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Climbing Behavior of Flatwoods Salamanders (Ambystoma bishopi /A. cingulatum)
Kelly C. Jones, Pierson Hill, Thomas A. Gorman, and Carola A. Haas

Southeastern Naturalist, Volume 11, Issue 3 (2012): 537–542

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Climbing Behavior of Flatwoods Salamanders (Ambystoma bishopi /A. cingulatum) Kelly C. Jones1,*, Pierson Hill2, Thomas A. Gorman1, and Carola A. Haas1 Abstract - As a result of declining populations and reduced availability of suitable habitat, Ambystoma bishopi (Reticulated Flatwoods Salamander) and A. cingulatum (Frosted Flatwoods Salamander) are federally listed as endangered and threatened, respectively, by the United States Fish and Wildlife Service under the Endangered Species Act. Recovery efforts are hindered by a lack of basic natural history information. Although the fossorial life history of ambystomatid salamanders often precludes direct observation of adults and juveniles, an incidental encounter of A. bishopi climbing on Aristida stricta (Wiregrass) sparked a new search effort. On 13 occasions from 29 April 2010 to 23 November 2011, we examined herbaceous ground cover in and around six known breeding wetlands in the Florida panhandle, documenting 36 observations of flatwoods salamanders (10 adults and 26 juveniles), 30 of which were climbing up to 0.5 m above the ground in Wiregrass. These observations led us to hypothesize that Wiregrass, a fire-dependent grass species associated with flatwoods salamander breeding habitat, may be used by juvenile and adult flatwoods salamanders as a foraging substrate (foliage) and/or as refugia (foliage and root base). Although these observations are interesting, further research is needed to determine if the salamanders are selectively using wiregrass and if a foraging and/or refugial advantage is gained by climbing. Among ambystomatid salamanders, the federally endangered Ambystoma bishopi Goin (Reticulated Flatwoods Salamander) and federally threatened A. cingulatum Cope (Frosted Flatwoods Salamander) are among the most poorly understood and least studied. Opportunities to observe behavior have been limited due to widespread extirpations over the last several decades (Palis 1997a) coupled with dramatic declines in local populations (Means et al. 1996, Palis et al. 2006, USFWS 2009). Furthermore, the two species are regarded as secretive and difficult to observe. Consequently, little is known about their activity and post-metamorphic habitat use. Considering that conservation efforts for flatwoods salamanders are currently underway, it is imperative that we increase our understanding of these species’ natural history so monitoring strategies and management decisions can be tailored accordingly. Herein we report on novel behaviors of flatwoods salamanders observed in the field in the Florida panhandle during spring and fall of 2010 and 2011. We observed that juvenile and adult A. bishopi and A. cingulatum were using Aristida stricta Michx. (Wiregrass), a fire-dependent grass species associated with suitable flatwoods salamander breeding habitat, possibly as a foraging substrate (foliage) and/or as refugia (foliage and root base). Field-site description and methods. On 13 occasions from 29 April 2010 to 23 November 2011, we searched for flatwoods salamanders at night by visually examining herbaceous ground cover in ecotones and basins of four known breeding wetlands on Eglin Air Force Base (AFB), one known breeding site in St. Marks National Wildlife Refuge (NWR), and one known breeding site in Apalachicola National Forest. We recorded dates and times of all searches; when a salamander was encountered, we recorded 1Department of Fish and Wildlife Conservation, Virginia Tech, Blacksburg, VA 24061. 2Department of Biological Sciences, Florida State University, Tallahassee, fl32306. *Corresponding author - kcjones7@vt.edu. Notes of the Southeastern Nat u ral ist, Issue 11/3, 2012 537 538 Southeastern Naturalist Vol. 11, No. 3 substrate (plant species or genus), salamander age (as determined by relative size), and presence (recent metamorph) or absence of residual larval stripes (J. Palis, , pers. comm.). We determined the sex of adults by visually inspecting for the presence (male) or absence (female) of cloacal swelling (Palis 1997b). In addition, we recorded any noteworthy behaviors for all observations. Our methods were designed to monitor climbing behavior as initially observed and were not intended to imply a greater frequency of use than other potential substrates (e.g., walking on the ground or using fossorial retreats). Three sites on Eglin AFB were dome swamps with Wiregrass-dominated ecotones, surrounded by mature Pinus palustris Mill. (Longleaf Pine) mesic flatwoods with a moderately dense, shrubby midstory. The fourth site on Eglin AFB was a highly disturbed dome swamp with little remaining ecotone, surrounded by Longleaf Pine mesic flatwoods beyond a paved road along 40% of its boundary and Pinus clausa [Chapm.] Vasey (Sand Pine) scrub habitat surrounding the remainder of the site. The site at St. Marks NWR was a Cladium sp. P. Br. (Sawgrass)-dominated ephemeral wetland with little herbaceous ecotone, surrounded by Pinus elliottii Englm. (Slash Pine) flatwoods with a dense, shrubby midstory. The site at Apalachicola National Forest was a dome swamp with a Wiregrass-dominated ecotone at the