Nerodia Fasciata Research Articles

Virulence factors of Salmonella spp. isolated from free-living grass snakes Natrix natrix.

Salmonellosis associated with reptiles is a well-researched topic, particularly in China and the United States, but it occurs less frequently in Europe. The growth of the human population and changes in the environment could potentially increase the interaction between humans and free-living reptiles, which are an unidentified source of Salmonella species. In this study, we sought to explore this issue by comparing the microbiota of free-living European grass snakes, scientifically known as Natrix natrix, with that of captive banded water snakes, or Nerodia fasciata. We were able to isolate 27 strains of Salmonella species from cloacal swabs of 59 N. natrix and 3 strains from 10 N. fasciata. Our findings revealed that free-living snakes can carry strains of Salmonella species that are resistant to normal human serum (NHS). In contrast, all the Salmonella species strains isolated from N. fasciata were sensitive to the action of the NHS, further supporting our findings. We identified two serovars from N. natrix: Salmonella enterica subspecies diarizonae and S. enterica subspecies houtenae. Additionally, we identified three different virulotypes (VT) with invA, sipB, prgH, orgA, tolC, iroN, sitC, sifA, sopB, spiA, cdtB and msgA genes, and β-galactosidase synthesised by 23 serovars. The identification of Salmonella species in terms of their VT is a relatively unknown aspect of their pathology. This can be specific to the serovar and pathovar and could be a result of adaptation to a new host or environment.

Mercury bioaccumulation and Hepatozoon spp. infections in two syntopic watersnakes in South Carolina.

Mercury (Hg) is a ubiquitous environmental contaminant known to bioaccumulate in biota and biomagnify in food webs. Parasites occur in nearly every ecosystem and often interact in complex ways with other stressors that their hosts experience. Hepatozoon spp. are intraerythrocytic parasites common in snakes. The Florida green watersnake (Nerodia floridana) and the banded watersnake (Nerodia fasciata) occur syntopically in certain aquatic habitats in the Southeastern United States. The purpose of this study was to investigate relationships among total mercury (THg) concentrations, body size, species, habitat type and prevalence and parasitemia of Hepatozoon spp. infections in snakes. In the present study, we sampled N. floridana and N. fasciata from former nuclear cooling reservoirs and isolated wetlands of the Savannah River Site in South Carolina. We used snake tail clips to quantify THg and collected blood samples for hemoparasite counts. Our results indicate a significant, positive relationship between THg and snake body size in N. floridana and N. fasciata in both habitats. Average THg was significantly higher for N. fasciata compared to N. floridana in bays (0.22 ± 0.02 and 0.08 ± 0.006 mg/kg, respectively; p < 0.01), but not in reservoirs (0.17 ± 0.02 and 0.17 ± 0.03 mg/kg, respectively; p = 0.29). Sex did not appear to be related to THg concentration or Hepatozoon spp. infections in either species. We found no association between Hg and Hepatozoon spp. prevalence or parasitemia; however, our results suggest that species and habitat type play a role in susceptibility to Hepatozoon spp. infection.

Trap Escape as a Driver of Capture Probability in Semiaquatic Snakes

Sampling biases resulting from capture methodology or animal behavioral responses can significantly skew our understanding of population size and structure. Behavioral responses to passive sampling with traps, such as trap-happy or trap-shy responses, can be accounted for in analyses using individual covariates but they require detailed understanding of the underlying behavioral mechanisms. We used a field experiment to investigate the impact of trap escape on capture rates and determine what traits influence trap escape rates between and within two species of semiaquatic snakes, Liodytes pygaea and Nerodia fasciata. We found that L. pygaea escaped significantly more often than did N. fasciata, smaller L. pygaea escaped at significantly higher rates than did larger conspecifics, and individual capture history significantly influenced escape rates in N. fasciata. Our findings highlight understudied sources of inter- and intraspecific capture heterogeneity in a common sampling technique, and we urge researchers to consider method-specific sampling biases when attempting to produce population parameter estimates for species that are difficult to detect.

The invasive pentastome Raillietiella orientalis in a banded water snake from the pet trade.

Raillietiella orientalis is established in Florida and rapidly spreading both geographically and in known host species. A banded water snake (Nerodia fasciata), purchased in Michigan at a regional reptile show, expectorated a pentastome whose morphology and DNA sequence indicated that it was R. orientalis. This event indicates that, through the pet trade, R. orientalis has been spread 1,500 km from its previously known distribution limit. Fecal sample analyses indicated that the snake was shedding large numbers of embryonated eggs for at least several months. The diversity of reptile species that are both known hosts of R. orientalis in Florida and are commonly sold in the pet trade indicates that this invasive pentastome may become a widespread health concern for pet owners and veterinarians.

