non-wall lizards (Donihue 2016). Parasites Urbanization has the potential to influence immunity and host–pathogen dynamics of urban-dwelling animals via the introduction of non-native parasites and pathogens along with other larger invasive species (Martin et al. 2010). Furthermore, changes in general ecology, including habitat size and fragmentation, can also alter disease transmission in urban habitats (Riley et al. 2014b). For example, Davis et al. (2012) found that more snakes had fecal parasites near the outer edges of an urban forest compared to snakes near the core of the forest. Similarly, Lazić et al. (2017) found that wall lizards (Podarcis muralis) had higher parasite loads and reduced body condition in urban areas compared to rural areas. These studies suggest that urbanization can potentially influence pathogen transmission among reptiles occupying previously natural habitat and adjacent areas. However, more studies are needed to further elucidate how urban cover influences transmission of parasites and susceptibility to reptile species. Abiotic factors Temperature, light, and noise Urbanization can alter abiotic features of an environment, such as temperature, light, and noise, but the direct impacts of these changes on animals is not well understood. Temperature is a dominant ecological variable for all animals that can be altered in urban environments. The majority of studies on temperature changes in urban environments have focused on endothermic species. However, known temperature changes in urban areas likely render ectotherms even more susceptible to urbanization. Decreased shade cover from plants can cause reptiles to be less active due to intense heat. Ackley et al. (2015a) demonstrated that irrigated and non-native shade planting increased lizard activity time in urban desert relative to native landscaping. Tiatragul et al. (2017) found that urban temperatures not only accelerated development of non-native anole embryos, but that non-native embryos were robust and survived well under urban temperatures. Temperature also has the potential to exacerbate other environmental stressors. For example, Talent (2005) found that temperature influenced the sensitivity of lizards to pyrethrin pesticides. Similar to temperature, light and photoperiod are critical for the timing of important life history events. Artificial light has also become a ubiquitous factor for most urban environments and while several studies have focused on birds (da Silva et al. 2014; Ouyang et al. 2017), the studies for reptile species are largely inconclusive or lacking (Perry et al. 2008). It has been suggested that no site in the continental United States is free from anthropogenic noise exposure, including remote protected areas such as national parks (Barber et al. 2011). Several studies have directly tested the effects of human noise on bird behavior and physiology (Rheindt 2003; Swaddle and Page 2007; Francis et al. 2009; Slabbekoorn 2013; Davies et al. 2017) and male tree frog (Hyla arborea) calling (Lengagne 2008) but few other animals have been studied this extensively. Alarcon and Fabiola (2016) tested different decibels and frequencies on behavior in blue-tongued skinks (T.scincoides) and found that loud, especially high frequency, noises resulted in animals spending more time freezing, a typical stress response in reptiles. Substrates and roads Some abiotic urban features may actually be beneficial to animal inhabitants, by providing access to more diverse substrates (e.g., greater refuge and perching options) and resources, allowing reptiles to persist under anthropogenic disturbances. Evidence for this has emerged through early work on comparative habitat preferences across the urban–rural landscape. For example, northern watersnakes (N.sipedon) occupying urbanized stream areas exhibit significantly greater site fidelity than those found in natural stream areas (Pattishall and Cundall 2008). Snakes in natural areas selected habitat with wide riparian zones and dense canopy cover, whereas snakes in urban areas more often occupied artificial substrates (e.g. piles of scrap metal, concrete, or holes in a railroad bed adjacent to streams) and areas with high human density (Pattishall and Cundall 2009). Artificial structures (e.g. broader and smoother substrates) are extensively utilized for perching and refuge among lizard species such as garden skinks (Lampropholis guichenoti), blue-tongued skinks (T.scincoides), crested anoles (A.cristatellus), and Gila monsters (Heloderma suspectum) (Koenig et al. 2001; Prosser et al. 2006; Winchell et al. 2016), whereas other species such as barred anoles (Anolis stratulus) tend to use more natural aspects of the urban environment (i.e., trees and other cultivated vegetation; (Winchell et al. 2018)). Artificial substrates also tend to be smoother than natural substrates for arboreal species which can impact running velocity as was demonstrated in two species of Anolis lizards (Kolbe et al. 2016a). This may in part explain why lizards display differing flight initiation distances, escape strategies, and performance levels from rural counterparts (Koenig et al. 2001; Prosser et al. 2006; Aviles-Rodriguez 2015; Winchell et al. 2016). As would be expected, urban changes in hydrology most significantly impact aquatic or riparian species, especially turtles. Rees et al. (2009) found that Australian freshwater turtles alter their behavior and are less likely to aestivate because the water supply does not seasonally dry up in urban areas. However, damming, which greatly alters riparian landscapes, reduces reptile occupancy and richness for many species, not only aquatic (Hunt et al. 2013). Perhaps most significantly, soil moisture in all habitats is critical for the development of reptile embryos of oviparous species, which constitutes the vast majority of reptile species (Ackerman 1991). Roads are a notable abiotic factor associated with urban environments that introduce changes in substrate, noise, and disturbance rates, which may in themselves also be a direct source of mortality (Ashley and Robinson 1996; Jochimsen et al. 2004; Andrews and Gibbons 2005; Andrews et al. 2008; Andrews et al. 2015). A synthesis of studies investigating the impact of roads on reptile abundance demonstrates generally negative impacts, as does a meta-analysis of life history traits and population responses to roads (Fahrig and Rytwinski 2009; Rytwinski and Fahrig 2012). Among snake species, those of smaller sizes such as ring-necked snakes (Diadophis punctatus), southeastern crown snakes (Tantilla coronata), and eastern hognose snakes (Heterodon platirhinos) more often avoid the presence of roads and traffic (Andrews and Gibbons 2005; Andrews et al. 2008; Robson and Blouin-Demers 2013). However, larger species such as gartersnakes (Thamnophis sirtalis parietalis), coachwhips (Masticophis flagellum), and brown tree snakes (Boiga irregularis) can exhibit alternative movement activities and pathways to avoid exposure and remain in natural habitats (Shine et al. 2004; Mitrovich et al. 2009; Siers et al. 2014). Similarly, turtle species including common snapping turtles and eastern painted turtles (Chrysemys picta picta) disperse into urbanized habitat less often than natural habitat, as they also exhibit avoidance behaviors toward areas with higher densities of roads (Patrick and Gibbs 2010). However, some species (Chrysemys picta and Chelydra serpentine) show sex-dependent differences in road mortality so much so that there are changes in population structure (Steen and Gibbs 2004). The sex-biased effect of roads seems to be driven by females nesting migrations that make them more likely to cross roads and be killed (Steen et al. 2006). Lizard species including blue-tongued skinks (T.scincoides), western fence lizards (Sceloporus occidentalis), orange-throated whiptails (Aspidoscelis hyperythra), and dunes sagebrush lizards (Sceloporus arenicolus) have been found to actively avoid crossing roads and instead utilize vegetation and other natural substrates for movement (Koenig et al. 2001; Brehme et al. 2013; Hibbitts et al. 2017). While the risk of mortality is reduced by road and traffic avoidance, changes in movement patterns and spatial distributions can contribute to genetic isolation and population sinks for reptiles (Forman and Alexander 1998; Shepard et al. 2008). However, the field of road ecology is growing and working to provide new planning strategies to mitigate the impacts on animals (Langen et al. 2012; Riley et al. 2014a). Pollution In many urban landscapes and adjacent developed areas, environmental toxin levels and air particulates are higher than surrounding rural areas (Cohen et al. 2004; Wei and Yang 2010). The few studies that test ecotoxicological outcomes of urban pollutants in reptiles demonstrate that the effects are not necessarily harmful. For example, yellow-bellied sliders (Trachemys scripta scripta) accumulate trace elements from coal combustion such as cadmium, copper, and arsenic as they grow, but these elements do not seem to adversely impact their immune systems measured via parasitism and responses to phytohemagglutinin, although bacterial killing ability was elevated in turtles from contaminated sites (Haskins et al. 2017). In gartersnakes (T.sirtalis), exposure to the pesticide indoxocarb induced an acute stress increase in corticosterone and immunity, whereas exposure to a similar natural toxin to which the gartersnakes have evolved resistance (i.e., tetrodoxin) did not induce a physiological response (Neuman-Lee et al. 2016). Moreover, exposure to polybrominated diphenyl ethers (PBDEs) which are used as flame retardants and are persistent contaminants found in practically every environment and organism tested, resulted in altered thyroid follicular height in female gartersnakes (Thamnophis elegans), suggesting thyroid dysfunction (Neuman-Lee et al. 2015). Neuman-Lee et al. (2017) also found an increase in body size of pregnant female gartersnakes exposed to PBDEs as well as their resulting offspring. Brasfield et al. (2004) demonstrated that exposure to cadmium, a byproduct of tire ware that is likely high in urban settings, could result in acute mortality and thyroid dysfunction in developing Eastern fence lizards (Sceloporus undulatus). Talent (2005) demonstrated that temperature influenced the sensitivity of green anoles to pyrethrin pesticides. Given that urbanization is known to alter ambient temperature and there are more pesticides in use in human-altered landscapes, this has important implications for urban reptiles. It is important to note that the effects of urban pollution can be wider reaching that just the urban footprint itself. American alligators (Alligator mississippiensis) from lakes contaminated with municipal and agricultural runoff show altered thyroid and sex steroid hormone levels and smaller phalluses (Crain et al. 1998; Guillette et al. 1999). Western pond turtles (Emys marmorata) from protected areas in California still show signs of both current- and historic-use pesticides in their blood (Meyer et al. 2016). Turtles in southwest VA, USA also show evidence of higher blood mercury levels when sampled at contaminated sites and depending on their feeding strategy (Bergeron et al. 2007). In common snapping turtles (C.serpentina), mercury levels were associated with reduced hatching success (Hopkins et al. 2013). This contamination in aquatic settings can also pass to species that prey upon aquatic animals, as observed in a viperine snake (Natrix maura) that preys on fish in France and has high mercury levels as a result (Lemaire et al. 2018). Yet, the overall evidence as to the effects of urban and anthropogenic pollutants on reptiles is limited and more research is needed (Croteau et al. 2008). In particular, researchers suggest that major ecotoxicological gaps for reptiles include better understanding the magnitude and mechanism of contaminant exposure (Weir et al. 2010; Riley et al. 2014b). Directionality of responses Individual responses In order to estimate the impact of urbanization on reptiles, understanding the directionality of how different species respond to the stressors of environmental change is pivotal. Measuring individual level responses can effectively provide real-time information concerning organismal viability in a particular environment, whereas population-level censuses may require long periods of time to yield insight. A large body of work has amassed in assessing individual responses to urbanization across several taxa, yet few studies thus far have included reptiles. Emerging findings suggest the impact of anthropogenic disturbance likely depends on habitat requirements and life history, whereby directionality for individual responses is either relatively consistent or species-specific. Behavior and morphology One of the main mechanisms through which animals respond to changing environmental conditions is by adjusting modes of behavior (Réale et al. 2007; Miranda et al. 2013; Sol et al. 2013). Differences in behavioral traits among urban and rural environments can either result from individual behavioral plasticity or microevolutionary changes (Miranda et al. 2013). Whether behavioral responses to urbanization rely on acclimation or adaptation in reptiles remains largely undetermined (Kang et al. 2018). Regardless, the directionality of behavioral responses to urbanization may be associated with life history strategies most appropriate for coping with changing environmental conditions (Huey et al. 2003; Sol and Maspons 2016; Sol et al. 2018). Across avian and mammalian taxa, urban and rural conspecifics generally vary in temperament, whereby behaviors involving neophobia or neophilia, exploration, aggression, and risk perception tend to differ (Miranda et al. 2013; Sol et al. 2013). Behavioral comparisons of urban and rural reptiles have so far been limited, but relatively consistent patterns of temperament shifts may be applicable to reptile species with similar habitat requirements and life histories (Table 1C). Of critical importance is understanding whether behavioral adjustments are occurring within a species and if such changes are beneficial in terms of survival and reproduction. Emerging studies suggest that at least some reptiles are more tolerant of particular anthropogenic factors. For example, brown anoles (A.sagrei) and crested anoles (A.cristatellus) from urban areas exhibit prolonged exploratory and foraging behaviors of new environments, as well as decreased risk perception and response rates towards predator and to human presence (Chejanovski et al. 2017; Lapiedra et al. 2017). Increased tolerance to urbanization is also evident in Indian rock agamas (P.dorsalis) and side-blotched lizards (Uta stansburiana) which exhibit decreased flight initiation distances or risk perception to anthropogenic stimuli (Batabyal et al. 2017; Keehn and Feldman 2018). However, Prosser et al. (2006) found that urban garden skinks (L.guichenoti) instead flee at a greater approach distance and exhibit greater sprint speed than conspecifics from natural habitats. Meanwhile, other studies yielded no behavioral response to urbanization including movement, exploratory, and foraging behaviors of delicate skinks (Lampropholis delicata) in the Sydney region (Moule et al. 2016). Overall activity of Gila monsters (H.suspectum) did not differ among rural and urban areas (Kwiatkowski et al. 2008). Similarly, risk-taking and neophobia of foraging behavior was not affected by urbanization in Dalmatian wall lizards (Podarcis melisellensis). Collectively, these findings are relatively in line with avian and mammal species that exhibit temperaments with more neophilic, exploratory, aggressive, and risk-taking behaviors in urban areas than in rural areas (Miranda et al. 2013; Sol et al. 2013; Greenberg and Holekamp 2017). Directionality of temperament shifts to urbanization may thus depend on whether anthropogenic conditions are beneficial, innocuous, or detrimental to the habitat