Thamnophis Eques Megalops) in North-Central Arizona
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U.S. Department of Energy Report, DOE/ of Energy Report, DOE/NV/11718–477, Las Vegas, Nevada. NV/11432-162, Las Vegas, Nevada. Herpetological Review, 2016, 47(4), 555–561. © 2016 by Society for the Study of Amphibians and Reptiles Prey Availability and Foraging Events of the Northern Mexican Gartersnake (Thamnophis eques megalops) in North-central Arizona The Northern Mexican Gartersnake (Thamnophis eques 2014; Boyarski et al. 2015). Based on these perceived threats, two megalops) is a generalist mesopredator and wetland and riparian of the primary constituent elements (PCEs) deemed essential obligate distributed in Arizona, New Mexico, and northern to the conservation of T. e. megalops in the proposed rule for Mexico (Degenhardt et al. 1996; Rossman et al. 1996; Brennan designation of critical habitat are: 1) a prey base consisting of and Holycross 2006). Extensive population declines have been viable populations of native amphibian and native fish species; documented throughout the range of the species in the United and 2) the absence or occurrence of nonnative aquatic predatory States with potential drivers including the introduction of species at “low enough levels such that recruitment of northern nonnative predatory species, loss of native prey, and habitat Mexican gartersnakes and maintenance of viable native fish or loss. Based on these declines, the U.S. Fish and Wildlife Service soft-rayed, nonnative fish populations (prey) is still occurring” (USFWS) listed T. e. megalops as federally threatened under the (USFWS 2013). Endangered Species Act on July 8, 2014 (USFWS 2014) with viable While an appropriate aquatic prey base is considered a populations occurring in north-central, western, and southern significant factor in T. e. megalops recovery and long-term Arizona. survival, dietary descriptions are limited to relatively few The USFWS identified “harmful nonnative species” including studies and records in Arizona and northern Mexico (García the American Bullfrog (Lithobates catesbeianus), spiny-rayed and Drummond 1988; d’Orgeix et al. 2013; T. Cotten, T. Sprague, predatory fish in the families Centrarchidae and Ictaluridae, pers. comm.). The species is thought to rely primarily on native, and the crayfish Orconectes virilis and Procambarus clarkii as small-bodied prey including annelids, leopard frog (Lithobates the leading threat to the survival of T. e. megalops throughout its sp.) adults and tadpoles, Mexican Spadefoot (Spea multiplicata) range in the United States (USFWS 2014). Confirmed depredation adults, Woodhouse’s Toad (Anaxyrus woodhousii) juveniles events on neonates and adults by L. catesbeianus and Largemouth and tadpoles, and Western Tiger Salamander (Ambystoma Bass (Micropterus salmoides) have been documented in previous mavortium) larvae (USFWS 2014). However, nonnative aquatic studies (Rosen and Schwalbe 1995; Young and Boyarski 2013). In vertebrates have also been identified as potential food sources, addition, potential negative interactions with nonnative species including L. catesbeianus juveniles and tadpoles, soft-rayed fish include competition for native prey species (Rosen and Schwalbe including the Western Mosquitofish (Gambusia affinis) and the 2002; Holycross et al. 2006) and injury or mortality when snakes Red Shiner (Cyprinella lutrensis) (S. Lashway, T. Sprague, pers. prey on centrarchid and ictalurid fish and swallow spines (USFWS comm.), and Goldfish (Carassius auratus), a nonnative spiny- rayed cyprinid fish (García and Drummond 1988). IAIN D. EMMONS* Building on previous dietary accounts, we investigated ERIKA M. NOWAK potential prey availability and confirmed prey use for T. e. KAYLA K. LAUGER megalops, as part of a larger study to determine population Biological Sciences Department and Colorado Plateau demography, habitat use, and spatial behavior for the species Research Station, Northern Arizona University, along the upper-middle Verde River in north-central Arizona Flagstaff, Arizona 86011-5614, USA (Emmons and Nowak 2016). Here, we present additional prey *Corresponding author; e-mail: [email protected] records that provide a greater understanding of T. e. megalops diets, with implications for critical habitat elements deemed Herpetological Review 47(4), 2016 556 ARTICLES essential for the survival of this federally threatened species in megalops and confirmed prey, including gartersnake snout–vent the United States. (SVL) length, mass, and sex, total length (snout to tail tip) for fish Methods.— Study sites were located along the upper-middle and tadpoles, snout–urostyle length (SUL) for metamorphosed Verde River (Yavapai County, Arizona) and featured relatively frogs and toads, and mass for snakes and prey species when wild, undeveloped riparian corridors along the perennial possible. Anurans were identified to age class using the same Verde River, which is listed as one of five remaining United categories as used in trap surveys. We also calculated prey-snake States localities that contain potentially viable populations ratios for length and mass and calculated ratios (mean ± 1 SE) of the species in question (USFWS 2014). One of the sites also for length and mass, when measurements were available for contained artificial lagoons fed by diversions from the main prey items. Telemetered snakes in the process of consuming prey stem river. Due to the vulnerable status of T. e. megalops we items contained surgically implanted internal radio transmitters; have withheld specific site name information to protect snakes surgical procedures and care followed those of Hardy and Greene at the study locations. We pooled data from sites because there (1999), as modified in Nowak (2006) and Boyarski et al. (2015). is potential genetic and reproductive connectivity between Results.—We conducted 22 minnow trap trips at the study subpopulations of gartersnakes (D. Wood, pers. comm.), they sites for an estimated 122,738 trap-hours from May 2012 through share a similar assemblage of potential aquatic prey species, and November 2014 and detected 15,519 total of 12 different potential they are included in the same sub-basin unit as described in the prey species, all aquatic vertebrates, including 10 nonnative designation of critical habitat proposed rule (USFWS 2013). species and two native species (Table 1). Gambusia affinis was We recorded total numbers of potential prey captured in the most frequently encountered nonnative potential prey species self-baiting Gee™ minnow traps (e.g., Nowak and Santana- in minnow traps, comprising 37% (N = 5780) of the total captures Bendix 2002; Holycross et al. 2006). We defined potential prey as with an availability rate of 0.046 ± 0.012 individuals/trap-hour invertebrates and vertebrates identified in previous diet studies (mean ± 1 SE), and Common Carp (Cyprinus carpio) was the least for T. e. megalops and similar aquatic gartersnake species, frequently encountered nonnative potential prey item, comprising including spiny-rayed fish. Furthermore, we categorized all L. < 0.1% (N = 2) of the total captures with an availability rate of 0.001 catesbeianus and A. woodhousii (the two potential anuran species ± 0.001 individuals/trap-hour (mean ± 1 SE). Anaxyrus woodhousii present at the study sites) by age class to maintain consistency tadpoles were the most frequently encountered native potential with the USFWS final listing rule, where juveniles and tadpoles, prey item, comprising 5% (N = 804) of the total captures with an but not subadults or adults, of both species were identified as availability rate of 0.004 ± 0.002 individuals/trap-hour (mean potential prey (USFWS 2014).