Conservation Assessment

R9 Conservation Assessment for the Green , Aneides aeneus (Cope and Packard)

TITLE PAGE – Green Salamander, Aneides aeneus in the Eastern Region of the U.S. Forest District 9 (populations in , Ohio, , Maryland and West ).

Robert Brodman Professor & Chair of Biology & Environmental Science Saint Joseph’s College, IN 47978

Date of Completion: January 6, 2004.

DISCLAIMER – This Conservation Assessment was prepared to compile the published and unpublished information on the subject taxon or community; or this document was prepared by another organization and provides information to serve as a Conservation Assessment for the Eastern Region of the Forest Service. It does not represent a management decision by the U.S. Forest Service. Though the best scientific information available was used and subject experts were consulted in preparation of this document, it is expected that new information will arise. In the spirit of continuous learning and adaptive management, if you have information that will assist in conserving the subject taxon, please contact the Eastern Region of the Forest Service Threatened and Program at 310 Wisconsin Avenue, Suite 580 Milwaukee, Wisconsin 53203.

1. EXECUTIVE SUMMARY: The Green salamander, Aneides aeneus is designated as a Regional Forester Sensitive Species on the Hoosier and Monongahela National Forests in the Eastern Region of the Forest Service. The species occurrence is documented but not designated as sensitive on the Hoosier and Monongahela National Forests. The purpose of this document is to provide the background information necessary to prepare Conservation Approaches and a Conservation Strategy that will include management actions to conserve the species.

The green salamander is more tolerant to dry conditions than other species of and is a good climber. Its includes crevices in rock outcrops and cliffs in forested landscapes. They need moist rock crevices to reproduce and hibernate but they are also arboreal, using trees and bark as substrates for foraging on insects.

The green salamander has a spotty distribution with several populations that are disjunct from the main distribution in the Allegheny Plateau. Much of the range includes isolated and peripheral populations that are vulnerable to statewide extinction in most of the states within it range. The range is likely to be more restrictive than currently recognized because the green-salamander-complex consists of several related species. Within the National Forest District 9 few populations have been documented from the Hoosier and Monongahela National Forests.

The main threat to green salamander populations is deforestation resulting in loss, degradation or fragmentation of habitat. Logging activities should be managed to keep at least 100m of buffered forest habitat around rock outcrops and cliffs.

Little is known about the population biology, lifespan, mortality rates, dispersal, colonization of , metapopulation dynamics, the extent of arboreal activity, and the phylogeography of significant evolutionary-units throughout the range. More species-specific surveys need to be conducted, especially within the National and State Forests within the Eastern Region of the National Forest Service.

2. ACKNOWLEDGEMENTS: I thank Jennifer Wykle for providing locality information, showing me green salamander habitats in the Monongahela National Forest and for comments on an earlier version of this manuscript. I thank Anne Timm for providing locality information and showing me green salamander habitats in the Hoosier National Forest. I thank Andrew Johnson, Jeffery Corser, and Jayme Waldron for providing me with unpublished data, thoughtful insights, and for comments on an earlier version of this manuscript. I thank Jim Petranka for comments on management recommendations.

3. NOMENCLATURE AND : The green salamander Aneides aeneus (Cope and Packard 1881) is a member of the tribe Plethodontini within the lungless salamander family , order Caudata. Phyogenetic studies have suggested that there is more than one species currently assigned to Aneides aeneus (Morescalchi 1975; Sessions and Kezer 1987). Jeff Corser’s group has found that nucleotide sequences, as well as ecological, and morphological traits, show that there are at least four species-level clades of green salamanders. Their mtDNA clades generally follow the karyological pattern detected by Sessions and Kezer (1987) and Dick Highton has unpublished allozyme data which also confirm to the same pattern (Corser, personal communication). The populations in Ohio, Pennsylvania, Maryland and will be in members of a newly named northern Cumberland species, however specimen from the disjunct Indiana populations have not been included in any phylogenetic analysis (Corser, personal communication).

4. DESCRIPTION OF SPECIES: The green salamander (Fig. 1) is a long-legged, slender salamander with bright green to yellow lichen-like patches on a dark background making this species easy to identify (Bishop 1943; Minton 2001). Its flattened body and head and its expanded, square-tipped toes are distinctive adaptations for climbing and living in rock crevices (Wake 1966; Juterbock 1989; Petranka 1998). The eyes are large and protuberant (Bishop 1943). The ventral surface is light bluish or yellowish gray and unmarked except for faint yellow coloration at base of each leg (Jutterbock 1989; Petranka 1998; Minton 2001). Adults measure 8-12 cm TL and have 14-15 costal grooves (Bishop 1943; Petranka 1998; Minton 2001). There is no sexual dimorphism of color or size (Juterbock 1989). Hatchlings look like small adults but have relatively shorter tails (Walker and Goodpaster 1941; Gordon 1952).