requirements of a given reptile species. Assessments of habitat selection and use have yielded preliminary evidence of how urban environments may meet the habitat requirements of some species, but may fail to do so for others (also see “Substrates and Roads” section above). For example, in the same urban environment, the barred anole (A.stratulus) utilizes more natural habitat compared to the crested anole (A.cristatellus), which utilizes more anthropogenic structures (Winchell et al. 2018). Additional work has demonstrated that urban crested anoles jump from perch to perch less than rural conspecifics, which instead move around more frequently on a given perch (Aviles-Rodriguez 2015). Such preferences likely depend on the degree of similarity in environmental conditions between both habitat types as this is predicted to determine the magnitude of selective pressures for behavioral differences that may be associated with urbanization. Lastly, it is important to consider that some of these behavioral changes may be the result of morphological shifts in response to urbanization, especially in species with short generation times. A recent study on the effects of urbanization on antipredator behaviors of fence lizards found urban environments to be associated with shorter limbs, lowered sprint speed, and more frequent tonic immobility (Sparkman et al. 2018). Interestingly, certain aspects of morphology have been found to be overall smaller (e.g., head size) and more asymmetric in urban common wall lizards, suggesting divergent size–shape allometries from those in rural environments (Lazić et al. 2013; 2015). However, other morphological components have been found to be larger, including limb length in urban agamid lizards (Lophognathus temporalis) and crested anoles, and greater subdigital lamellae (i.e., footpad scales) in urban crested anoles (Iglesias et al. 2012; Winchell et al. 2016). Urban brown anoles and crested anoles both tend to exhibit greater body sizes (i.e., snout-vent lengths and masses) and body condition than those in natural environments (Chejanovski et al. 2017; Hall and Warner 2017). Yet others find no difference in body condition or growth rates, such as the lesser Antillean iguanas (Iguana delicatissima) (Knapp and Perez-Heydrich 2012). Finally, Tyler et al. (2016) found greater rates of tail autonomy and regrowth in urban anoles than their rural counterparts. In the case of all of these morphological changes, there is the potential for downstream effects on behavior or locomotion, and for the animal to incur inherent costs. Physiology Just as with behavioral research, studies investigating physiological responses to urbanization also yield mixed results (Table 1D). One common metric utilized across studies is the endocrine stress response involving activation of the hypothalamic–pituitary–adrenal axis (Saplosky 1992) and ultimately the release of glucocorticoids (i.e., corticosterone in the case of reptiles; CORT) (Moore and Jessop 2003). As compared to reptiles occurring in natural habitat, those residing in urbanized areas have been found to exhibit either similar or contrasting levels of baseline stress and stress reactivity. When considering stress physiology in snakes, no difference in baseline levels of CORT is evident in copperheads (Agkistrodon contortrix) residing in forests compared to urbanized habitat (i.e., road development and traffic) (Owen et al. 2014). Similarly, northwestern garter snakes (Thamnophis ordinoides) generally do not exhibit blood heterophil-lymphocyte ratios, indicative of chronic stress, in urbanized habitat (i.e., increased human and predator presence) (Bell 2013). Further, urban snakes, such as copperheads, exhibit reduced stress-induced CORT levels compared to forest conspecifics (Owen et al. 2014). However, copperheads in urbanized areas demonstrate a negative association between anthropogenic activity and baseline, stress-induced, and magnitude of CORT response (Owen et al. 2014). Similar trends are evident in turtles, as no differences in baseline or stress-induced CORT levels were found in painted turtles exposed to urban features (i.e., road development and traffic) as compared to those in natural areas (Baxter-Gilbert et al. 2014; Polich 2016). Stress physiology in lizard species, however, exhibits dissimilarities in response to urbanized areas. For example, tree lizards (Urosaurus ornatus) inhabiting Phoenix, AZ, USA have lower baseline and stress-induced levels of corticosterone than their rural counterparts, suggesting they have habituated to urban living (French et al. 2008). These same animals also show evidence of elevated immunity (i.e., higher leukocyte counts) perhaps to deal with increased incidence of wounding in the city (2008). However, side-blotched lizards (U.stansburiana), a not too distant relative of the tree lizard, show differential responses to city life in Saint George, UT, USA. Lucas and French (2012) found both increased corticosterone response to a stressor and elevated oxidative stress in urban side-blotched lizards. However, these same urban animals also have lower immunity (bacterial killing ability) and higher reproductive investment relative to rural side-blotched lizards. Laboratory studies on these same urban lizards demonstrated that there is direct competition for protein resources between the eggs and immunity in reproductive females (Durso and French 2018), and that immune-challenged lizards alter their energetic strategy by down-regulating metabolism (Smith et al. 2017). Taken together these results suggest urbanization may be causing a life history shift in investment from self-maintenance to reproduction, a viable strategy in a short-lived reptile (Smith and French 2017). Finally, in this context, the degree to which phenotypic plasticity, genetic evolution, or a combination thereof may underlie differences in physiology, behavior, and morphology among populations across the urban–rural landscape remains unclear. Regardless, significant changes in physiology, behavior, and morphology in urban reptiles should yield the potential to induce long-lasting effects on population size and performance over time. Population responses Although aspects of urbanization are known to place reptile populations directly at risk, whether individual responses result in additional threats to population viability is largely undetermined. Changes in individual physiology and behavior in response to urbanization can affect survival and reproduction, and thus ultimately affect populations. Linking physiological and behavioral measures to demographic parameters may elucidate undetected effects of urbanization, yet few studies have pursued such endeavors (e.g., Lucas and French 2012). Of upmost concern thus far has been the abundance of reptile populations, as individual survival and reproduction often vary with respect to biotic and abiotic factors of the urban-rural landscape (Table 1A, 1E). Anthropogenic impacts generally appear to have neutral effects, and in some cases, positive effects on the population dynamics of semi-aquatic reptiles, although detrimental effects may arise under severe cases of habitat disturbance. Despite close proximities to anthropogenic activity, abundances of northern water snake populations were similar to those in rural environments (Pattishall and Cundall 2009). Such findings are thought to be due to variable refuge and thermal opportunities provided by both terrestrial and aquatic features of the urban environment. Greater complexity and stability in urban habitats may also explain how eastern long-necked turtles (Chelodina longicollis) maintain and even increase survival, reproductive output, and population abundance in spite of temporal fluctuations in environmental conditions (Roe et al. 2011; Stokeld et al. 2014; Ferronato et al. 2017). This seems to be congruent with high survival estimates for other turtles, such as yellowbelly sliders, common snapping turtles, and spiny softshell turtles (A.spinifera), although eastern mud turtles (Kinosternon subrubrum) tend to exhibit lower survival estimates (Eskew et al. 2010; Plummer and Mills 2008). Populations of mangrove salt marsh snakes (Nerodia clarkii compressicauda) in St. Petersburg, FL, USA were also higher in abundance in anthropogenic habitats until their decline after severe disturbance (Ackley and Meylan 2010). Interestingly terrestrial and aquatic populations tend to differ in their responses. Populations of terrestrial reptiles instead seem to be more variable in their sensitivity to anthropogenic perturbations compared to semi-aquatic species. This may be due to relative differences in the urban modifications of aquatic versus terrestrial habitats (e.g., varying facets, intensities, and frequencies) or differences in survey methods and species detection rates. For example, population abundance is often related to the degree of fragmentation, size, and quality of habitat. However, different species have particular ecological requirements in habitats that determine the directionality of response. Urban disturbance in the form of increasingly fragmented landscapes often causes fast declines and local extirpations in reptile populations, such as in the lesser Antillean iguana (I.delicatissima) (Knapp and Perez-Heydrich 2012). In the case of Texas horned lizards (Phrynosoma cornutum) in central Oklahoma, urban development caused declines in the abundance due to increased mortality despite moderate reproductive output (Endriss et al. 2007; Wolf et al. 2013). Populations of dunes sagebrush lizards vary significantly in abundance and this variation could be explained by habitat patch size and quality, which are affected by human development, oil, and gas industry (Smolensky and Fitzgerald 2011). Similarly, declines in population abundance for the invasive crested anole (A.cristatellus) in Miami, FL, USA are strongly associated with losses in habitat size and quality (Kolbe et al. 2016b). In other urban habitats with limited refuge and plant food density, population abundance of common chuckwallas (Sauromalus ater) is dependent on the availability of plant food diversity (Sullivan and Sullivan 2008; Sullivan and Williams 2010). Collectively, these studies suggest that urbanization can lead to population level changes but that some species, and perhaps environments, are more sensitive. Over multiple generations, differences in the selective pressures of an environment and the resulting physiological and population-level changes could potentially lead to genetic differentiation. Genetic Urbanization may have genetic consequences among reptile populations, as land-use changes can lead to intense forms of habitat alteration (Table 1F). Herpetofauna that are particularly sensitive to habitat degradation can rapidly become extirpated from urban locales (Gibbon et al. 2000; Cushman 2006; Hamer and McDonnell 2009). For those that persist across fragmented urban landscapes, connectivity and gene flow between populations may be hindered or inhibited, which can ultimately lead to reductions in genetic diversity, inbreeding depression, and even local extinction (Reed et al. 2002; Reed et al. 2003; Reed 2004; Cushman 2006; Frankham 2006). Degraded and fragmented habitats are also less likely to be recolonized by extinguished species. Additionally, the loss of genetic diversity in remaining populations can reduce adaptive potential in response to environmental changes. Although there are population and behavioral studies to document thriving and mobile urban reptiles species, preliminary genetic data provides little evidence of continuous gene flow for various reptile species located along fragmented urban landscapes (e.g., Delaney et al. 2010; Krawiec et al. 2015; Beninde et al. 2016; Richmond et al. 2016; Thomassen et al. 2018). Inhibited or decreased gene flow, in turn, seems to either reduce genetic diversity levels (Rubin et al. 2001; Delaney et al. 2010) or yield no effect (Parham and Papenfuss 2009; Richmond et al. 2009; Cureton et al. 2014; Krawiec et al. 2015; Sunny et al. 2015). There are often remarkable amounts of genetic divergence of populations across the urban–rural landscape, which may be occurring over relatively short geographic and temporal scales (Hoehn et al. 2007; Moore et al. 2008; Delaney et al. 2010; Sunny et al. 2015). Implications for population genetics may thus depend on the size, configuration, age, and isolation of habitat fragments. Studies are revealing evidence of selection for divergent phenotypes and suggest a potential for reptile populations to adapt to urban environments (Winchell et al. 2016; 2018). Conclusions Overall, this review has identified complex and diverse results that are variable both within and among all scales of ecological organization. There is variability in the directionality of the responses to urbanization, whereby some studies find inert or even positive effects of urbanization, while others show the opposite. This discrepancy is due in part to the heterogeneity of urban landscapes and to the fact that species responses are also different. Another important factor leading to inconsistencies in study outcomes is that urbanization includes many different factors interacting simultaneously. While there are a handful of studies testing the interactive effects of urban qualities, most are focused on non-reptile taxonomic groups (Isaksson 2015). With a growing body of studies investigating specific urban features or stressors, it is becoming increasingly important to measure interactive effects (Talent 2005), physiological responses, and multiple health indicators simultaneously. Testing only one endpoint can produce misleading results due to tradeoffs which can occur between physiological systems (Lucas and French 2012). Also, the large number of different approaches used in these studies make it difficult to compare outcomes and to assess directionality of the response. Given the methodologies and results of most experiments, we can only assess whether or not there is an effect of the urban environment or stressor, and not the fitness implications of that given effect. To conclude, based on our findings there is an apparent lack of urban research in reptilian species (1) investigating interactive or additive urban factors which more accurately represent the reality of urbanization; (2) measuring multiple morphological, behavioral, and physiological responses, because a more comprehensive approach will allow researchers to better assess directionality; (3) linking individual to population-level responses to identify the mechanisms for population changes, especially declines, and (4) testing genetic/genomic differences across an urban environment as evidence for selective pressures. These important gaps will need to be filled in the near future as urbanization and animal population declines continue. Finally, there is an imperative need for better community outreach, involvement, and education to make conservation of all species possible (McKinney 2002). Funding This work was supported by the National Science Foundation [(IOS)-1350070 to S.S.F.]. 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Published by Oxford University Press on behalf of the Society for Integrative and Comparative Biology. All rights reserved. For permissions please email: journals.permissions@oup.com. This article is published and distributed under the terms of the Oxford University Press, Standard Journals Publication Model (https://academic.oup.com/journals/pages/about_us/legal/notices) http://www.deepdyve.com/assets/images/DeepDyve-Logo-lg.png Integrative and Comparative Biology Oxford University Press

Town and Country Reptiles: A Review of Reptilian Responses to Urbanization

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Oxford University Press
Copyright
© The Author(s) 2018. Published by Oxford University Press on behalf of the Society for Integrative and Comparative Biology. All rights reserved. For permissions please email: journals.permissions@oup.com.