Divergent Genital Morphologies and Female-Male Covariation in Watersnakes.

Genital evolution can be driven by diverse selective pressures. Across taxa we see evidence of covariation between males and females, as well as divergent genital morphologies between closely related species. Quantitative analyses of morphological changes in coevolving male and female genitalia have not yet been shown in vertebrates. This study uses 2D and 3D geometric morphometrics to quantitatively compare the complex shapes of vaginal pouches and hemipenes across three species of watersnakes (the sister taxa Nerodia fasciata, N. sipedon, and a close relative N. rhombifer) to address the relationship between genital morphology and divergence time in a system where sexual conflict may have driven sexually antagonistic coevolution of genital traits. Our pairwise comparisons of shape differences across species show that the sister species have male and female genitalia that are significantly different from each other, but more similar to each other than to N. rhombifer. We also determine that the main axes of shape variation are the same for males and females, with changes that relate to deeper bilobation of the vaginal pouch and hemipenes. In males, the protrusion of the region of spines at the base of the hemipene trades off with the degree of bilobation, suggesting amelioration of sexual conflict, perhaps driven by changes in the relative size of the entrance of the vaginal pouch that could have made spines less effective.

Plasma and tissue enzyme activities of banded water snakes (Nerodia fasciata) and diamondback water snakes (Nerodia rhombifer).

To measure plasma and tissue activities of alkaline phosphatase, alanine aminotransferase, aspartate aminotransferase (AST), creatine kinase, and γ-glutamyltransferase in 2 snake species. 6 banded water snakes (Nerodia fasciata) and 6 diamondback water snakes (Nerodia rhombifer). Blood was collected via the ventral tail vein to measure plasma enzyme activities. Animals were then euthanized, and samples of 9 tissues were collected from each snake: skeletal muscle, cardiac muscle, liver, spleen, lung, kidney, testicle, pancreas, and gallbladder. Tissues were frozen for 30 days, then homogenized and processed. Supernatants were collected and analyzed within 24 hours of processing. A linear mixed model was used to determine differences in enzyme activity between tissues and species and assess interactions between tissues and species. Activities of all enzymes were found to differ significantly among tissues. There were also significant differences between species for all enzyme activities, except AST activity. The kidney had the highest alanine aminotransferase and γ-glutamyltransferase activities. Alkaline phosphatase activity was significantly highest in liver and kidney tissues than in other tissue. Creatine kinase activity was highest in skeletal muscle, followed by cardiac muscle and kidney. AST activity was present in all tissues evaluated, but was highest in liver, kidney, and cardiac muscle in both species. Results reinforced the importance of characterizing the origin of tissue enzymes in reptiles to improve our understanding of biochemistry results and highlighted the differences that can exist in tissue enzyme activities between closely related species.

SUCCESSFUL PREDATION OF INVASIVE SOUTH AMERICAN CANE TOADS (RHINELLA MARINA) BY SOUTHERN WATERSNAKES (NERODIA FASCIATA)

South American Cane Toads (Rhinella marina) have been introduced into several regions outside of their native distribution. Outside of their native range, few predators have been documented preying upon R.marina due to their potent toxins secreted in defense. However, prevalence of a toxin resistance gene makes it possible for some snakes of the sub-family Natricinae to consume native toads. We documented successful consumption of the invasive cane toad by the Southern Watersnake (Nerodia fasciata) in southwest Florida, both in the wild and in the laboratory. Southern Watersnakes from populations that existed both with and without cane toads successfully consumed toad prey without obvious ill-effect. Southern Watersnakes in southwest Florida are thus resistant to, and readily consume cane toads, an otherwise relatively predator-free invasive species in Florida. More dietary field data and controlled experiments that measure resistance to multiple prey items, sizes, and frequency will serve to determine the extent to which Southern Watersnakes can impact the size and structure of sympatric populations of cane toads.

Sex-specific growth is mirrored in feeding rate but not moulting frequency in a sexually dimorphic snake.