Figure 1. Green salamander from West Virginia (from Johnson (2002) and taken by Mizuki Takahashi).

The green salamander is the only salamander in eastern North America that has bright green coloration making this the key characteristic for identification.

5. LIFE HISTORY: 5.1. Reproduction: Reproduction is terrestrial with direct development that lacks a larval stage (Pauley and Watson 2004). Mating usually occurs in May and June (Brooks 1948, Gordon 1952; Snyder 1971; Cupp 1971; Cupp 1991; Canterbury and Pauley 1994) with some observations of breeding activity in autumn (Snyder 1971; Cupp 1971; Cupp 1991; Canterbury and Pauley 1994). Females produce eggs on a biennial cycle (Snyder 1991; Canterbury and Pauley 1994). Eggs are laid in clusters of 10-32 attached to the roof of a moist rock crevice where females brood the eggs for 2-3 months (Gordon 1952; Lee and Norden 1973; Green and Pauley 1987; Juttebock, 1989; Cupp 1991; Canterbury and Pauley 1994). Females that were experimentally removed and displaced up to 9 m away returned to their eggs within 24 hours (Gordon 1961) and females may use the same breeding crevice in successive years (Cupp 1991). Nest failure is common when females abandon clutches (Gordon 1952; Snyder 1973). Eggs hatch in September and October (Walker and Goodpaster 1941; Gordon 1952; Woods 1969; Juterbock 1989; Cupp 1991; Canterbury and Pauley 1994). Females also attend newly hatched young for 3-5 weeks after hatching (Walker and Goodpaster 1941; Gordon 1952; Norden and Groves 1974; Cupp 1991). Size at sexual maturity is 4.5 cm SVL (Juterbock, 1989) in the spring-summer of their third year (Canterbury 1991). An on-going study by Jeff Humphries and Jayme Waldron suggests that WV and Blue Ridge Escarpment populations have similar life history strategies.

5.2. Ecology: Little is known about the longevity of green salamanders. They probably live longer than 3 years since sexual maturity is not reached until the third year (Canterbury 1991). Corser (2001) reports that Castanet (unpublished data) suggests that they may live 10 or more years. Little is know about the mortality rate of green salamanders.

Little is known about the diet of green salamanders but they are likely to be invertebrate specialists (Corser, personal communication). Specimen from West Virginia have been found with snails, slugs, arachnids, and small insects in their stomachs (Gordon 1952; Rubin 1969; Lee and Norden 1973; Canterbury and Pauley 1990).

Predators include ring-neck snakes (Diadophis punctatus) and common garter snakes (Thamnophis sirtalis) and possible adult Allegheny Mountain dusky salamanders (Pauley and Watson 2004). Defenses include moving deep into crevices (Petranka 1998) , undulating the tail from side-to-side (Brodie 1977), and remaining immobile when disturbed (Brodie et al. 1974). No diseases are documented in green salamanders but the parasitic nematode Batacholandros magnavulgaris has been reported from green salamanders (Baker 1987).

Home range size has not been determined for green salamanders, however there is some data on linear movements. Juveniles move more than adults and most movements are short distances (< 4 m) within a single rock outcrop as animals leave crevices to forage (Canterbury 1991, Cupp 2000). Green salamanders are sometimes found on dry rocks but tend to move into deeper crevices during hot and dry stretches (Pauley 1993). However there are larger movements (16-107 m) between rocky outcrops during dispersal prior to hibernation in late autumn (Gordon 1952; Canterbury 1991; Waldron 2000). At least 12% of the marked in these studies moved distances ranging from 46-107 m site the point of original capture. Green salamanders are sometimes found crossing roads (Williams and Gordon 1961; Cupp 1991) and are often found 30-50 m and up to 400 m from rocky outcrops (Waldron personal communication). A molecular study within a metapopulation in West Virginia suggests that forested habitats between rock outcrops do not act as barriers to gene flow (Johnson, 2002).

Male green salamanders are territorial and aggressively defend their rock crevices against other males (Cupp 1980; Canterbury and Pauley 1991). Resident males won 95.6% of encounters with conspecific males (Cupp 1980), however laboratory experiments show that male green salamanders cannot successfully defend crevices from northern slimy salamanders and salamanders (Canterbury and Pauley 1991).