ISSN
1540-7063
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1557-7023
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10.1093/icb/icy052
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Abstract

Abstract The majority of the world population is now inhabiting urban areas, and with staggering population growth, urbanization is also increasing. While the work studying the effects of changing landscapes and specific urban pressures on wildlife is beginning to amass, the majority of this work focuses on avian or mammalian species. However, the effects of urbanization likely vary substantially across taxonomic groups due to differences in habitat requirements and life history. The current article aims first to broaden the review of urban effects across reptilian species; second, to summarize the responses of reptilian fauna to specific urban features; and third, to assess the directionality of individual and population level responses to urbanization in reptile species. Based on our findings, urban research in reptilian taxa is lacking in the following areas: (1) investigating interactive or additive urban factors, (2) measuring multiple morphological, behavioral, and physiological endpoints within an animal, (3) linking individual to population-level responses, and (4) testing genetic/genomic differences across an urban environment as evidence for selective pressures. Overview Increasing human population growth necessitates the development and expansion of urban areas. The urban environment poses novel and diverse challenges for species that inhabit the landscape. Abiotic factors such as noise, artificial light, hydrology, and temperature changes (e.g., urban heat island) can cause stress, alter timing of life history events, and affect behavior and basic physiological functioning. Human-built structures can also provide habitat options for reptile species. Biotic factors, such as invasive species, can also alter community and trophic interactions as well as pathogen exposure. Thus, the potential number of urban effectors for native species is large. While the general consensus is that urbanization reduces species richness, the mechanisms for that reduction are unclear (McKinney 2008). Studies have documented responses of wildlife to specific urban areas (e.g., changes within one municipality over time), or specific species to different urban areas (e.g., passerines in different cities across the United States and Europe), but the results are often conflicting. This discrepancy is due in part to the heterogeneity of urban landscapes, to the large number of interacting and coinciding stressors in an urban landscape, but perhaps most significantly to the fact that species responses vary considerably. While most work has focused on avian and mammalian responses to urbanization, there are a growing number of studies investigating other taxonomic groups (Mitchell et al. 2008), although much of the urban work in amphibians and reptiles is investigating the abundance and spread of invasive species (Gibbon et al. 2000). Assessing the impacts on multiple taxonomic groups and a diversity of ecosystems is critical due to differences in dispersal, habitat, ecology, physiology, and life history of species inhabiting urban landscapes as well as those that are unable to inhabit these areas (McKinney 2008; Allen et al. 2017). The current review is poised to accomplish three main aims. First, is to broaden the review of urban effects across reptilian species. It is critical to assess a diversity of taxonomic groups and not solely focus on a few models species for the health of the overall ecosystem. Second, it is critical to summarize the responses of reptilian fauna to specific urban features. This is necessary because the majority of the research on urban reptiles to date assesses presence or abundance, which is important, but understanding how animals respond to specific features (e.g., contaminants, invasive species, light, noise, etc.) will allow for better management moving forward. Third, it is essential for researchers to assess the directionality of behavioral, physiological, and population level responses to urbanization in reptiles. By doing so, we will gain a better understanding of the directionality of responses, both individual and population level, providing mechanisms for the effects of urban features on wildlife. This is critical as there is surmounting evidence that reptiles are declining worldwide for a number of reasons including urbanization (Gibbon et al. 2000; Mitchell et al. 2008; Todd et al. 2010). Responses to specific urban features Largescale landscape changes and habitat fragmentation The majority of research available regarding the effects of urbanization on reptiles focuses primarily on species richness or presence/absence data (Table 1A). While most work demonstrates a decrease in population size or species richness with urbanization, a few studies instead find the opposite relationship. Rodda and Tyrrell (2008) review life history characteristics of invasive, urban, and pet herpetofauna and found that many invasive species thrive in urban settings. However, some native species such as snakes also persist and even thrive in urban environments (Schlauch 1978). Moreno-Rueda and Pizarro (2007) found that reptile species richness is positively correlated with human populations, although their focus was primarily on agricultural landscapes associated with urban environments. Similarly, Barrett and Guyer (2008) found that unlike amphibian species, reptile species significantly increased in urban watersheds in western Georgia USA, likely because of changes in canopy cover. Ackley et al. (2015b) have taken this one step further and looked at microhabitat differences in Phoenix, AZ, USA. The authors did find a negative impact of building cover, and also found that affluent areas including patches of desert remnants still retained relatively high lizard diversity and abundance (Ackley et al. 2015b). Whereas, when identifying species factors of impact on herpetofauna in northern Italy, litter and direct disturbance are negatively related to species richness (Ficetola et al. 2007). Table 1 Reptilian responses to urban features and general outcomes General response measured Species Urban factor measured Outcome Reference (A) Abundance and diversity Green anole (A. carolinensis) Cats Cats ate a lot of lizards Loyd et al. (2013) Crested anole (A. cristatellus) Urbanization Decreased presence and abundance Kolbe et al. (2016b) Lesser Antillean iguana (I. delicatissima) Urbanization Decreased abundance and local extirpations, similar density Knapp and Perez-Heydrich (2012) Dunes sagebrush lizard (S. arenicolus) Urbanization Decreased abundance Smolensky and Fitzgerald (2011) Desert grassland whiptail (Aspidoscelis uniparens) and Lesser earless lizard (Holbrookia maculata) Urbanization Decreased abundance coincided with increased roadrunner abundance Audsley et al. (2006) Fence lizard (S. occidentalis) Urbanization More frequent tonic immobility and lower sprint speeds; Significantly shorter limbs in females. Sparkman et al. (2018) Common chuckwalla (S. ater) Urbanization Similar abundance Sullivan and Sullivan (2008) Common chuckwalla (S. ater) Urbanization Similar abundance Sullivan and Williams (2010) Northern watersnake (N. sipedon) Urbanization Altered habitat use and abundance Pattishall and Cundall (2009) Eastern long-necked turtle (C. longicollis) Urbanization during dry period Increased abundance Roe et al. (2011) Eastern long-necked turtle (C. longicollis) Urbanization Similar abundance Stokeld et al. (2014) Reptile species in southeastern Spain Population density Increased species richness Moreno-Rueda and Pizarro (2007) Reptile species in western Georgia, USA Watershed development Increased species richness Barrett and Guyer (2008) Reptile species in Oxford, UK Urbanization and habitat fragmentation Species richness decreased with increasing distance from permanent water and increased with patch size. Dickman (1987) Reptile species in Australia Urbanization Vegetation remnant size correlated positively with species number How and Dell (2000) Reptile species in South Bulgaria Urbanization Reptile species richness was highest in rural zone, 2nd highest was the urban zone, and last was the suburban zone. Mollov et al. (2009) Riparian reptile species Damming Decreased species richness and occupancy Hunt et al. (2013) Reptile species in Brisbane, Australia Urbanization Landscape structure and local scale habitat were most important for species assemblages Garden et al. (2010) Lizard species in Phoenix, AZ, USA Socioeconomic status and land cover Building cover negatively affected diversity Ackley et al. (2015b) Reptile species in Indianapolis, IN, USA Altered waterways Turtles disappeared and snakes decreased Minton (1968) Reptile species in Melbourne, Australia Urbanization Decreased presence Hamer and McDonnell (2009) Snake species in NJ, USA Urbanization Species-dependent effects Zappalorti and Mitchell (2008) (B) Diet Coastal horned lizard (P. coronatum) Invasive ants Altered prey selection in areas invaded by non-native ants Suarez et al. (2000) Dugite (P. affinis) Urbanization Smaller in mass and less likely to have food in stomach Wolfe et al. (2017) (C) Behavior Side-blotched lizard (U. stansburiana) and ornate tree lizard (U. ornatus) Temperature Urban vegetation allowed for extended lizard activity Ackley et al. (2015a) Puerto Rican crested anole (A. cristatellus) Predation Increased tail autonomy and regrowth Tyler et al. (2016) Anoles (A. sagrei and A. cristatellus) Urbanization Increased body size, longer latency to feeding when offered food, and lower overall response rates Chejanovski et al. (2017) Anoles (A. cristatellus and A. stratulus) Artifical substrates Artificial substrates slowed running speed but were still used frequently Kolbe et al. (2016a) Brown anole (A. sagrei) Urbanization More tolerant to humans, less aggressive, and spent more time exploring new habitat Lapiedra et al. (2017) Common wall lizard (P. muralis) Urbanization More asymmetric traits Lazić et al. (2015) Aegean wall lizard (P. erhardii) Human built structures Switched foraging mode in new environment Donihue (2016) Dalmatian wall lizard (P. melisellensis) Urbanization Similar risk-taking and neophobia of foraging behavior De Meester et al. (2018) Delicate skink (L. delicata) Urbanization No differences in learning metrics Kang et al. (2018) Delicate skink (L. delicata) Urbanization Similar activity, exploratory, and foraging behaviors Moule et al. (2016) Garden skink (L. guichenoti) Urbanization Greater flight initiation distance, approach distance, and sprint speed Prosser et al. (2006) Side-blotched lizard (U. stansburiana) Human presence Shorter flight initiation Keehn and Feldman (2018) South Indian rock agama (P. dorsalis) Urbanization Better body condition, less diverse diet, and altered hunting strategies Balakrishna et al. (2016) Peninsular rock agama (P. dorsalis) Urbanization Shorter flight initiation distance Batabyal et al. (2017) Gila monster (H. suspectum) Urbanization No difference in home range size and movement parameters; Population sex ratio was female-biased. Kwiatkowski et al. (2008) Blue-tongued skinks (Tiliqua spp.) Noise (decible frequency) Altered movement behavior Alarcon and Fabiola (2016) Sleepy lizard (T. rugosa) Human presence and handling Increases stride frequency for up to an hour Kerr et al. (2004) Liolaemus lizards Human presence Shorter approach distance Labra and Leonard (1999) Snake species in TN, USA Urbanization and habitat fragmentation Higher fecal parasite counts Davis et al. (2012) Australian freshwater turtle (C. longicollis) Urbanization and drought Less aestivation due to increased water Rees et al. (2009) (D) Physiology Eastern fence lizard (S. undulatus) Cadmium tire byproduct Acute mortality and altered thyroid hormone Brasfield et al. (2004) Ornate tree lizard (U. ornatus) Urbanization Lower baseline and stress-induced CORT levels and altered leukocyte counts French et al. (2008) Side-blotched lizard (U. stansburiana) Urbanization Higher CORT response, reproductive investment, and oxidative stress; lower survival, innate immunity, and antioxidants Lucas and French (2012) Anoles (A. sagrei and A. cristatellus) Temperature Higher urban temperatures accelerated development of non-native anole embryos Tiatragul et al. (2017) Eurpoean wall lizard (P. muralis) Urbanization Increased parasite loads and reduced body condition Lazić et al. (2017) Lesser Antillean iguana (I. delicatissima) Urbanization No change in growth rate or body condition; asymptotic body condition Knapp and Perez-Heydrich (2012) Galápagos marine iguana (A. cristatus) Human development and tourism (including urban) Higher oxidative stress, lower immunity, and sex-dependent responses in CORT and sex hormones French et al. 2017) Common gartersnake (T. sirtalis) Indoxocarb pesticide Acute increase in CORT and immunity Neuman-Lee et al. (2016) Western terrestrial gartersnake (T. elegans) Polybrominated diphenyl ethers (PBDEs) flame retardants Altered thyroid morphology and increased body size of reproductive females and offspring Neuman-Lee et al. (2015) Copperhead (A. contortrix) Roads and traffic Reduced CORT response and no difference in baseline CORT Owen et al. 2014) Eastern long-necked turtle (C. longicollis) Urbanization during wet period Similar reproductive output and growth rate Ferronato et al. (2017) Eastern long-necked turtle (C. longicollis) Urbanization during dry period Increased growth rate Roe et al. (2011) Yellow-bellied slider (T. scripta scripta) Trace elements of coal combustion (cadmium, copper, and arsenic) Increased bactericidal ability and no change in PHA response or parasitism Haskins et al. (2017) Painted turtle (C. picta) Roads and traffic No differences in baseline or stress-induced CORT levels Polich (2016) Painted turtle (C. picta) Roads and traffic No differences in baseline or stress-induced CORT levels Baxter-Gilbert et al. (2014) Common snapping turtles (C. serpentina) Mercury exposure Reduced hatching success Hopkins et al. (2013) American Alligator (A. mississippiensis) Chemicals from pristine and contaminated lakes Thyroid and sex steroid hormone abnormalities in contaminated lakes; Smaller phallus sizes Crain et al. (1998); Guillette et al. (1999) (E) Survival Common blue-tongued skink (T. scincoides) Domestic pets and habitat loss Increased injury and mortality with increased domestic pets and habitat loss Koenig et al. (2002) Green anole (A. carolinensis) Interaction of temperature and pyrethrin pesticide Temperature and dose of pesticide interact to affect mortality Talent (2005) Texas horned lizard (P. cornutum) Urbanization Increased survival Endriss et al. (2007) Texas horned Lizard (P. cornutum) Urbanization Decreased survival Wolf et al. (2013) Salt marsh snake (N. clarkii compressicauda) Herbicide Decreased survival Ackley and Meylan (2010) Spiny softshell turtle (A. spinifera) Urbanization Decreased survival Plummer and Mills (2008) Eastern long-necked turtle (C. longicollis) Urbanization during wet period Similar abundance and survival Ferronato et al. (2017) Semi-aquatic turtle species in NC, USA Urbanization Species dependent survival Eskew et al. (2010) Reptile species in the Southwestern USA Urbanization