Sexual size dimorphism (SSD), commonly observed in snakes, may arise from a different growth rate between the sexes. This indicates a sex-specific resource intake that is in fact observable in free-living snakes. It is not so well known whether the sexes can express differential feeding rates under conditions unconstrained by spatial accessibility, competition, etc. Here, I studied sex-specific variation in growth, its correlate-moulting frequency, and feeding rate in a captive group of sexually dimorphic banded water snakes (Nerodia fasciata) with access to food unconstrained by predation, competition or space. I showed that the sexes did indeed differ in relative mass growth in that females grew faster than males (p = 0.02), but such differences were not apparent in the moulting rate (p = 0.19). Such differential growth was mirrored in the sex-specific feeding rate, with females ingesting a larger number of meals than males (p = 0.004). Such variation in feeding rate may be governed by an individual's energy expenditure and can be interpreted as a behavioural tendency that contributes to SSD development, independently of other behavioural characteristics. Sex-specific resource demands may drive the differential effects of increasing resource scarcity on both sexes.

Case Report: Invasive Pentastomes, Raillietiella orientalis (Sambon, 1922), in a Free-Ranging Banded Water Snake (Nerodia fasciata) in North Central Florida, USA

Raillietiella orientalis is an obligate, crustacean parasite that resides in the respiratory tract of definitive snake hosts. Common throughout southeastern Asia and Australia, R. orientalis is believed to have been introduced into southern Florida, United States along with Burmese pythons (Python bivittatus) in the 1990s. While the invasive range of Burmese pythons is restricted to southern Florida, R. orientalis has advanced north in the state in native snake species. R. orientalis were recovered from the lungs, trachea, oral cavity, and esophagus of an emaciated adult female free-ranging banded water snake (Nerodia fasciata) in north central (Alachua County), Florida, USA. Concurrent findings included the recovery of Ochetosoma sp. trematodes from the oral cavity, and multifocal dermal lesions consistent with snake fungal disease (Ophidiomyces ophiodiicola). This is the first report of R. orientalis in north central Florida, well outside the invasive range of the Burmese python, documenting the substantial northward expansion of the known geographical range of this invasive pentastome in Florida.

A Test of Tradeoffs between Performance and Immunity in Water Snakes (Nerodia clarkii X Nerodia fasciata)

All organisms are constrained by limited energy and thus must allocate resources in such a way to maximize fitness. Due to their importance to survival, immunity and performance are each often favored in selective environments and thus may be tradeoffs of each other. To test this tradeoff hypothesis, we compared performance to three constitutive measures of immunity in a wild hybrid population of water snakes (Nerodia clarkii X Nerodia fasciata): agglutination response to sheep red blood cells (SRBC), heterophil: lymphocyte ratio (HLR), and bactericidal ability (BA). Performance was quantified by exercising snakes up to the point of loss of righting response (LRR) and measuring plasma lactate concentration at this point of fatigue. Support for the tradeoff hypothesis was mixed among the immune assays. Bactericidal ability had a significant positive correlation with LRR (P = 0.003, R2 = 0.263); this was counter to our tradeoff hypothesis and suggests that individuals invest in both traits simultaneously. We failed to detect a significant relationship between SRBC and performance. A significant negative relationship was found between HLR and LRR (P = 0.005, R2 = 0.237). Heterophil: lymphocyte ratio is a balance between innate and acquired immunity that is also indicative of baseline stress. This result showed that snakes with higher HLR are constrained and perform poorly compared to snakes with lower HLR, thus indicating a potential role of stress in affecting performance. Alternatively, this result may denote that internal tradeoffs in the immune system between innate and acquired immunity drive tradeoffs between immunity and performance. Snout–vent length (SVL) was significantly positively correlated with our performance measure (P = 0.012, R2 = 0.192). This result may be explained by selection driving increases in performance and/or ontogenetic development of the locomotor apparatus. Similarly, SVL was an important driver of immunity in snakes. Together, these results clearly show an important link exists between multiple physiological traits that are each vital to survival.

Traditional trapping methods outperform eDNA sampling for introduced semi-aquatic snakes.