5.3. Dispersal/Migration: There are pre-hibernation and post-hibernation migrations of green salamanders (Gordon 1952; Cupp 1991; Canterbury 1991). Hibernation occurs in deep interconnecting crevices (Gordon 1952) and may involve large groups of green salamanders sharing winter hibernacula (Waldron, personal communication).

5.4. Obligate Associations: Because of there microhabitat preferences, there are no obligate associations of green salamanders with other species of (Petranka 1998). Green salamanders, Alleghany Mountain dusky salamanders (Desmognathus ochrophaeus), and northern slimy salamanders ( glutinosus) stratify vertically with the green salamanders selecting crevices that are higher on the rock faces (Baltar 1983; Cliburn and Porter 1986, 1987; Canterbury 1991; Waldron 2000). This is expected to reduce competition (Petranka 1998; Pauley and Watson 2004). Laboratory studies suggest that green salamanders compete with Cumberland Plateau salamanders (P. kentucki) and Wehrle’s salamanders (P. wehrlei) (Cupp 1980; Canterbury and Pauley 1994). In Indiana cave salamanders (Eurycea lucifuga) were the most common found at sites inhabited by green salamanders and may be competing for limited crevice space (Madej 1998).

6. HABITAT: 6.1. Range-wide: Green salamanders mostly inhabit forested rock outcrops and cliffs (Gordon 1952; Juterbock 1989; Madej 1998). They prefer deep moist crevices that do not have running or dripping water or moss (Gordon and Smith 1949). Rocky areas inhabited by green salamanders are usually well- shaded within mixed mesophytic forests (Gordan 1952; Corser 1991; Petranka 1998). The type of rock is less important than crevice size and moisture (Gordon and Smith 1949). The green salamanders tolerate drier condition than any other species of salamander (Gordon 1952). Green salamanders are occasionally found in caves (Cope and Packard 1881; Juterbock 1998) and are rarely found under rocks or under ground cover (Minton 2001).

The green salamander was considered to be weakly arboreal (Bishop 1928). Many early observations (1920s-1950s) of green salamanders report them under bark in dead trees and fallen logs or on the bark of living trees (Brimley 1927; Bishop 1928; Pope 1928; Welter and Barbour 1940; Brimley 1941; Fowler 1947; Barbour 1949; Gordon 1952; Schwartz 1954) when substantial amounts of old- growth forests covered much of the range of the green salamander (Petranka 1998). Most of these observations were from the exfoliating bark of American (Castenea dentata). Pope (1928) also reported observing a brooding female green salamander within a small cavity of an oak (Quercus) tree. However, there have been a number of recent observations suggesting that green salamanders are more arboreal than previously thought (Minton 2001; Wilson 2003; Waldron personal communication). Green salamanders have been observed on a variety of tree species and as high up as 20 m (Waldron, personal communication) and include additional observations of breeding activity in trees (Wilson 2003; Waldron personal communication). Loss of old-growth forests and arboreal foraging sites due to the chestnut blight and logging may have primarily restricted the green salamanders to crevices of rock outcrops in the last half century (Gordon 1952; Corser 1991; Petranka 1998; Corser 2001, Wilson 2003).

6.2. National Forests: In the Monongahela National Forest there were 3 reports at 2 sites in the 1930’s and 6 reports at 6 sites in the 1990’s of green salamanders (National Heritage Database). On October 20, 2003 I visited four of these sites with Jennifer Wykle and found a total of 3 green salamanders at two sites (Fig. 2). Most of the habitat that I saw consisted of isolated rocky outcrops surrounded by mature mixed mesophytic forests. There appears to be ample appropriate habitat for green salamanders at Monongahela, however the sparse records appears to be an artifact created by a lack of survey work. Most green salamander research in West Virginia has been conducted by researchers from Marshall University in areas south and west of the Monongahela National Forest.

Figure 2. Green salamander habitat in the Monongahela National Forest, WV. The arrow in the bottom panel points to the face of a green salamander within a rock crevice. In the Hoosier National Forest there have been two populations of green salamanders reported from Crawford County (Madej 1994; Madej 1998; Minton 2001). The first population discovered in 1993 was the first occurrence of this species in Indiana. On October 25, 2003 I visited the two known sites with Anne Timm (Fig.3). We did not observe any green salamanders but there appears to be a substantial amount of appropriate habitat at each site. Most of the habitats that I saw were rocky cliffs and outcrops within mature mixed mesophytic forests.

Figure 3. Green salamander habitat in the Hoosier National Forest, IN.