and altered riparian habitat Species dependent extinctions and declines Lowe (1985) Lacertid lizard species in Poland Cats Cats killed more lizards in rural areas Krauze-Gryz et al. (2017) Skink species in Australia Habitat fragmentation Increased bird predation on the edge of fragmented remnants Anderson and Burgin (2008) Painted turtles (C. picta) and common snapping turtles (C. serpentine) Roads Sex-dependent mortality on roads changes population structure to male biased Steen and Gibbs (2004); Steen et al. (2006) (F) Genetic responses Crested anole (A. cristatellus) Urbanization Phenotypic shifts Winchell et al. (2016) Common wall lizard (P. muralis) Urbanization Decreased gene flow Beninde et al. (2016) West coast laterite ctenotus (C. fallens) Urbanization Decreased gene flow, similar genetic diversity, and no genetic differentiation Krawiec et al. (2015) Florida sand skink (Plestiodon reynoldsi) Urbanization Decreased gene flow and similar genetic diversity Richmond et al. (2009) California legless lizard (Anniella pulchra) Urbanization Similar genetic diversity Parham and Papenfuss (2009) Reticulated velvet gecko (Hesperoedura reticulata) Tree Dtella (Gehyra variegata) Urbanization Increased genetic differentiation Hoehn et al. 2007) Alameda striped racer (Coluber lateralis euryxanthus) Urbanization Decreased gene flow Richmond et al. (2016) Mexican dusky rattlesnake (Crotalus triseriatus) Urbanization Increased genetic differentiation and normal gene flow Sunny et al. (2015) Blanding's turtle (Emydoidea blandingii) Urbanization Decreased gene flow and genetic diversity Rubin et al. (2001) Ornate box turtle (Terrapene ornata) Urbanization Decreased gene flow, similar genetic diversity, and no genetic differentiation Cureton et al. (2014) Tuatara (Sphenodon punctatus) Urbanization Increased genetic differentiation Moore et al. (2008) Lizard species in Southern California, USA Urbanization Decreased gene flow, decreased genetic diversity, and increased genetic differentiation Delaney et al. (2010) Lizard species in Southern California, USA Urbanization Decreased gene flow and similar genetic diversity Thomassen et al. (2018) General response measured Species Urban factor measured Outcome Reference (A) Abundance and diversity Green anole (A. carolinensis) Cats Cats ate a lot of lizards Loyd et al. (2013) Crested anole (A. cristatellus) Urbanization Decreased presence and abundance Kolbe et al. (2016b) Lesser Antillean iguana (I. delicatissima) Urbanization Decreased abundance and local extirpations, similar density Knapp and Perez-Heydrich (2012) Dunes sagebrush lizard (S. arenicolus) Urbanization Decreased abundance Smolensky and Fitzgerald (2011) Desert grassland whiptail (Aspidoscelis uniparens) and Lesser earless lizard (Holbrookia maculata) Urbanization Decreased abundance coincided with increased roadrunner abundance Audsley et al. (2006) Fence lizard (S. occidentalis) Urbanization More frequent tonic immobility and lower sprint speeds; Significantly shorter limbs in females. Sparkman et al. (2018) Common chuckwalla (S. ater) Urbanization Similar abundance Sullivan and Sullivan (2008) Common chuckwalla (S. ater) Urbanization Similar abundance Sullivan and Williams (2010) Northern watersnake (N. sipedon) Urbanization Altered habitat use and abundance Pattishall and Cundall (2009) Eastern long-necked turtle (C. longicollis) Urbanization during dry period Increased abundance Roe et al. (2011) Eastern long-necked turtle (C. longicollis) Urbanization Similar abundance Stokeld et al. (2014) Reptile species in southeastern Spain Population density Increased species richness Moreno-Rueda and Pizarro (2007) Reptile species in western Georgia, USA Watershed development Increased species richness Barrett and Guyer (2008) Reptile species in Oxford, UK Urbanization and habitat fragmentation Species richness decreased with increasing distance from permanent water and increased with patch size. Dickman (1987) Reptile species in Australia Urbanization Vegetation remnant size correlated positively with species number How and Dell (2000) Reptile species in South Bulgaria Urbanization Reptile species richness was highest in rural zone, 2nd highest was the urban zone, and last was the suburban zone. Mollov et al. (2009) Riparian reptile species Damming Decreased species richness and occupancy Hunt et al. (2013) Reptile species in Brisbane, Australia Urbanization Landscape structure and local scale habitat were most important for species assemblages Garden et al. (2010) Lizard species in Phoenix, AZ, USA Socioeconomic status and land cover Building cover negatively affected diversity Ackley et al. (2015b) Reptile species in Indianapolis, IN, USA Altered waterways Turtles disappeared and snakes decreased Minton (1968) Reptile species in Melbourne, Australia Urbanization Decreased presence Hamer and McDonnell (2009) Snake species in NJ, USA Urbanization Species-dependent effects Zappalorti and Mitchell (2008) (B) Diet Coastal horned lizard (P. coronatum) Invasive ants Altered prey selection in areas invaded by non-native ants Suarez et al. (2000) Dugite (P. affinis) Urbanization Smaller in mass and less likely to have food in stomach Wolfe et al. (2017) (C) Behavior Side-blotched lizard (U. stansburiana) and ornate tree lizard (U. ornatus) Temperature Urban vegetation allowed for extended lizard activity Ackley et al. (2015a) Puerto Rican crested anole (A. cristatellus) Predation Increased tail autonomy and regrowth Tyler et al. (2016) Anoles (A. sagrei and A. cristatellus) Urbanization Increased body size, longer latency to feeding when offered food, and lower overall response rates Chejanovski et al. (2017) Anoles (A. cristatellus and A. stratulus) Artifical substrates Artificial substrates slowed running speed but were still used frequently Kolbe et al. (2016a) Brown anole (A. sagrei) Urbanization More tolerant to humans, less aggressive, and spent more time exploring new habitat Lapiedra et al. (2017) Common wall lizard (P. muralis) Urbanization More asymmetric traits Lazić et al. (2015) Aegean wall lizard (P. erhardii) Human built structures Switched foraging mode in new environment Donihue (2016) Dalmatian wall lizard (P. melisellensis) Urbanization Similar risk-taking and neophobia of foraging behavior De Meester et al. (2018) Delicate skink (L. delicata) Urbanization No differences in learning metrics Kang et al. (2018) Delicate skink (L. delicata) Urbanization Similar activity, exploratory, and foraging behaviors Moule et al. (2016) Garden skink (L. guichenoti) Urbanization Greater flight initiation distance, approach distance, and sprint speed Prosser et al. (2006) Side-blotched lizard (U. stansburiana) Human presence Shorter flight initiation Keehn and Feldman (2018) South Indian rock agama (P. dorsalis) Urbanization Better body condition, less diverse diet, and altered hunting strategies Balakrishna et al. (2016) Peninsular rock agama (P. dorsalis) Urbanization Shorter flight initiation distance Batabyal et al. (2017) Gila monster (H. suspectum) Urbanization No difference in home range size and movement parameters; Population sex ratio was female-biased. Kwiatkowski et al. (2008) Blue-tongued skinks (Tiliqua spp.) Noise (decible frequency) Altered movement behavior Alarcon and Fabiola (2016) Sleepy lizard (T. rugosa) Human presence and handling Increases stride frequency for up to an hour Kerr et al. (2004) Liolaemus lizards Human presence Shorter approach distance Labra and Leonard (1999) Snake species in TN, USA Urbanization and habitat fragmentation Higher fecal parasite counts Davis et al. (2012) Australian freshwater turtle (C. longicollis) Urbanization and drought Less aestivation due to increased water Rees et al. (2009) (D) Physiology Eastern fence lizard (S. undulatus) Cadmium tire byproduct Acute mortality and altered thyroid hormone Brasfield et al. (2004) Ornate tree lizard (U. ornatus) Urbanization Lower baseline and stress-induced CORT levels and altered leukocyte counts French et al. (2008) Side-blotched lizard (U. stansburiana) Urbanization Higher CORT response, reproductive investment, and oxidative stress; lower survival, innate immunity, and antioxidants Lucas and French (2012) Anoles (A. sagrei and A. cristatellus) Temperature Higher urban temperatures accelerated development of non-native anole embryos Tiatragul et al. (2017) Eurpoean wall lizard (P. muralis) Urbanization Increased parasite loads and reduced body condition Lazić et al. (2017) Lesser Antillean iguana (I. delicatissima) Urbanization No change in growth rate or body condition; asymptotic body condition Knapp and Perez-Heydrich (2012) Galápagos marine iguana (A. cristatus) Human development and tourism (including urban) Higher oxidative stress, lower immunity, and sex-dependent responses in CORT and sex hormones French et al. 2017) Common gartersnake (T. sirtalis) Indoxocarb pesticide Acute increase in CORT and immunity Neuman-Lee et al. (2016) Western terrestrial gartersnake (T. elegans) Polybrominated diphenyl ethers (PBDEs) flame retardants Altered thyroid morphology and increased body size of reproductive females and offspring Neuman-Lee et al. (2015) Copperhead (A. contortrix) Roads and traffic Reduced CORT response and no difference in baseline CORT Owen et al. 2014) Eastern long-necked turtle (C. longicollis) Urbanization during wet period Similar reproductive output and growth rate Ferronato et al. (2017) Eastern long-necked turtle (C. longicollis) Urbanization during dry period Increased growth rate Roe et al. (2011) Yellow-bellied slider (T. scripta scripta) Trace elements of coal combustion (cadmium, copper, and arsenic) Increased bactericidal ability and no change in PHA response or parasitism Haskins et al. (2017) Painted turtle (C. picta) Roads and traffic No differences in baseline or stress-induced CORT levels Polich (2016) Painted turtle (C. picta) Roads and traffic No differences in baseline or stress-induced CORT levels Baxter-Gilbert et al. (2014) Common snapping turtles (C. serpentina) Mercury exposure Reduced hatching success Hopkins et al. (2013) American Alligator (A. mississippiensis) Chemicals from pristine and contaminated lakes Thyroid and sex steroid hormone abnormalities in contaminated lakes; Smaller phallus sizes Crain et al. (1998); Guillette et al. (1999) (E) Survival Common blue-tongued skink (T. scincoides) Domestic pets and habitat loss Increased injury and mortality with increased domestic pets and habitat loss Koenig et al. (2002) Green anole (A. carolinensis) Interaction of temperature and pyrethrin pesticide Temperature and dose of pesticide interact to affect mortality Talent (2005) Texas horned lizard (P. cornutum) Urbanization Increased survival Endriss et al. (2007) Texas horned Lizard (P. cornutum) Urbanization Decreased survival Wolf et al. (2013) Salt marsh snake (N. clarkii compressicauda) Herbicide Decreased survival Ackley and Meylan (2010) Spiny softshell turtle (A. spinifera) Urbanization Decreased survival Plummer and Mills (2008) Eastern long-necked turtle (C. longicollis) Urbanization during wet period Similar abundance and survival Ferronato et al. (2017) Semi-aquatic turtle species in NC, USA Urbanization Species dependent survival Eskew et al. (2010) Reptile species in the Southwestern USA Urbanization and altered riparian habitat Species dependent extinctions and declines Lowe (1985) Lacertid lizard species in Poland Cats Cats killed more lizards in rural areas Krauze-Gryz et al. (2017) Skink species in Australia Habitat fragmentation Increased bird predation on the edge of fragmented remnants Anderson and Burgin (2008) Painted turtles (C. picta) and common snapping turtles (C. serpentine) Roads Sex-dependent mortality on roads changes population structure to male biased Steen and Gibbs (2004); Steen et al. (2006) (F) Genetic responses Crested anole (A. cristatellus) Urbanization Phenotypic shifts Winchell et al. (2016) Common wall lizard (P. muralis) Urbanization Decreased gene flow Beninde et al. (2016) West coast laterite ctenotus (C. fallens) Urbanization Decreased gene flow, similar genetic diversity, and no genetic differentiation Krawiec et al. (2015) Florida sand skink (Plestiodon reynoldsi) Urbanization Decreased gene flow and similar genetic diversity Richmond et al. (2009) California legless lizard (Anniella pulchra) Urbanization Similar genetic diversity Parham and Papenfuss (2009) Reticulated velvet gecko (Hesperoedura reticulata) Tree Dtella (Gehyra variegata) Urbanization Increased genetic differentiation Hoehn et al. 2007) Alameda striped racer (Coluber lateralis euryxanthus) Urbanization Decreased gene flow Richmond et al. (2016) Mexican dusky rattlesnake (Crotalus triseriatus) Urbanization Increased genetic differentiation and normal gene flow Sunny et al. (2015) Blanding's turtle (Emydoidea blandingii) Urbanization Decreased gene flow and genetic diversity Rubin et al. (2001) Ornate box turtle (Terrapene ornata) Urbanization Decreased gene flow, similar genetic diversity, and no genetic differentiation Cureton et al. (2014) Tuatara (Sphenodon punctatus) Urbanization Increased genetic differentiation Moore et al. (2008) Lizard species in Southern California, USA Urbanization Decreased gene flow, decreased genetic diversity, and increased genetic differentiation Delaney et al. (2010) Lizard species in Southern California, USA Urbanization Decreased gene flow and similar genetic diversity Thomassen et al. (2018) Notes: An organized look at original research investigating reptilian responses to urban features organized by (A) abundance and diversity, (B) diet, (C) behavior, (D) physiology, (E) survival, and (F) genetic. Within each subsection of the table, studies are ordered by taxonomic group and then followed by multispecies studies. Table 1 Reptilian responses to urban features and general outcomes General response measured Species Urban factor measured Outcome Reference (A) Abundance and diversity Green anole (A. carolinensis) Cats Cats ate a lot of lizards Loyd et al. (2013) Crested anole (A. cristatellus) Urbanization Decreased presence and abundance Kolbe et al. (2016b) Lesser Antillean iguana (I. delicatissima) Urbanization Decreased abundance and local extirpations, similar density Knapp and Perez-Heydrich (2012) Dunes sagebrush lizard (S. arenicolus) Urbanization Decreased abundance Smolensky and Fitzgerald (2011) Desert grassland whiptail (Aspidoscelis uniparens) and Lesser earless lizard (Holbrookia maculata) Urbanization Decreased abundance coincided with increased roadrunner abundance Audsley et al. (2006) Fence lizard (S. occidentalis) Urbanization More frequent tonic immobility and lower sprint speeds; Significantly shorter limbs in females. Sparkman et al. (2018) Common chuckwalla (S. ater) Urbanization Similar abundance Sullivan and Sullivan (2008) Common chuckwalla (S. ater) Urbanization Similar abundance Sullivan and Williams (2010) Northern watersnake (N. sipedon) Urbanization Altered habitat use and abundance Pattishall and Cundall (2009) Eastern long-necked turtle (C. longicollis) Urbanization during dry period Increased abundance Roe et al. (2011) Eastern long-necked turtle (C. longicollis) Urbanization Similar abundance Stokeld et al. (2014) Reptile species in southeastern Spain Population density Increased species richness Moreno-Rueda and Pizarro (2007) Reptile species in western