head of a cypress strand, surrounded by Longleaf Pine mesic flatwoods. Observations. On 13 occasions from 29 April 2010 to 23 November 2011, we observed 32 A. bishopi and 4 A. cingulatum near breeding sites in both spring and fall, including 10 observations (8 bishopi and 2 cingulatum) of adults and 26 observations (24 bishopi and 2 cingulatum) of juveniles (Table 1). For 30 of the 36 observations of salamanders climbing vegetation, salamanders (22 juvenile and 6 adult A. bishopi, including 1 gravid female, and 2 juvenile A. cingulatum) were found up to 0.5 m above the ground in Wiregrass (Fig. 1). No individual salamanders were moving when initially observed, but several began climbing, both vertically and horizontally, among the blades of Wiregrass during observation. All four legs were engaged during locomotion through Wiregrass, making contact with vertical Wiregrass blades as well as horizontal pine needles and dead Wiregrass blades from prior years’ growth. Salamanders always appeared to use their tails for support while crossing larger horizontal gaps, and often curled their tails while vertically descending in a controlled fashion. Individuals that descended rapidly seemed to move Table 1. Summary of behavioral observations of Ambystoma bishopi and A. cingulatum. Start/end times encompass all observations but also include some time not spent searching. Start/end # of # of Date Location times observers salamander observations 29 April 2010 Eglin AFB 2045-2200 3 10 juvenile bishopi 30 April 2010 Eglin AFB 0135–0205 2 1 juvenile bishopi 5 May 2010 Eglin AFB 2025–2400 1 10 juvenile bishopi 12 May 2010 St. Marks NWR 2130–2300 2 1 juvenile cingulatum 13 May 2010 Eglin AFB 2130–2330 3 2 juvenile bishopi 22 May 2010 Eglin AFB 2300–2400 1 1 juvenile and 1 adult bishopi 3 Nov 2010 Apalachicola NF - 1 2 adult cingulatum 29 Nov 2010 Eglin AFB 2200–2210 1 1 adult bishopi Spring 2011 Apalachicola NF - 1 1 juvenile cingulatum 15 Nov 2011 Eglin AFB 2245–2315 3 2 adult bishopi 16 Nov 2011 Eglin AFB 1915–1950 2 2 adult bishopi 16 Nov 2011 Eglin AFB 2130–2215 1 1 adult bishopi 23 Nov 2011 Eglin AFB 0315–0415 2 1 adult bishopi 2012 Southeastern Naturalist Notes 539 in a way that would be best described as wriggling. Several individuals of both species (juvenile and adult) remained in virtually the same location for up to 1 hr after the initial observation. No individuals were monitored for more than 1 hr. On 6 occasions, individuals were observed climbing vegetation other than Wiregrass; 3 were found perched in vegetation, and 3 were observed actively climbing in response to being disturbed by observers. The perching salamanders included 1 juvenile A. bishopi observed on top of a Taxodium ascendens Brongn. (Pond Cypress) knee, another juvenile climbing on Dichanthelium sp. [Hitchc. & Chase] Gould (Rosette Grass), and 1 adult male A. bishopi perched in a relatively dense patch of Aristida palustris [Chapm.] Vasey (Longleaf Threeawn). On 3 November 2010, during the fall breeding season, several adult male Ambystoma cingulatum were observed crawling amidst patches of herbs within a dry pond basin. When alerted to the observer’s presence, 2 individuals quickly retreated up into the nearest complex elevated structure provided by blades of Carex sp. (sedges). The salamanders remained motionless 10–15 cm above the ground for approximately 10 min before descending and resuming crawling. Similarly, on 23 November 2011, during the next year’s fall breeding season, 2 adult male and 1 adult female A. bishopi were all observed actively moving about within a dry pond basin in the vicinity (less than 2 m) of several oviposition sites from the previous year. After being accidentally disturbed by the observer, the female quickly retreated up the nearest complex elevated structure provided by Xyris sp. (Yellow-Eyed Grass) and Aristida palustris. She remained motionless 15–20 cm above the ground for approximately 6 min before climbing back down and disappearing from view in the vegetation. On this same night, simultaneous drift-fence monitoring revealed movement of adult flatwoods salamanders into 2 breeding sites on Eglin AFB, indicating that breeding activity was likely occurring. On 9 occasions, the Wiregrass supporting the salamanders was accidentally disturbed by the observer; when this occurred, juveniles and adults retreated down through the Figure 1. Juvenile Ambystoma bishopi perched high in Wiregrass (left) and close-up of same individual (right). 540 Southeastern Naturalist Vol. 11, No. 3 Wiregrass and out of sight. One juvenile that had disappeared out of sight returned to within 10 cm of its original location high in the Wiregrass within 30 minutes of being disturbed. One juvenile was observed descending over the edge of a recently burned Wiregrass hummock, after which it entered a small hole (<1 cm diameter) at the base of the clump. One individual retreated straight over the periphery of a grass clump and was observed falling into water below. On 2 additional occasions, at least 1 juvenile Ambystoma cingulatum and 1 adult A. bishopi did not demonstrate a retreat response when the Wiregrass supporting them was lightly disturbed. Discussion. Although an association between flatwoods salamander larval occurrence and herbaceous vegetation structure has been documented (Gorman et al. 2009, Palis 1995), and Wiregrass (when inundated) has been identified as an important