Given limited resources for managing invasive species, traditional survey methods may not be feasible to implement at a regional scale. Environmental DNA (eDNA) sampling has proven to be an effective method for detecting some invasive species, but comparisons between the detection probability of eDNA and traditional survey methods using modern occupancy modeling methods are rare. We developed a qPCR assay to detect two species of watersnake (Nerodia fasciata and Nerodia sipedon) introduced to California, USA, and we compared the efficacy of eDNA and aquatic trapping. We tested 3–9 water samples each from 30 sites near the known range of N. fasciata, and 61 sites near the known range of N. sipedon. We also deployed aquatic funnel traps at a subset of sites for each species. We detected N. fasciata eDNA in three of nine water samples from just one site, but captured N. fasciata in traps at three of ten sites. We detected N. sipedon eDNA in five of six water samples from one site, which was also the only site of nine at which this species was captured in traps. Traditional trapping surveys had a higher probability of detecting watersnakes than eDNA surveys, and both survey methods had higher detection probability for N. sipedon than N. fasciata. Occupancy models that integrated both trapping and eDNA surveys estimated that 5 sites (95% Credible Interval: 4–10) of 91 were occupied by watersnakes (both species combined), although snakes were only detected at four sites (three for N. fasciata, one for N. sipedon). Our study shows that despite the many successes of eDNA surveys, traditional sampling methods can have higher detection probability for some species. We recommend those tasked with managing species invasions explicitly compare eDNA and traditional survey methods in an occupancy framework to inform their choice of the best method for detecting nascent populations.

Landscape-Scale Effects of Supra-Seasonal Drought on Semi-Aquatic Snake Assemblages

Climate change is predicted to alter the frequency and intensity of precipitation events, placing stress on freshwater aquatic ecosystems and their associated wildlife. Thus, understanding interspecific variation in drought sensitivity and the repeatability of those responses across heterogeneous landscapes is critical. Semi-aquatic snakes serve important roles within aquatic ecosystems and several species are threatened. Yet, little is known about the effects of drought on semi-aquatic snake populations or assemblages. We systematically trapped 20 isolated wetlands in South Carolina before (2006) and after (2013) a multi-year supra-seasonal drought to determine drought-induced shifts in occupancy and detection for five semi-aquatic snake species. Our results confirm that supra-seasonal drought differentially affects semi-aquatic snake species across landscape scales. Specifically, site occupancy decreased dramatically following drought for banded watersnakes (Nerodia fasciata) (0.95 to 0.69) and Florida green watersnakes (Nerodia floridana) (0.32 to 0.05), but was relatively unchanged for black swamp snakes (Seminatrix pygaea), mudsnakes (Farancia abacura) and glossy crayfish snakes (Regina rigida). Species lacking adaptations that make them resistant or resilient to drought may become locally extirpated if climate change projections are realized or landscapes are degraded or fragmented in ways that prevent drought recovery.

Ontogenetic prey size selection in snakes: predator size and functional limitations to handling minimum prey sizes

As body size increases, some predators eliminate small prey from their diet exhibiting an ontogenetic shift toward larger prey. In contrast, some predators show a telescoping pattern of prey size in which both large and small prey are consumed with increasing predator size. To explore a functional explanation for the two feeding patterns, I examined feeding effort as both handling time and number of upper jaw movements during ingestion of fish of consistent size. I used a range of body sizes from two snake species that exhibit ontogenetic shifts in prey size (Nerodia fasciata and N. rhombifer) and a species that exhibits telescoping prey size with increased body size (Thamnophis proximus). For the two Nerodia species, individuals with small or large heads exhibited greater difficulty in feeding effort compared to snakes of intermediate size. However, for T. proximus measures of feeding effort were negatively correlated with head length and snout–vent length (SVL). These data indicate that ontogenetic shifters of prey size develop trophic morphology large enough that feeding effort increases for disproportionately small prey. I also compared changes in body size among the two diet strategies for active foraging snake species using data gleaned from the literature to determine if increased change in body size and thereby feeding morphology is observable in snakes regardless of prey type or foraging habitat. Of the 30 species sampled from literature, snakes that exhibit ontogenetic shifts in prey size have a greater magnitude of change in SVL than species that have telescoping prey size patterns. Based upon the results of the two data sets above, I conclude that ontogenetic shifts away from small prey occur in snakes due, in part, to growth of body size and feeding structures beyond what is efficient for handling small prey.

Ecology and Control of an Introduced Population of Southern Watersnakes (Nerodia fasciata) in Southern California

Abstract: Native to the southeastern United States, Southern Watersnakes (Nerodia fasciata) are known from two sites in California, but their ecological impacts are poorly understood. We investigated the ecology of Southern Watersnakes in Machado Lake, Harbor City, Los Angeles County, California, including an assessment of control opportunities. We captured 306 watersnakes as a result of aquatic trapping and hand captures. We captured snakes of all sizes (162–1063 mm snout–vent length [SVL], 3.5–873.3 g), demonstrating the existence of a well-established population. The smallest reproductive female was 490 mm SVL and females contained 12–46 postovulatory embryos (mean = 21). Small watersnakes largely consumed introduced Western Mosquitofish (Gambusia affinis), while larger snakes specialized on larval and metamorph American Bullfrogs (Lithobates catesbeianus) and Green Sunfish (Lepomis cyanellus). Overall capture per unit effort (CPUE) in traps declined with time during an intensive 76-d trapping bout,...