Georgia, USA Watershed development Increased species richness Barrett and Guyer (2008) Reptile species in Oxford, UK Urbanization and habitat fragmentation Species richness decreased with increasing distance from permanent water and increased with patch size. Dickman (1987) Reptile species in Australia Urbanization Vegetation remnant size correlated positively with species number How and Dell (2000) Reptile species in South Bulgaria Urbanization Reptile species richness was highest in rural zone, 2nd highest was the urban zone, and last was the suburban zone. Mollov et al. (2009) Riparian reptile species Damming Decreased species richness and occupancy Hunt et al. (2013) Reptile species in Brisbane, Australia Urbanization Landscape structure and local scale habitat were most important for species assemblages Garden et al. (2010) Lizard species in Phoenix, AZ, USA Socioeconomic status and land cover Building cover negatively affected diversity Ackley et al. (2015b) Reptile species in Indianapolis, IN, USA Altered waterways Turtles disappeared and snakes decreased Minton (1968) Reptile species in Melbourne, Australia Urbanization Decreased presence Hamer and McDonnell (2009) Snake species in NJ, USA Urbanization Species-dependent effects Zappalorti and Mitchell (2008) (B) Diet Coastal horned lizard (P. coronatum) Invasive ants Altered prey selection in areas invaded by non-native ants Suarez et al. (2000) Dugite (P. affinis) Urbanization Smaller in mass and less likely to have food in stomach Wolfe et al. (2017) (C) Behavior Side-blotched lizard (U. stansburiana) and ornate tree lizard (U. ornatus) Temperature Urban vegetation allowed for extended lizard activity Ackley et al. (2015a) Puerto Rican crested anole (A. cristatellus) Predation Increased tail autonomy and regrowth Tyler et al. (2016) Anoles (A. sagrei and A. cristatellus) Urbanization Increased body size, longer latency to feeding when offered food, and lower overall response rates Chejanovski et al. (2017) Anoles (A. cristatellus and A. stratulus) Artifical substrates Artificial substrates slowed running speed but were still used frequently Kolbe et al. (2016a) Brown anole (A. sagrei) Urbanization More tolerant to humans, less aggressive, and spent more time exploring new habitat Lapiedra et al. (2017) Common wall lizard (P. muralis) Urbanization More asymmetric traits Lazić et al. (2015) Aegean wall lizard (P. erhardii) Human built structures Switched foraging mode in new environment Donihue (2016) Dalmatian wall lizard (P. melisellensis) Urbanization Similar risk-taking and neophobia of foraging behavior De Meester et al. (2018) Delicate skink (L. delicata) Urbanization No differences in learning metrics Kang et al. (2018) Delicate skink (L. delicata) Urbanization Similar activity, exploratory, and foraging behaviors Moule et al. (2016) Garden skink (L. guichenoti) Urbanization Greater flight initiation distance, approach distance, and sprint speed Prosser et al. (2006) Side-blotched lizard (U. stansburiana) Human presence Shorter flight initiation Keehn and Feldman (2018) South Indian rock agama (P. dorsalis) Urbanization Better body condition, less diverse diet, and altered hunting strategies Balakrishna et al. (2016) Peninsular rock agama (P. dorsalis) Urbanization Shorter flight initiation distance Batabyal et al. (2017) Gila monster (H. suspectum) Urbanization No difference in home range size and movement parameters; Population sex ratio was female-biased. Kwiatkowski et al. (2008) Blue-tongued skinks (Tiliqua spp.) Noise (decible frequency) Altered movement behavior Alarcon and Fabiola (2016) Sleepy lizard (T. rugosa) Human presence and handling Increases stride frequency for up to an hour Kerr et al. (2004) Liolaemus lizards Human presence Shorter approach distance Labra and Leonard (1999) Snake species in TN, USA Urbanization and habitat fragmentation Higher fecal parasite counts Davis et al. (2012) Australian freshwater turtle (C. longicollis) Urbanization and drought Less aestivation due to increased water Rees et al. (2009) (D) Physiology Eastern fence lizard (S. undulatus) Cadmium tire byproduct Acute mortality and altered thyroid hormone Brasfield et al. (2004) Ornate tree lizard (U. ornatus) Urbanization Lower baseline and stress-induced CORT levels and altered leukocyte counts French et al. (2008) Side-blotched lizard (U. stansburiana) Urbanization Higher CORT response, reproductive investment, and oxidative stress; lower survival, innate immunity, and antioxidants Lucas and French (2012) Anoles (A. sagrei and A. cristatellus) Temperature Higher urban temperatures accelerated development of non-native anole embryos Tiatragul et al. (2017) Eurpoean wall lizard (P. muralis) Urbanization Increased parasite loads and reduced body condition Lazić et al. (2017) Lesser Antillean iguana (I. delicatissima) Urbanization No change in growth rate or body condition; asymptotic body condition Knapp and Perez-Heydrich (2012) Galápagos marine iguana (A. cristatus) Human development and tourism (including urban) Higher oxidative stress, lower immunity, and sex-dependent responses in CORT and sex hormones French et al. 2017) Common gartersnake (T. sirtalis) Indoxocarb pesticide Acute increase in CORT and immunity Neuman-Lee et al. (2016) Western terrestrial gartersnake (T. elegans) Polybrominated diphenyl ethers (PBDEs) flame retardants Altered thyroid morphology and increased body size of reproductive females and offspring Neuman-Lee et al. (2015) Copperhead (A. contortrix) Roads and traffic Reduced CORT response and no difference in baseline CORT Owen et al. 2014) Eastern long-necked turtle (C. longicollis) Urbanization during wet period Similar reproductive output and growth rate Ferronato et al. (2017) Eastern long-necked turtle (C. longicollis) Urbanization during dry period Increased growth rate Roe et al. (2011) Yellow-bellied slider (T. scripta scripta) Trace elements of coal combustion (cadmium, copper, and arsenic) Increased bactericidal ability and no change in PHA response or parasitism Haskins et al. (2017) Painted turtle (C. picta) Roads and traffic No differences in baseline or stress-induced CORT levels Polich (2016) Painted turtle (C. picta) Roads and traffic No differences in baseline or stress-induced CORT levels Baxter-Gilbert et al. (2014) Common snapping turtles (C. serpentina) Mercury exposure Reduced hatching success Hopkins et al. (2013) American Alligator (A. mississippiensis) Chemicals from pristine and contaminated lakes Thyroid and sex steroid hormone abnormalities in contaminated lakes; Smaller phallus sizes Crain et al. (1998); Guillette et al. (1999) (E) Survival Common blue-tongued skink (T. scincoides) Domestic pets and habitat loss Increased injury and mortality with increased domestic pets and habitat loss Koenig et al. (2002) Green anole (A. carolinensis) Interaction of temperature and pyrethrin pesticide Temperature and dose of pesticide interact to affect mortality Talent (2005) Texas horned lizard (P. cornutum) Urbanization Increased survival Endriss et al. (2007) Texas horned Lizard (P. cornutum) Urbanization Decreased survival Wolf et al. (2013) Salt marsh snake (N. clarkii compressicauda) Herbicide Decreased survival Ackley and Meylan (2010) Spiny softshell turtle (A. spinifera) Urbanization Decreased survival Plummer and Mills (2008) Eastern long-necked turtle (C. longicollis) Urbanization during wet period Similar abundance and survival Ferronato et al. (2017) Semi-aquatic turtle species in NC, USA Urbanization Species dependent survival Eskew et al. (2010) Reptile species in the Southwestern USA Urbanization and altered riparian habitat Species dependent extinctions and declines Lowe (1985) Lacertid lizard species in Poland Cats Cats killed more lizards in rural areas Krauze-Gryz et al. (2017) Skink species in Australia Habitat fragmentation Increased bird predation on the edge of fragmented remnants Anderson and Burgin (2008) Painted turtles (C. picta) and common snapping turtles (C. serpentine) Roads Sex-dependent mortality on roads changes population structure to male biased Steen and Gibbs (2004); Steen et al. (2006) (F) Genetic responses Crested anole (A. cristatellus) Urbanization Phenotypic shifts Winchell et al. (2016) Common wall lizard (P. muralis) Urbanization Decreased gene flow Beninde et al. (2016) West coast laterite ctenotus (C. fallens) Urbanization Decreased gene flow, similar genetic diversity, and no genetic differentiation Krawiec et al. (2015) Florida sand skink (Plestiodon reynoldsi) Urbanization Decreased gene flow and similar genetic diversity Richmond et al. (2009) California legless lizard (Anniella pulchra) Urbanization Similar genetic diversity Parham and Papenfuss (2009) Reticulated velvet gecko (Hesperoedura reticulata) Tree Dtella (Gehyra variegata) Urbanization Increased genetic differentiation Hoehn et al. 2007) Alameda striped racer (Coluber lateralis euryxanthus) Urbanization Decreased gene flow Richmond et al. (2016) Mexican dusky rattlesnake (Crotalus triseriatus) Urbanization Increased genetic differentiation and normal gene flow Sunny et al. (2015) Blanding's turtle (Emydoidea blandingii) Urbanization Decreased gene flow and genetic diversity Rubin et al. (2001) Ornate box turtle (Terrapene ornata) Urbanization Decreased gene flow, similar genetic diversity, and no genetic differentiation Cureton et al. (2014) Tuatara (Sphenodon punctatus) Urbanization Increased genetic differentiation Moore et al. (2008) Lizard species in Southern California, USA Urbanization Decreased gene flow, decreased genetic diversity, and increased genetic differentiation Delaney et al. (2010) Lizard species in Southern California, USA Urbanization Decreased gene flow and similar genetic diversity Thomassen et al. (2018) General response measured Species Urban factor measured Outcome Reference (A) Abundance and diversity Green anole (A. carolinensis) Cats Cats ate a lot of lizards Loyd et al. (2013) Crested anole (A. cristatellus) Urbanization Decreased presence and abundance Kolbe et al. (2016b) Lesser Antillean iguana (I. delicatissima) Urbanization Decreased abundance and local extirpations, similar density Knapp and Perez-Heydrich (2012) Dunes sagebrush lizard (S. arenicolus) Urbanization Decreased abundance Smolensky and Fitzgerald (2011) Desert grassland whiptail (Aspidoscelis uniparens) and Lesser earless lizard (Holbrookia maculata) Urbanization Decreased abundance coincided with increased roadrunner abundance Audsley et al. (2006) Fence lizard (S. occidentalis) Urbanization More frequent tonic immobility and lower sprint speeds; Significantly shorter limbs in females. Sparkman et al. (2018) Common chuckwalla (S. ater) Urbanization Similar abundance Sullivan and Sullivan (2008) Common chuckwalla (S. ater) Urbanization Similar abundance Sullivan and Williams (2010) Northern watersnake (N. sipedon) Urbanization Altered habitat use and abundance Pattishall and Cundall (2009) Eastern long-necked turtle (C. longicollis) Urbanization during dry period Increased abundance Roe et al. (2011) Eastern long-necked turtle (C. longicollis) Urbanization Similar abundance Stokeld et al. (2014) Reptile species in southeastern Spain Population density Increased species richness Moreno-Rueda and Pizarro (2007) Reptile species in western Georgia, USA Watershed development Increased species richness Barrett and Guyer (2008) Reptile species in Oxford, UK Urbanization and habitat fragmentation Species richness decreased with increasing distance from permanent water and increased with patch size. Dickman (1987) Reptile species in Australia Urbanization Vegetation remnant size correlated positively with species number How and Dell (2000) Reptile species in South Bulgaria Urbanization Reptile species richness was highest in rural zone, 2nd highest was the urban zone, and last was the suburban zone. Mollov et al. (2009) Riparian reptile species Damming Decreased species richness and occupancy Hunt et al. (2013) Reptile species in Brisbane, Australia Urbanization Landscape structure and local scale habitat were most important for species assemblages Garden et al. (2010) Lizard species in Phoenix, AZ, USA Socioeconomic status and land cover Building cover negatively affected diversity Ackley et al. (2015b) Reptile species in Indianapolis, IN, USA Altered waterways Turtles disappeared and snakes decreased Minton (1968) Reptile species in Melbourne, Australia Urbanization Decreased presence Hamer and McDonnell (2009) Snake species in NJ, USA Urbanization Species-dependent effects Zappalorti and Mitchell (2008) (B) Diet Coastal horned lizard (P. coronatum) Invasive ants Altered prey selection in areas invaded by non-native ants Suarez et al. (2000) Dugite (P. affinis) Urbanization Smaller in mass and less likely to have food in stomach Wolfe et al. (2017) (C) Behavior Side-blotched lizard (U. stansburiana) and ornate tree lizard (U. ornatus) Temperature Urban vegetation allowed for extended lizard activity Ackley et al. (2015a) Puerto Rican crested anole (A. cristatellus) Predation Increased tail autonomy and regrowth Tyler et al. (2016) Anoles (A. sagrei and A. cristatellus) Urbanization Increased body size, longer latency to feeding when offered food, and lower overall response rates Chejanovski et al. (2017) Anoles (A. cristatellus and A. stratulus) Artifical substrates Artificial substrates slowed running speed but were still used frequently Kolbe et al. (2016a) Brown anole (A. sagrei) Urbanization More tolerant to humans, less aggressive, and spent more time exploring new habitat Lapiedra et al. (2017) Common wall lizard (P. muralis) Urbanization More asymmetric traits Lazić et al. (2015) Aegean wall lizard (P. erhardii) Human built structures Switched foraging mode in new environment Donihue (2016) Dalmatian wall lizard (P. melisellensis) Urbanization Similar risk-taking and neophobia of foraging behavior De Meester et al. (2018) Delicate skink (L. delicata) Urbanization No differences in learning metrics Kang et al. (2018) Delicate skink (L. delicata) Urbanization Similar activity, exploratory, and foraging behaviors Moule et al. (2016) Garden skink (L. guichenoti) Urbanization Greater flight initiation distance, approach distance, and sprint speed Prosser et al. (2006) Side-blotched lizard (U. stansburiana) Human presence Shorter flight initiation Keehn and Feldman (2018) South Indian rock agama (P. dorsalis) Urbanization Better body condition, less diverse diet, and altered hunting strategies Balakrishna et al. (2016) Peninsular rock agama (P. dorsalis) Urbanization Shorter flight initiation distance Batabyal et al. (2017) Gila monster (H. suspectum) Urbanization No difference in home range size and movement parameters; Population sex ratio was female-biased. Kwiatkowski et al. (2008) Blue-tongued skinks (Tiliqua spp.) Noise (decible frequency) Altered movement behavior Alarcon and Fabiola (2016) Sleepy lizard (T. rugosa) Human presence and handling Increases stride frequency for up to an hour Kerr et al. (2004) Liolaemus lizards Human presence Shorter approach distance Labra and Leonard (1999) Snake species in TN, USA Urbanization and habitat fragmentation Higher fecal parasite counts Davis et al. (2012) Australian freshwater turtle (C. longicollis) Urbanization and drought Less aestivation due to increased water Rees et al. (2009) (D) Physiology Eastern fence lizard (S. undulatus) Cadmium tire byproduct Acute mortality and altered thyroid hormone Brasfield et al. (2004) Ornate tree lizard (U. ornatus) Urbanization Lower baseline and stress-induced CORT levels and altered leukocyte counts French et al. (2008) Side-blotched lizard (U. stansburiana) Urbanization Higher CORT response, reproductive investment, and oxidative stress; lower survival, innate immunity, and antioxidants Lucas and French (2012) Anoles (A. sagrei and A. cristatellus) Temperature Higher urban temperatures accelerated development of non-native anole embryos Tiatragul et al. (2017) Eurpoean wall lizard (P. muralis) Urbanization Increased parasite loads and reduced body condition Lazić et al. (2017) Lesser Antillean iguana (I. delicatissima) Urbanization No change in growth rate or body condition; asymptotic body condition Knapp and Perez-Heydrich (2012) Galápagos marine iguana (A. cristatus) Human development and tourism (including urban) Higher oxidative stress, lower immunity, and sex-dependent responses in CORT and sex hormones French et al. 2017) Common gartersnake (T. sirtalis) Indoxocarb pesticide Acute increase in CORT and immunity Neuman-Lee et al. (2016) Western terrestrial gartersnake (T. elegans) Polybrominated diphenyl ethers (PBDEs) flame retardants Altered thyroid morphology and increased body size of reproductive females and offspring Neuman-Lee et al. (2015) Copperhead (A. contortrix) Roads and traffic Reduced CORT response and no difference in baseline CORT Owen et al. 2014) Eastern long-necked turtle (C. longicollis) Urbanization during wet period Similar reproductive output and growth rate Ferronato et al. (2017) Eastern long-necked turtle (C. longicollis) Urbanization during dry period Increased growth rate Roe et al. (2011) Yellow-bellied slider (T. scripta scripta) Trace elements of coal combustion (cadmium, copper, and arsenic) Increased bactericidal ability and no change in PHA response or parasitism Haskins et al. (2017) Painted turtle (C. picta) Roads and traffic No differences in baseline or stress-induced CORT levels Polich (2016) Painted turtle (C. picta) Roads and traffic No differences in baseline or stress-induced CORT levels Baxter-Gilbert et al. (2014) Common snapping turtles (C. serpentina) Mercury exposure Reduced hatching success Hopkins et al. (2013) American Alligator (A. mississippiensis) Chemicals from pristine and contaminated lakes Thyroid and sex steroid hormone abnormalities in contaminated lakes; Smaller phallus sizes Crain et al. (1998); Guillette et al. (1999) (E) Survival Common blue-tongued skink (T. scincoides) Domestic pets and habitat loss Increased injury and mortality with increased domestic pets and habitat loss Koenig et al. (2002) Green anole (A. carolinensis) Interaction of temperature and pyrethrin pesticide Temperature and dose of pesticide interact to affect mortality Talent (2005) Texas horned lizard (P. cornutum) Urbanization Increased survival Endriss et al. (2007) Texas horned Lizard (P. cornutum) Urbanization Decreased survival Wolf et al. (2013) Salt marsh snake (N. clarkii compressicauda) Herbicide Decreased survival Ackley and Meylan (2010) Spiny softshell turtle (A. spinifera) Urbanization Decreased survival Plummer and Mills (2008) Eastern long-necked turtle (C. longicollis) Urbanization during wet period Similar abundance and survival Ferronato et al. (2017) Semi-aquatic turtle species in NC, USA Urbanization Species dependent survival Eskew et al. (2010) Reptile species in the Southwestern USA Urbanization and altered riparian habitat Species dependent extinctions and declines Lowe (1985) Lacertid lizard species in Poland Cats Cats killed more lizards in rural areas Krauze-Gryz et al. (2017) Skink species in Australia Habitat fragmentation Increased bird predation on the edge of fragmented remnants Anderson and Burgin (2008) Painted turtles (C. picta) and common snapping turtles (C. serpentine) Roads Sex-dependent mortality on roads changes population structure to male biased Steen and Gibbs (2004); Steen et al. (2006) (F) Genetic responses Crested anole (A. cristatellus) Urbanization Phenotypic shifts Winchell et al. (2016) Common wall lizard (P. muralis) Urbanization Decreased gene flow Beninde et al. (2016) West coast laterite ctenotus (C. fallens) Urbanization Decreased gene flow, similar genetic diversity, and no genetic differentiation Krawiec et al. (2015) Florida sand skink (Plestiodon reynoldsi) Urbanization Decreased gene flow and similar genetic diversity Richmond et al. (2009) California legless lizard (Anniella pulchra) Urbanization Similar genetic diversity Parham and Papenfuss (2009) Reticulated velvet gecko (Hesperoedura reticulata) Tree Dtella (Gehyra variegata) Urbanization Increased genetic differentiation Hoehn et al. 2007) Alameda striped racer (Coluber lateralis euryxanthus) Urbanization Decreased gene flow Richmond et al. (2016) Mexican dusky rattlesnake (Crotalus triseriatus) Urbanization Increased genetic differentiation and normal gene flow Sunny et al. (2015) Blanding's turtle (Emydoidea blandingii) Urbanization Decreased gene flow and genetic diversity Rubin et al. (2001) Ornate box turtle (Terrapene ornata) Urbanization Decreased gene flow, similar genetic diversity, and no genetic differentiation Cureton et al. (2014) Tuatara (Sphenodon punctatus) Urbanization Increased genetic differentiation Moore et al. (2008) Lizard species in Southern California, USA Urbanization Decreased gene flow, decreased genetic diversity, and increased genetic differentiation Delaney et al. (2010) Lizard species in Southern California, USA Urbanization Decreased gene flow and similar genetic diversity Thomassen et al. (2018) Notes: An organized look at original research investigating reptilian responses to urban features organized by (A) abundance and diversity, (B) diet, (C) behavior, (D) physiology, (E) survival, and (F) genetic. Within each subsection of the table, studies are ordered by taxonomic group and then followed by multispecies studies. Habitat fragmentation also plays an important role in species richness of herpetofauna (Dickman 1987; Irwin et al. 2010). Patch size in particular has repeatedly been shown as an important predictor of reptile species richness in many different environments (Dickman 1987; Garden et al. 2007; 2010) and of species evenness (i.e., species diversity) in others (Sullivan et al. 2014). For example, remnants in metropolitan Perth, Australia were found to be an important predictor of species richness with bigger remnants having more reptile species (How and Dell 2000). Similarly, by using specialized habitats in NJ, USA (i.e., Pine Barrens) some species of snakes are able to thrive (Zappalorti and Mitchell 2008). Importantly, many of these studies show that maintaining structural complexity and remnant habitat patches can help protect some species (Hamer and McDonnell 2008). The effects of patch size appear to be species dependent, whereby habitat fragmentation in the Midwestern United States adversely effects some species of turtles (red-eared sliders, Trachemys scripta elegans) more than other species (e.g., midland painted turtles, Chrysemys picta marginata, eastern spiny softshells, Apalone spinifera spinifera, common snapping turtles, Chelydra serpentina serpentina) (Rizkalla and Swihart 2006; Ryan et al. 2014). The vast majority of work, however, demonstrates a negative impact of urbanization on reptiles. For example, Lowe found that encroaching urbanization was associated with altered riparian habitat and the extinction of several reptiles (e.g., Thamnophis species) in the American southwest (1985). These effects are seen worldwide, as for example, reptile species richness in Bulgaria was found to be highest in rural zones and followed by urban and suburban zones (Mollov et al. 2009; Mollov 2011). Many other studies also document apparent adverse effects of urbanization on reptile abundance and species richness (Table 1; Ackley et al. 2009; Banville and Bateman 2012; Hunt et al. 2013; Sullivan et al. 2017), whereby development beyond moderate levels leads to decreased number of species and abundance for those that remain (Germaine and Wakeling 2001). Modification of aquatic habitat in IN, USA, also led to declines in both turtles and snakes from the area (Minton 1968). Finally, by comparing wildlife databases of herpetofauna, researchers demonstrated that reptiles are negatively impacted by urbanization (Hamer and McDonnell 2008). While distribution studies are a critical first step in determining what species are most impacted, an important next step is to begin investigating specific factors within the urban environment that may be influencing reptile species using a combination of field and controlled laboratory studies. Biotic factors Human presence and invasive species In general, urbanization tends to decrease native species richness but promotes diversity of exotic and/or non-native species (McKinney 2006; 2008). Because urbanization can promote the establishment of non-native and generalist species, there is the potential for urbanization to influence species interactions through the introduction of novel predators, prey, and parasites. The number of invasive species is significantly increasing and thought to be one of the predominate risks for native species (Pimentel et al. 2005). For example, the presence of native crested anoles (Anolis cristatellus) is negatively associated with the presence of a non-native competitor species, brown anoles (Anolis sagrei). Here, brown anoles not only reduced the abundance of crested anoles, but also caused a shift in their use of perches (Kolbe et al. 2016b). Humans are one of the predominant invasive species present in urban environments that have been shown to alter reptilian behavior and movement patterns. Lizards from areas of high human density have shorter approach distances before initiating flight compared to lizards from areas of low human density (Labra and Leonard 1999). Marine iguanas close to towns come in more frequent contact with people including tourists in the Galapagos Islands and also show behavioral and physiological responses to intensity of exposure, including elevated stress reactivity, oxidative stress, and suppressed immunity (French et al. 2010; 2017). Another study found that lizards which have been observed, briefly handled, or extensively handled all increased stride frequency following exposure to humans (Kerr et al. 2004). While allowing humans to get closer before running away could be a result of habituation to human presence, increased stride frequency would suggest an increased cost of movement associated with human presence. While humans are the leading factor altering environments for native species, the introduction of other invasive species can also alter the landscape and ecology, having significant implications for native species. There is evidence of increased predation for reptiles in urbanized areas primarily by invasive species and pets (Fischer et al. 2012). Household pets, such as domestic cats, are known to prey upon local fauna. For example, video monitoring of free-roaming cats in southeastern United States showed that 33% of prey recovered from outdoor cats were reptiles, specifically the green anoles (Anolis carolinensis) (Loyd et al. 2013). Similarly, Koenig et al. (2002) found that blue-tongued skinks (Tiliqua scincoides) were more likely to be attacked by household pets in suburban areas compared to urban areas. However, Krauze-Gryz et al. (2017) found that reptiles in Poland were more likely to be predated by cats in rural areas. With native predators, skinks were more likely to be predated by birds on the edge of fragmented remnants compared to the core (Anderson and Burgin 2008). Similarly, in southeast Arizona, terrestrial lizard abundance decreased in urban areas along with an increase in roadrunners abundance, a common natural predator (Audsley et al. 2006), suggesting a causal link. These results provide evidence to suggest that urbanization generally increases chances of predation from both native and non-native predators. Diet Urbanization can affect reptile diet composition and frequency of feeding by altering the availability of native food sources and introducing non-native prey species (Table 1B), which, in some instances, can be beneficial. In the case of the threatened Lake Erie Water Snake (Nerodia sipedon insularum), the invasive round goby (Neogobius melanostomus) is an important food source allowing for increased growth rates and body size which can reduce predation risk during vulnerable developmental stages (King et al. 2006). Similarly, Balakrishna et al. (2016) found that Indian rock agamas (Psammophilus dorsalis) in urban environments had better body condition, had a less diverse diet, and even altered their foraging strategy compared to rural conspecifics. Not all species perform better on urban diets or with novel diet sources. For example, a study comparing diet of road-killed and museum-collected specimens showed that dugites (Pseudonaja affinis) occupying urban areas in Australia were less likely to contain a meal and were smaller in mass compared to their rural counterparts (Wolfe et al. 2017). Suarez et al. (2000) found that invasive Argentine ants (Linepithema humile) originating from urban areas displaced native ant species and significantly altered the diet composition for coastal horned lizard (Phrynosoma coronatum). Furthermore, ontogenetic differences in diet suggest the need for a diverse ant community to sustain populations, and raise concern that the documented decline in native ant species and diversity through displacement by the Argentine ant could potentially affect survival and population persistence of many ant predator species. Feeding behaviors may also differ for lizards in urban versus forested environments. Anolis lizards in urban environments of Puerto Rico have been observed to be larger and to have longer latency to feeding when offered food (Chejanovski et al. 2017). Some lizards have even switched foraging modes in response to habitat changes occurring with human presence. Aegean wall lizards (Podarcis erhardii) which utilized rock walls were more sedentary, exhibited morphological changes, and ate less sedentary prey compared to non-wall lizards (Donihue 2016). Parasites Urbanization has the potential to influence immunity and host–pathogen dynamics of urban-dwelling animals via the introduction of non-native parasites and pathogens along with other larger invasive species (Martin et al. 2010). Furthermore, changes in general ecology, including habitat size and fragmentation, can also alter disease transmission in urban habitats (Riley et al. 2014b). For example, Davis et al. (2012) found that more snakes had fecal parasites near the outer edges of an urban forest compared to snakes near the core of the forest. Similarly, Lazić et al. (2017) found that wall lizards (Podarcis muralis) had higher parasite loads and reduced body condition in urban areas compared to rural areas. These studies suggest that urbanization can potentially influence pathogen transmission among reptiles occupying previously natural habitat and adjacent areas. However, more studies are needed to further elucidate how urban cover influences transmission of parasites and susceptibility to reptile species. Abiotic factors