7. DISTRIBUTION AND ABUNDANCE: 7.1. Range-wide Distribution: The green salamander is distributed throughout the Allegheny Plateau from southwest Pennsylvania, Maryland and Virginia to northern and (Dunn 1926; Bishop 1943; Ferguson 1961b; Richmond 1952; Harris and Lyons 1968; Bier 1985; Mitchell and Reay 1999) at elevations < 1340 m (Petranka 1998). There are also disjunct populations in extreme southern Indiana (Madej 1994), Blue-Ridge Mountain populations in southwestern , northwestern and northeastern (Petranka 1998), the Great Smoky Mountains in (Weller 1931), central Tennessee (Redmond and Scott 1996), and extreme southern Ohio (Gordon 1967). Several parts of the geographic range probably represent distinct species with restricted ranges (Corser, personal communication). There has been a recent discovery of a green salamander population across the river from Indiana in Breckinridge County by John MacGregor (Corser, personal communication).

The Great Smoky Mountain population has not been located since its discovery in the 1930s (Redmond and Scott 1996). The spotty distribution in Ohio could have resulted from range retraction caused by lumbering and land clearing (Gordon 1952). Green salamanders have declined in North Carolina during the 1970s (Mitchell et al. 1999). Snyder (1991) attributed this decline to prolonged cold weather. Populations in the Blue-Ridge Escarpment had a 98% decline in abundance since the 1970s (Snyder 1983; Corser 2001). Corser (2001) suggested that this decline was range-wide within the Blue- Ridge Escarpment and that this was due to habitat loss, over-collecting, epidemic disease and climate change. Green salamanders had very large population densities in Kentucky during the 1930s but declined afterward (Barbour 1971, Snyder 1991). Snyder (1991) suggests that creating openings in the canopy, as caused by forest thinning for construction, could have caused salamanders to leave the suddenly unshaded rocky habitats.

Population densities range from 0.1to 0.25 individuals per square meter and populations are often less than 50 individuals (Gordon 1952; Juterbock 1998; Madej 1998). The unique habitat requirements and deforestation likely contribute to the patchy distribution and uncommon abundance of green salamanders (Petranka 1998; Madej 1998; Wilson 2003). A closely related species, the clouded salamander (Aneides ferreus), shares similar microhabitat preferences (Petranka 1998) and declines after clear-cut logging (Bury and Corn, 1988). A 100 m buffer of forested habitat surrounding rock outcrops is suggested for survival and dispersal between rock outcrops (Petranka 1998; Pauley and Watson 2004).

7.2. Region-wide Distribution (including state and National Forest specific information): Within the Eastern Region of the U.S. Forest District the green salamander has been found in Crawford County Indiana, within the Hoosier National Forest, Adams and Lawrence Counties in extreme southern Ohio, Fayette County in southern Pennsylvania, Garrett County in extreme western Maryland, and throughout the central counties of the Alleghany Plateau in West Virginia including the Monogahela National Forest (Fig 4, from USGS ARMI). While more than 50 populations of cave salamanders were found in recent surveys of cave fauna in the Blue River area, and other areas in southern Indiana, no green salamanders were found (Ewert at al, 1993; Lewis 1998; Brodman 2003).

Figure 4. The historic range of the green salamander (adapted from USGS ARMI).

8. RANGE WIDE STATUS: The TNC global rank of the green salamander is G3G4 and the national rank is N3N4. The state heritage program rankings of the green salamander are summarized in table 1.

Table 1. State Heritage program rankings of green salamanders (adapted from Madej 1998; Pauley and Watson 2004). State listed species are listed in bold (SR = State Rare and protected , SC = State Special Concern. ST = State Threatened, SE = State Endangered). State Ranking and rational for ranking Alabama S3 – rare or uncommon Georgia S2 – imperiled because of rarity, SR Indiana S1 – critically imperiled because of extreme rarity, SE Kentucky S4 – apparently secure Maryland S2 – imperiled because of rarity, SE Mississippi S1- critically imperiled because of extreme rarity, SE North Carolina S2 – imperiled because of rarity, SE Ohio S2 – imperiled because of rarity, SE Pennsylvania S1 – critically imperiled because of extreme rarity, ST South Carolina S1 – critically imperiled because of extreme rarity, SC Tennessee S4 – possibly in peril but status uncertain Virginia S3 – rare or uncommon West Virginia S3 – rare or uncommon, SC

The risk factors for green salamanders occurring in the Monongahela National Forest are as follows. The abundance risk factor is B3 the habitat integrity risk factor is B2, and the population vulnerability risk factor is A2 based on limited populations on MNF (Evans 2000).

The risk factors for green salamanders occurring in the Hoosier National Forest are as follows. The abundance risk factor is A3 based on just three observations along the Ohio River from 1993-1998. The distribution risk factor is B2 and the population trend risk factor is B2 because populations are at the edge of the range. The habitat integrity risk factor is B2 because there is no specific protection of damp rock crevices and outcrops on HNF, but activities usually avoid such habitat. The population vulnerability risk factor is B2 because they are somewhat resistant to dry conditions (Reynolds 1999).