Herpetofaunal community change in multiple habitats after fifteen years in a southwest Florida preserve, USA.

Herpetofaunal declines have been documented globally, and southern Florida, USA, is an especially vulnerable region because of high impacts from hydrological perturbations and nonindigenous species. To assess the extent of recent change in herpetofauna community composition, we established a baseline inventory during 1995-97 at a managed preserve in a habitat rich area of southwest Florida, and repeated our sampling methods fifteen years later (2010-11). Nine drift fence arrays were placed in four habitat types: mesic flatwood, mesic hammock, depression marsh, and wet prairie. Trapping occurred daily for one week during 7-8 sampling runs in each period (57 and 49 total sampling days, respectively). Species richness was maintained in mesic hammock habitats but varied in the others. Catch rates of several native species (Anaxyrus terrestris, Lithobates grylio, Anolis carolinensis, Nerodia fasciata) declined significantly. Other native species (Lithobates sphenocephalus, Siren lacertian, and Notophthalmus viridescens piaropicola) that were abundant in 1995-97 declined by greater than 50%. Catch rate of only two species (the nonindigenous Anolis sagrei and the native Diadophis punctatus) increased significantly. Hierarchical cluster analysis indicated similarity within habitat types but significant dissimilarity between sampling periods, confirming shifts in community composition. Analysis of individual species’ contributions to overall similarity across habitats shows a shift from dominance of native species in the 1990s to increased importance of nonindigenous species in 2010-11. Although natural population fluctuations may have influenced differences between the two sampling periods, our results suggest considerable recent change in the structure and composition of this southwest Florida herpetofaunal community. The causes are unknown, but hydrological shifts and ecological impacts of nonindigenous species may have contributed.

Projecting invasion risk of non-native watersnakes (Nerodia fasciata and Nerodia sipedon) in the western United States.

Species distribution models (SDMs) are increasingly used to project the potential distribution of introduced species outside their native range. Such studies rarely explicitly evaluate potential conflicts with native species should the range of introduced species expand. Two snake species native to eastern North America, Nerodia fasciata and Nerodia sipedon, have been introduced to California where they represent a new stressor to declining native amphibians, fish, and reptiles. To project the potential distributions of these non-native watersnakes in western North America, we built ensemble SDMs using MaxEnt, Boosted Regression Trees, and Random Forests and habitat and climatic variables. We then compared the overlap between the projected distribution of invasive watersnakes and the distributions of imperiled native amphibians, fish, and reptiles that can serve as prey or competitors for the invaders, to estimate the risk to native species posed by non-native watersnakes. Large areas of western North America were projected to be climatically suitable for both species of Nerodia according to our ensemble SDMs, including much of central California. The potential distributions of both N. fasciata and N. sipedon overlap extensively with the federally threatened Giant Gartersnake, Thamnophis gigas, which inhabits a similar ecological niche. N. fasciata also poses risk to the federally threatened California Tiger Salamander, Ambystoma californiense, whereas N. sipedon poses risk to some amphibians of conservation concern, including the Foothill Yellow-legged Frog, Rana boylii. We conclude that non-native watersnakes in California can likely inhabit ranges of several native species of conservation concern that are expected to suffer as prey or competing species for these invaders. Action should be taken now to eradicate or control these invasions before detrimental impacts on native species are widespread. Our methods can be applied broadly to quantify the risk posed by incipient invasions to native biodiversity.

Intraspecific and interspecific variation of female genitalia in two species of watersnake

The morphological differences in female genitalia within and between species are little studied and poorly understood, yet understanding patterns of variation in female genitalia can provide insights into mechanisms of genital evolution. The present study aimed to explore the patterns of intraspecific and interspecific variation in female genitalia in two sister taxa of watersnake (Nerodia sipedon and Nerodia fasciata) that have similar genital shape. We used a geometric morphometric (GM) approach to study variation in shape of the vagina between and within two sister species. We examined genital shape in female watersnakes ranging from small, sexually immature females to large reproductive females that had recently given birth. We found that shape variation of genitalia is strongly correlated with body size, where larger but not smaller females have a bifurcation in the vagina. However, we also found significant shape variation in the structure of the vagina between the two species, where N. fasciata has narrower genitalia with more prominent bifurcation, whereas N.sipedon has wider genitalia with less marked bifurcation. Using GM allowed us to detect significant differences in genital shape that were not apparent upon visual examination alone, suggesting that shape variation in female genitalia may be greater than previously assumed. Additional study of morphological differences in male reproductive organs for these species would help to determine whether there has been genital co-evolution, and potentially mechanical reproductive isolation, in these two closely-related and occasionally sympatric species. © 2013 The Linnean Society of London.