Temperature, light, and noise Urbanization can alter abiotic features of an environment, such as temperature, light, and noise, but the direct impacts of these changes on animals is not well understood. Temperature is a dominant ecological variable for all animals that can be altered in urban environments. The majority of studies on temperature changes in urban environments have focused on endothermic species. However, known temperature changes in urban areas likely render ectotherms even more susceptible to urbanization. Decreased shade cover from plants can cause reptiles to be less active due to intense heat. Ackley et al. (2015a) demonstrated that irrigated and non-native shade planting increased lizard activity time in urban desert relative to native landscaping. Tiatragul et al. (2017) found that urban temperatures not only accelerated development of non-native anole embryos, but that non-native embryos were robust and survived well under urban temperatures. Temperature also has the potential to exacerbate other environmental stressors. For example, Talent (2005) found that temperature influenced the sensitivity of lizards to pyrethrin pesticides. Similar to temperature, light and photoperiod are critical for the timing of important life history events. Artificial light has also become a ubiquitous factor for most urban environments and while several studies have focused on birds (da Silva et al. 2014; Ouyang et al. 2017), the studies for reptile species are largely inconclusive or lacking (Perry et al. 2008). It has been suggested that no site in the continental United States is free from anthropogenic noise exposure, including remote protected areas such as national parks (Barber et al. 2011). Several studies have directly tested the effects of human noise on bird behavior and physiology (Rheindt 2003; Swaddle and Page 2007; Francis et al. 2009; Slabbekoorn 2013; Davies et al. 2017) and male tree frog (Hyla arborea) calling (Lengagne 2008) but few other animals have been studied this extensively. Alarcon and Fabiola (2016) tested different decibels and frequencies on behavior in blue-tongued skinks (T.scincoides) and found that loud, especially high frequency, noises resulted in animals spending more time freezing, a typical stress response in reptiles. Substrates and roads Some abiotic urban features may actually be beneficial to animal inhabitants, by providing access to more diverse substrates (e.g., greater refuge and perching options) and resources, allowing reptiles to persist under anthropogenic disturbances. Evidence for this has emerged through early work on comparative habitat preferences across the urban–rural landscape. For example, northern watersnakes (N.sipedon) occupying urbanized stream areas exhibit significantly greater site fidelity than those found in natural stream areas (Pattishall and Cundall 2008). Snakes in natural areas selected habitat with wide riparian zones and dense canopy cover, whereas snakes in urban areas more often occupied artificial substrates (e.g. piles of scrap metal, concrete, or holes in a railroad bed adjacent to streams) and areas with high human density (Pattishall and Cundall 2009). Artificial structures (e.g. broader and smoother substrates) are extensively utilized for perching and refuge among lizard species such as garden skinks (Lampropholis guichenoti), blue-tongued skinks (T.scincoides), crested anoles (A.cristatellus), and Gila monsters (Heloderma suspectum) (Koenig et al. 2001; Prosser et al. 2006; Winchell et al. 2016), whereas other species such as barred anoles (Anolis stratulus) tend to use more natural aspects of the urban environment (i.e., trees and other cultivated vegetation; (Winchell et al. 2018)). Artificial substrates also tend to be smoother than natural substrates for arboreal species which can impact running velocity as was demonstrated in two species of Anolis lizards (Kolbe et al. 2016a). This may in part explain why lizards display differing flight initiation distances, escape strategies, and performance levels from rural counterparts (Koenig et al. 2001; Prosser et al. 2006; Aviles-Rodriguez 2015; Winchell et al. 2016). As would be expected, urban changes in hydrology most significantly impact aquatic or riparian species, especially turtles. Rees et al. (2009) found that Australian freshwater turtles alter their behavior and are less likely to aestivate because the water supply does not seasonally dry up in urban areas. However, damming, which greatly alters riparian landscapes, reduces reptile occupancy and richness for many species, not only aquatic (Hunt et al. 2013). Perhaps most significantly, soil moisture in all habitats is critical for the development of reptile embryos of oviparous species, which constitutes the vast majority of reptile species (Ackerman 1991). Roads are a notable abiotic factor associated with urban environments that introduce changes in substrate, noise, and disturbance rates, which may in themselves also be a direct source of mortality (Ashley and Robinson 1996; Jochimsen et al. 2004; Andrews and Gibbons 2005; Andrews et al. 2008; Andrews et al. 2015). A synthesis of studies investigating the impact of roads on reptile abundance demonstrates generally negative impacts, as does a meta-analysis of life history traits and population responses to roads (Fahrig and Rytwinski 2009; Rytwinski and Fahrig 2012). Among snake species, those of smaller sizes such as ring-necked snakes (Diadophis punctatus), southeastern crown snakes (Tantilla coronata), and eastern hognose snakes (Heterodon platirhinos) more often avoid the presence of roads and traffic (Andrews and Gibbons 2005; Andrews et al. 2008; Robson and Blouin-Demers 2013). However, larger species such as gartersnakes (Thamnophis sirtalis parietalis), coachwhips (Masticophis flagellum), and brown tree snakes (Boiga irregularis) can exhibit alternative movement activities and pathways to avoid exposure and remain in natural habitats (Shine et al. 2004; Mitrovich et al. 2009; Siers et al. 2014). Similarly, turtle species including common snapping turtles and eastern painted turtles (Chrysemys picta picta) disperse into urbanized habitat less often than natural habitat, as they also exhibit avoidance behaviors toward areas with higher densities of roads (Patrick and Gibbs 2010). However, some species (Chrysemys picta and Chelydra serpentine) show sex-dependent differences in road mortality so much so that there are changes in population structure (Steen and Gibbs 2004). The sex-biased effect of roads seems to be driven by females nesting migrations that make them more likely to cross roads and be killed (Steen et al. 2006). Lizard species including blue-tongued skinks (T.scincoides), western fence lizards (Sceloporus occidentalis), orange-throated whiptails (Aspidoscelis hyperythra), and dunes sagebrush lizards (Sceloporus arenicolus) have been found to actively avoid crossing roads and instead utilize vegetation and other natural substrates for movement (Koenig et al. 2001; Brehme et al. 2013; Hibbitts et al. 2017). While the risk of mortality is reduced by road and traffic avoidance, changes in movement patterns and spatial distributions can contribute to genetic isolation and population sinks for reptiles (Forman and Alexander 1998; Shepard et al. 2008). However, the field of road ecology is growing and working to provide new planning strategies to mitigate the impacts on animals (Langen et al. 2012; Riley et al. 2014a). Pollution In many urban landscapes and adjacent developed areas, environmental toxin levels and air particulates are higher than surrounding rural areas (Cohen et al. 2004; Wei and Yang 2010). The few studies that test ecotoxicological outcomes of urban pollutants in reptiles demonstrate that the effects are not necessarily harmful. For example, yellow-bellied sliders (Trachemys scripta scripta) accumulate trace elements from coal combustion such as cadmium, copper, and arsenic as they grow, but these elements do not seem to adversely impact their immune systems measured via parasitism and responses to phytohemagglutinin, although bacterial killing ability was elevated in turtles from contaminated sites (Haskins et al. 2017). In gartersnakes (T.sirtalis), exposure to the pesticide indoxocarb induced an acute stress increase in corticosterone and immunity, whereas exposure to a similar natural toxin to which the gartersnakes have evolved resistance (i.e., tetrodoxin) did not induce a physiological response (Neuman-Lee et al. 2016). Moreover, exposure to polybrominated diphenyl ethers (PBDEs) which are used as flame retardants and are persistent contaminants found in practically every environment and organism tested, resulted in altered thyroid follicular height in female gartersnakes (Thamnophis elegans), suggesting thyroid dysfunction (Neuman-Lee et al. 2015). Neuman-Lee et al. (2017) also found an increase in body size of pregnant female gartersnakes exposed to PBDEs as well as their resulting offspring. Brasfield et al. (2004) demonstrated that exposure to cadmium, a byproduct of tire ware that is likely high in urban settings, could result in acute mortality and thyroid dysfunction in developing Eastern fence lizards (Sceloporus undulatus). Talent (2005) demonstrated that temperature influenced the sensitivity of green anoles to pyrethrin pesticides. Given that urbanization is known to alter ambient temperature and there are more pesticides in use in human-altered landscapes, this has important implications for urban reptiles. It is important to note that the effects of urban pollution can be wider reaching that just the urban footprint itself. American alligators (Alligator mississippiensis) from lakes contaminated with municipal and agricultural runoff show altered thyroid and sex steroid hormone levels and smaller phalluses (Crain et al. 1998; Guillette et al. 1999). Western pond turtles (Emys marmorata) from protected areas in California still show signs of both current- and historic-use pesticides in their blood (Meyer et al. 2016). Turtles in southwest VA, USA also show evidence of higher blood mercury levels when sampled at contaminated sites and depending on their feeding strategy (Bergeron et al. 2007). In common snapping turtles (C.serpentina), mercury levels were associated with reduced hatching success (Hopkins et al. 2013). This contamination in aquatic settings can also pass to species that prey upon aquatic animals, as observed in a viperine snake (Natrix maura) that preys on fish in France and has high mercury levels as a result (Lemaire et al. 2018). Yet, the overall evidence as to the effects of urban and anthropogenic pollutants on reptiles is limited and more research is needed (Croteau et al. 2008). In particular, researchers suggest that major ecotoxicological gaps for reptiles include better understanding the magnitude and mechanism of contaminant exposure (Weir et al. 2010; Riley et al. 2014b). Directionality of responses Individual responses In order to estimate the impact of urbanization on reptiles, understanding the directionality of how different species respond to the stressors of environmental change is pivotal. Measuring individual level responses can effectively provide real-time information concerning organismal viability in a particular environment, whereas population-level censuses may require long periods of time to yield insight. A large body of work has amassed in assessing individual responses to urbanization across several taxa, yet few studies thus far have included reptiles. Emerging findings suggest the impact of anthropogenic disturbance likely depends on habitat requirements and life history, whereby directionality for individual responses is either relatively consistent or species-specific. Behavior and morphology One of the main mechanisms through which animals respond to changing environmental conditions is by adjusting modes of behavior (Réale et al. 2007; Miranda et al. 2013; Sol et al. 2013). Differences in behavioral traits among urban and rural environments can either result from individual behavioral plasticity or microevolutionary changes (Miranda et al. 2013). Whether behavioral responses to urbanization rely on acclimation or adaptation in reptiles remains largely undetermined (Kang et al. 2018). Regardless, the directionality of behavioral responses to urbanization may be associated with life history strategies most appropriate for coping with changing environmental conditions (Huey et al. 2003; Sol and Maspons 2016; Sol et al. 2018). Across avian and mammalian taxa, urban and rural conspecifics generally vary in temperament, whereby behaviors involving neophobia or neophilia, exploration, aggression, and risk perception tend to differ (Miranda et al. 2013; Sol et al. 2013). Behavioral comparisons of urban and rural reptiles have so far been limited, but relatively consistent patterns of temperament shifts may be applicable to reptile species with similar habitat requirements and life histories (Table 1C). Of critical importance is understanding whether behavioral adjustments are occurring within a species and if such changes are beneficial in terms of survival and reproduction. Emerging studies suggest that at least some reptiles are more tolerant of particular anthropogenic factors. For example, brown anoles (A.sagrei) and crested anoles (A.cristatellus) from urban areas exhibit prolonged exploratory and foraging behaviors of new environments, as well as decreased risk perception and response rates towards predator and to human presence (Chejanovski et al. 2017; Lapiedra et al. 2017). Increased tolerance to urbanization is also evident in Indian rock agamas (P.dorsalis) and side-blotched lizards (Uta stansburiana) which exhibit decreased flight initiation distances or risk perception to anthropogenic stimuli (Batabyal et al. 2017; Keehn and Feldman 2018). However, Prosser et al. (2006) found that urban garden skinks (L.guichenoti) instead flee at a greater approach distance and exhibit greater sprint speed than conspecifics from natural habitats. Meanwhile, other studies yielded no behavioral response to urbanization including movement, exploratory, and foraging behaviors of delicate skinks (Lampropholis delicata) in the Sydney region (Moule et al. 2016). Overall activity of Gila monsters (H.suspectum) did not differ among rural and urban areas (Kwiatkowski et al. 2008). Similarly, risk-taking and neophobia of foraging behavior was not affected by urbanization in Dalmatian wall lizards (Podarcis melisellensis). Collectively, these findings are relatively in line with avian and mammal species that exhibit temperaments with more neophilic, exploratory, aggressive, and risk-taking behaviors in urban areas than in rural areas (Miranda et al. 2013; Sol et al. 2013; Greenberg and Holekamp 2017). Directionality of temperament shifts to urbanization may thus depend on whether anthropogenic conditions are beneficial, innocuous, or detrimental to the habitat requirements of a given reptile species. Assessments of habitat selection and use have yielded preliminary evidence of how urban environments may meet the habitat requirements of some species, but may fail to do so for others (also see “Substrates and Roads” section above). For example, in the same urban environment, the barred anole (A.stratulus) utilizes more natural habitat compared