component of larval habitat (Sekerak et al. 1996), these new observations suggest that herbaceous vegetation may also play an important role in terrestrial components of juvenile and adult flatwoods salamander natural history. We hypothesize that graminaceous vegetation, particularly Wiregrass, may be used by juvenile and adult flatwoods salamanders as a foraging substrate (foliage) and/or as refugia (foliage and root base). The apparently novel behavior of climbing on vegetation in a primarily fossorial animal is important because it broadens our knowledge of the biphasic life-history requirements of these species and points to a broader connection, beyond the larval phase, to intact understory vegetation. Although climbing behavior has been observed by 3 captive adult A. cingulatum which climbed up to 25 cm above the substrate in a planted bunch of wiregrass at night on multiple occasions (J. Palis, pers. comm.), these observations represent the first time this behavior has been observed in the field. It has been proposed that plethodontid salamanders often climb on plants on wet nights to gain a foraging advantage (Jaeger 1978). Although there does not appear to be any record of direct observation of ambystomatid salamanders climbing for this reason, Judd (1957) found insects that feed or rest on plants above ground, including larvae of Tortricidae and adults of Curculionidae, Miridae, and Chironomidae, in the stomachs of A. jeffersonianum Green (Jefferson Salamanders), and Smith et al. (2004) found adults of Curculionidae in the stomachs of newly metamorphosed A. mavortium Baird (Barred Tiger Salamanders). Furthermore, over the course of the study we did observe several potential prey items on clumps of Wiregrass. These prey items included spiders, small roaches (order Blattodea), early instars of orthopterans, and lepidopteran adults and larvae, including members of the moth family Geometridae. In addition to potentially providing foraging opportunities, Wiregrass may also be important for reducing risk of exposure to terrestrial predators, as it appears to do for plethodontid salamanders (Roberts and Liebgold 2008). However, little is known about the predators of flatwoods salamanders. Because all of our observations occurred in the vicinity of breeding sites where breeding adults and pre-dispersal juveniles are likely to be concentrated, we may be at an advantage for observing this behavior. For example, the relatively large number of individuals observed on 29–30 April and 5 May 2010 (Table 1) was likely due to an abundance of newly metamorphosed juveniles. The precipitous drop in number of individuals found following those two nights may be due to their dispersal into the surrounding uplands. Observing climbing activity away from breeding sites would be interesting but probably very difficult without the aid of radio transmitters or in the context of a fairly natural captive setting outside of the breeding season. The fact that both juveniles and adults engaged in climbing behavior appears to indicate that this is a normal part of flatwoods salamander post-metamorphic life history and underscores the importance of intact habitat conditions for all life stages of these two 2012 Southeastern Naturalist Notes 541 rare and declining species. Intact herbaceous ground cover, especially in and near breeding sites, has been shown to be an important component of breeding habitat (Gorman et al. 2009, Sekerak et al. 1996) and will be essential for the continued existence of these species. Efforts to maintain herbaceous ground cover, such as application of prescribed burning, at these breeding sites may be most productive during the growing season, because it coincides with the historic timing of wildfires. In addition, the growing season is the time period when these ephemeral wetlands are more often dry and conducive to burning (see Frost 1995). Growing season fires are more likely to increase seeding of wiregrass and other herbaceous plant species (Brewer and Platt 1994, Outcalt 1994). However, in many locations within the southeastern US, dormant season fire is still the most widely used method, because conditions are better for controlling fires (Bishop and Haas 2005). Our observations have provided a valuable new method for observing flatwoods salamanders and may offer an additional method for confirming occupancy at breeding sites. Additional research is needed to determine if the salamanders are selectively using wiregrass and if they are gaining an advantage for foraging and/or predator avoidance by climbing. Acknowledgments. We thank the Natural Resources Branch (Jackson Guard) of Eglin Air Force Base and the Department of Fish and Wildlife Conservation at Virginia Tech for their financial and logistical support. We thank B. Rincon, S. Goodman, D. Allison, W. McDearman, K. Wray, M. Olmstead, and J. Bente for their field assistance, J. Palis for sharing his observations with us, S. Jones for her editing assistance, and D. Steen and 2 anonymous reviewers for their helpful comments. Literature Cited Bishop, D.C., and C.A. Haas. 2005. Burning trends and potential negative effects of suppressing wetland fires on flatwoods salamanders. Natural Areas Journal 25:290–294. Brewer, J.S., and W.J. Platt. 1994. Effects of fire season and herbivory on reproductive success in a clonal forb, Pityopsis graminifolia. 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