Feeding in natricines: relationships among feeding morphology, behavior, performance and preferred prey type

AbstractThe diversity of feeding mechanisms among predators reflects phenotypic modifications that may improve feeding performance on a preferred prey type. I compared trophic morphology, feeding performance (time and upper‐jaw walks) and behavior (initial bite and ingestion directions) among three species of natricine snakes that were fed fish and frogs over a broad range of relative prey sizes. Feeding behavior was influenced by prey type but did not differ among the snake species. Both bite and ingestion directions influenced the number of upper‐jaw movements (lateral excursions and protractions) required to consume fish, but only initial bite position significantly affected the number of upper‐jaw movements required to ingest frogs. Within snake species, feeding performance did not differ between fish and frogs for Nerodia fasciata and Nerodia rhombifer; however, Thamnophis proximus consumed fish with fewer upper‐jaw movements. Feeding time differed significantly among snake species when fed both fish and frogs. Trophic morphology did not significantly affect ingestion costs for fish but did influence ingestion when fed frogs. In general, differences in trophic morphology among the three species are not correlated to handling and ingestion performance.

Allometry of cranial morphology, gape size and ingestion performance in the banded watersnake (Nerodia fasciata) feeding on two types of prey

Small body size imposes limitations on the feeding capabilities of juveniles, particularly in species that consume their prey whole. It has been hypothesized that juveniles exhibit exceptional performance measures to compensate for their small size. However, few studies have examined whether juveniles have better feeding performance relative to adults and investigations of snake feeding ontogeny have not shown enhanced performance in smaller snakes. I tested the hypothesis that juvenile snakes have better feeding performance by comparing maximum gape circumference and ingestion performance (time and number of pterygoid protractions) in a series of banded watersnakes (Nerodia fasciata) of different sizes fed fish and frogs. I also measured several external and osteological dimensions of the head and used Akaike's information criterion to determine which morphological measurements were the strongest predictors of relative gape. All skull measurements and maximum gape circumference showed negative allometry relative to snout-vent length (SVL). Given the available models, Akaike information criterion (AIC) analysis indicated that both skull length and mandible length were the strongest predictors of gape circumference for both external and osteological measurements. Multiple regression analysis of ingestion performance indicated SVL was negatively correlated with the time and number of pterygoid protractions required to consume fish or frogs, indicating that juveniles do not have a higher ingestion performance than adults. While exaggerated morphology in juvenile snakes does not appear to improve ingestion performance, a larger gape should increase the ability of juvenile snakes to consume a wide range of encountered prey shapes and sizes.

The Function of Oscillatory Tongue-Flicks in Snakes: Insights from Kinematics of Tongue-Flicking in the Banded Water Snake (Nerodia fasciata)

Tongue-flicking is an important sensory behavior unique to squamate reptiles in which chemical stimuli gathered by the tongue are delivered the vomeronasal organ situated in the roof of the mouth. Because tongue-flick numbers can easily be quantified, this behavior has been widely used as a measure of vomeronasal sampling in snakes using related variables such as tongue-flick rate or tongue-flick/attack score. Surprisingly, the behavior itself and especially the function of the oscillatory tongue-flicks remains poorly understood. To describe the overall kinematics of tongue-flicking in the colubrid snake Nerodia fasciata and to test predictions on the function of oscillatory tongue-flicks, we filmed the tongue-flicks of 8 adult Nerodia fasciata using 4 synchronized high-speed cameras. Three-dimensional kinematic and performance variables were extracted from the videos in order to quantify tongue movements. Based on the kinematic analysis, we demonstrate the existence of 2 functional and behavioral tongue-flick categories. Tongue-flicks with oscillations meet all the criteria for being adapted to the collection of odorants; simple downward extensions appear better suited for the rapid pick up of nonvolatile chemical stimuli from the substrate or a food item. External stimuli such as tactile and/or vomeronasal stimulation can induce a shift between these categories.