to the crested anole (A.cristatellus), which utilizes more anthropogenic structures (Winchell et al. 2018). Additional work has demonstrated that urban crested anoles jump from perch to perch less than rural conspecifics, which instead move around more frequently on a given perch (Aviles-Rodriguez 2015). Such preferences likely depend on the degree of similarity in environmental conditions between both habitat types as this is predicted to determine the magnitude of selective pressures for behavioral differences that may be associated with urbanization. Lastly, it is important to consider that some of these behavioral changes may be the result of morphological shifts in response to urbanization, especially in species with short generation times. A recent study on the effects of urbanization on antipredator behaviors of fence lizards found urban environments to be associated with shorter limbs, lowered sprint speed, and more frequent tonic immobility (Sparkman et al. 2018). Interestingly, certain aspects of morphology have been found to be overall smaller (e.g., head size) and more asymmetric in urban common wall lizards, suggesting divergent size–shape allometries from those in rural environments (Lazić et al. 2013; 2015). However, other morphological components have been found to be larger, including limb length in urban agamid lizards (Lophognathus temporalis) and crested anoles, and greater subdigital lamellae (i.e., footpad scales) in urban crested anoles (Iglesias et al. 2012; Winchell et al. 2016). Urban brown anoles and crested anoles both tend to exhibit greater body sizes (i.e., snout-vent lengths and masses) and body condition than those in natural environments (Chejanovski et al. 2017; Hall and Warner 2017). Yet others find no difference in body condition or growth rates, such as the lesser Antillean iguanas (Iguana delicatissima) (Knapp and Perez-Heydrich 2012). Finally, Tyler et al. (2016) found greater rates of tail autonomy and regrowth in urban anoles than their rural counterparts. In the case of all of these morphological changes, there is the potential for downstream effects on behavior or locomotion, and for the animal to incur inherent costs. Physiology Just as with behavioral research, studies investigating physiological responses to urbanization also yield mixed results (Table 1D). One common metric utilized across studies is the endocrine stress response involving activation of the hypothalamic–pituitary–adrenal axis (Saplosky 1992) and ultimately the release of glucocorticoids (i.e., corticosterone in the case of reptiles; CORT) (Moore and Jessop 2003). As compared to reptiles occurring in natural habitat, those residing in urbanized areas have been found to exhibit either similar or contrasting levels of baseline stress and stress reactivity. When considering stress physiology in snakes, no difference in baseline levels of CORT is evident in copperheads (Agkistrodon contortrix) residing in forests compared to urbanized habitat (i.e., road development and traffic) (Owen et al. 2014). Similarly, northwestern garter snakes (Thamnophis ordinoides) generally do not exhibit blood heterophil-lymphocyte ratios, indicative of chronic stress, in urbanized habitat (i.e., increased human and predator presence) (Bell 2013). Further, urban snakes, such as copperheads, exhibit reduced stress-induced CORT levels compared to forest conspecifics (Owen et al. 2014). However, copperheads in urbanized areas demonstrate a negative association between anthropogenic activity and baseline, stress-induced, and magnitude of CORT response (Owen et al. 2014). Similar trends are evident in turtles, as no differences in baseline or stress-induced CORT levels were found in painted turtles exposed to urban features (i.e., road development and traffic) as compared to those in natural areas (Baxter-Gilbert et al. 2014; Polich 2016). Stress physiology in lizard species, however, exhibits dissimilarities in response to urbanized areas. For example, tree lizards (Urosaurus ornatus) inhabiting Phoenix, AZ, USA have lower baseline and stress-induced levels of corticosterone than their rural counterparts, suggesting they have habituated to urban living (French et al. 2008). These same animals also show evidence of elevated immunity (i.e., higher leukocyte counts) perhaps to deal with increased incidence of wounding in the city (2008). However, side-blotched lizards (U.stansburiana), a not too distant relative of the tree lizard, show differential responses to city life in Saint George, UT, USA. Lucas and French (2012) found both increased corticosterone response to a stressor and elevated oxidative stress in urban side-blotched lizards. However, these same urban animals also have lower immunity (bacterial killing ability) and higher reproductive investment relative to rural side-blotched lizards. Laboratory studies on these same urban lizards demonstrated that there is direct competition for protein resources between the eggs and immunity in reproductive females (Durso and French 2018), and that immune-challenged lizards alter their energetic strategy by down-regulating metabolism (Smith et al. 2017). Taken together these results suggest urbanization may be causing a life history shift in investment from self-maintenance to reproduction, a viable strategy in a short-lived reptile (Smith and French 2017). Finally, in this context, the degree to which phenotypic plasticity, genetic evolution, or a combination thereof may underlie differences in physiology, behavior, and morphology among populations across the urban–rural landscape remains unclear. Regardless, significant changes in physiology, behavior, and morphology in urban reptiles should yield the potential to induce long-lasting effects on population size and performance over time. Population responses Although aspects of urbanization are known to place reptile populations directly at risk, whether individual responses result in additional threats to population viability is largely undetermined. Changes in individual physiology and behavior in response to urbanization can affect survival and reproduction, and thus ultimately affect populations. Linking physiological and behavioral measures to demographic parameters may elucidate undetected effects of urbanization, yet few studies have pursued such endeavors (e.g., Lucas and French 2012). Of upmost concern thus far has been the abundance of reptile populations, as individual survival and reproduction often vary with respect to biotic and abiotic factors of the urban-rural landscape (Table 1A, 1E). Anthropogenic impacts generally appear to have neutral effects, and in some cases, positive effects on the population dynamics of semi-aquatic reptiles, although detrimental effects may arise under severe cases of habitat disturbance. Despite close proximities to anthropogenic activity, abundances of northern water snake populations were similar to those in rural environments (Pattishall and Cundall 2009). Such findings are thought to be due to variable refuge and thermal opportunities provided by both terrestrial and aquatic features of the urban environment. Greater complexity and stability in urban habitats may also explain how eastern long-necked turtles (Chelodina longicollis) maintain and even increase survival, reproductive output, and population abundance in spite of temporal fluctuations in environmental conditions (Roe et al. 2011; Stokeld et al. 2014; Ferronato et al. 2017). This seems to be congruent with high survival estimates for other turtles, such as yellowbelly sliders, common snapping turtles, and spiny softshell turtles (A.spinifera), although eastern mud turtles (Kinosternon subrubrum) tend to exhibit lower survival estimates (Eskew et al. 2010; Plummer and Mills 2008). Populations of mangrove salt marsh snakes (Nerodia clarkii compressicauda) in St. Petersburg, FL, USA were also higher in abundance in anthropogenic habitats until their decline after severe disturbance (Ackley and Meylan 2010). Interestingly terrestrial and aquatic populations tend to differ in their responses. Populations of terrestrial reptiles instead seem to be more variable in their sensitivity to anthropogenic perturbations compared to semi-aquatic species. This may be due to relative differences in the urban modifications of aquatic versus terrestrial habitats (e.g., varying facets, intensities, and frequencies) or differences in survey methods and species detection rates. For example, population abundance is often related to the degree of fragmentation, size, and quality of habitat. However, different species have particular ecological requirements in habitats that determine the directionality of response. Urban disturbance in the form of increasingly fragmented landscapes often causes fast declines and local extirpations in reptile populations, such as in the lesser Antillean iguana (I.delicatissima) (Knapp and Perez-Heydrich 2012). In the case of Texas horned lizards (Phrynosoma cornutum) in central Oklahoma, urban development caused declines in the abundance due to increased mortality despite moderate reproductive output (Endriss et al. 2007; Wolf et al. 2013). Populations of dunes sagebrush lizards vary significantly in abundance and this variation could be explained by habitat patch size and quality, which are affected by human development, oil, and gas industry (Smolensky and Fitzgerald 2011). Similarly, declines in population abundance for the invasive crested anole (A.cristatellus) in Miami, FL, USA are strongly associated with losses in habitat size and quality (Kolbe et al. 2016b). In other urban habitats with limited refuge and plant food density, population abundance of common chuckwallas (Sauromalus ater) is dependent on the availability of plant food diversity (Sullivan and Sullivan 2008; Sullivan and Williams 2010). Collectively, these studies suggest that urbanization can lead to population level changes but that some species, and perhaps environments, are more sensitive. Over multiple generations, differences in the selective pressures of an environment and the resulting physiological and population-level changes could potentially lead to genetic differentiation. Genetic Urbanization may have genetic consequences among reptile populations, as land-use changes can lead to intense forms of habitat alteration (Table 1F). Herpetofauna that are particularly sensitive to habitat degradation can rapidly become extirpated from urban locales (Gibbon et al. 2000; Cushman 2006; Hamer and McDonnell 2009). For those that persist across fragmented urban landscapes, connectivity and gene flow between populations may be hindered or inhibited, which can ultimately lead to reductions in genetic diversity, inbreeding depression, and even local extinction (Reed et al. 2002; Reed et al. 2003; Reed 2004; Cushman 2006; Frankham 2006). Degraded and fragmented habitats are also less likely to be recolonized by extinguished species. Additionally, the loss of genetic diversity in remaining populations can reduce adaptive potential in response to environmental changes. Although there are population and behavioral studies to document thriving and mobile urban reptiles species, preliminary genetic data provides little evidence of continuous gene flow for various reptile species located along fragmented urban landscapes (e.g., Delaney et al. 2010; Krawiec et al. 2015; Beninde et al. 2016; Richmond et al. 2016; Thomassen et al. 2018). Inhibited or decreased gene flow, in turn, seems to either reduce genetic diversity levels (Rubin et al. 2001; Delaney et al. 2010) or yield no effect (Parham and Papenfuss 2009; Richmond et al. 2009; Cureton et al. 2014; Krawiec et al. 2015; Sunny et al. 2015). There are often remarkable amounts of genetic divergence of populations across the urban–rural landscape, which may be occurring over relatively short geographic and temporal scales (Hoehn et al. 2007; Moore et al. 2008; Delaney et al. 2010; Sunny et al. 2015). Implications for population genetics may thus depend on the size, configuration, age, and isolation of habitat fragments. Studies are revealing evidence of selection for divergent phenotypes and suggest a potential for reptile populations to adapt to urban environments (Winchell et al. 2016; 2018). Conclusions Overall, this review has identified complex and diverse results that are variable both within and among all scales of ecological organization. There is variability in the directionality of the responses to urbanization, whereby some studies find inert or even positive effects of urbanization, while others show the opposite. This discrepancy is due in part to the heterogeneity of urban landscapes and to the fact that species responses are also different. Another important factor leading to inconsistencies in study outcomes is that urbanization includes many different factors interacting simultaneously. While there are a handful of studies testing the interactive effects of urban qualities, most are focused on non-reptile taxonomic groups (Isaksson 2015). With a growing body of studies investigating specific urban features or stressors, it is becoming increasingly important to measure interactive effects (Talent 2005), physiological responses, and multiple health indicators simultaneously. Testing only one endpoint can produce misleading results due to tradeoffs which can occur between physiological systems (Lucas and French 2012). Also, the large number of different approaches used in these studies make it difficult to compare outcomes and to assess directionality of the response. Given the methodologies and results of most experiments, we can only assess whether or not there is an effect of the urban environment or stressor, and not the fitness implications of that given effect. To conclude, based on our findings there is an apparent lack of urban research in reptilian species (1) investigating interactive or additive urban factors which more accurately represent the reality of urbanization; (2) measuring multiple morphological, behavioral, and physiological responses, because a more comprehensive approach will allow researchers to better assess directionality; (3) linking individual to population-level responses to identify the mechanisms for population changes, especially declines, and (4) testing genetic/genomic differences across an urban environment as evidence for selective pressures. These important gaps will need to be filled in the near future as urbanization and animal population declines continue. Finally, there is an imperative need for better community outreach, involvement, and education to make conservation of all species possible (McKinney 2002). Funding This work was supported by the National Science Foundation [(IOS)-1350070 to S.S.F.]. 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Published by Oxford University Press on behalf of the Society for Integrative and Comparative Biology. All rights reserved. For permissions please email: journals.permissions@oup.com. This article is published and distributed under the terms of the Oxford University Press, Standard Journals Publication Model (https://academic.oup.com/journals/pages/about_us/legal/notices)

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Integrative and Comparative BiologyOxford University Press

Published: Jul 6, 2018

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