9. POPULATION BIOLOGY AND VIABILITY: Little is known about the population biology or population viability of green salamanders (Juterbock 1989; Petranka 1998).

10. POTENTIAL THREATS: 10.1. Present or Threatened Risks to Habitat: Habitat loss, degradation and fragmentation are the greatest threats to green salamander populations. Logging near rock outcrops is the greatest risk to green salamander habitat. Preservation of hibernacula and forested corridors are also high priorities.

10.2. Over utilization: Over collecting green salamanders is listed as a probable cause of declines in some populations (Corser 2001). The bright green color and tolerance to dry conditions could make green salamanders targets of the pet trade.

10.3. Disease or Predation: Little is known about disease or predation on green salamanders.

10.4. Inadequacy of Existing Regulatory Mechanisms: There are currently no management plans to keep buffered of forested habitat around rock outcrops.

10.5. Other Natural or Human Factors: Pollution and climate change could be factors.

11. SUMMARY OF LAND OWNERSHIP AND EXISTING HABITAT PROTECTION: All currently known green salamander populations in Indiana are currently on land owned by the National Forest Service in the Hoosier National Forest. In West Virginia about half of the green salamander habitat is owned by the state (primarily State Forests) and the rest is private or federally owned within the Monongahela National Forest and the New River National River (Wykle, personal communication). These habitats are protected as long as management allows for at least a 100 m buffer around rock outcrops that remain forested and forested corridors between populations (Petranka 1998; Pauley and Watson 2004). The buffer provides core arboreal habitat (Waldron, personal communication) and protection against edge effects and desiccation of rock crevices (Petranka, personal communication).

12. SUMMARY OF EXISTING MANAGEMENT ACTIVITIES: Existing management activities that most influence the conservation of green salamanders are the preservation of habitat and the threat of logging near rock outcrops.

13. PAST AND CURRENT CONSERVATION ACTIVITIES: The green salamander is protected in states that have listed the species as endangered or threatened (Indiana, Ohio, Maryland, Pennsylvania, North Carolina, Georgia, Mississippi). The National Forest Service purchased land in Indiana to protect the first population of green salamanders that was discovered in the state in 1993.

14. RESEARCH AND MONITORING 14.1. Existing Surveys, Monitoring, and Research: Much of the current ecological research on the green salamander in district 9 is being conducted by Thomas Pauley and his students at Marshall University. Survey sites are rangewide in West Virginia but with no specific emphasis on the Monongahela National Forest. No current surveys or population monitoring is occurring in the Hoosier and Monongahela National Forests. Other current ecological studies in the south are being conducted by Jeff Humpheries, Jayme Waldron and students at Clemson University and phylogeographic studies are being conducted by Jeff Corser at the Tennessee Department of Environment and Conservation and Joseph Collins at University of Kansas.

14.2. Survey Protocol: Because of the unusual habitat preferences of green salamanders, they can be missed by community-level survey methods and protocols. Fauna-level surveys in and near the Hoosier National Forest did not detect the species (Ewert 1993; Lewis 1998; Brodman 2003). Crevices in rock outcrops and cliffs need to be systematically searched. Foggy summer evenings are the best times to search (Waldron personal communication).

14.3. Research Priorities: Little is known about the population biology or the extent to which individual green salamanders disperse from isolated rock outcrops (Juterbock 1989; Petranka 1998). Studies need to be conducted to determine lifespan, mortality rates, and dispersal. More studies are needed on the metapopulation dynamics of green salamanders including dispersal and colonization of habitat and factors that contribute to sink and source populations. The arboreal nature of green salamanders needs to be further investigated to determine what portion of time is spent in trees vs. rock crevices. These data are necessary to better determine the amount of habitat and size of buffer zone that is needed to conserve populations.

An unpublished 13 year (Corser, personal communication) study from several populations in North Carolina shows that green salamanders exist in metapopulations and movement between habitable rock outcrops is key to their population dynamics and viability. The extinction and colonization dynamics observed by Corser suggests that buffers around rock outcrops and forested corridors are essential for their conservation.

Phylogeographic studies suggest that the green salamander-complex comprises of several species (Morescalchi 1975; Sessions and Kezer 1987; Corser personal communication). However the newly discovered disjunct populations in Indiana have not been studied to determine if they are a unique species or evolutionary-unit. More species-specific surveys are needed, especially in the Hoosier and Monongahela National Forests to determine the abundance and distribution of green salamanders and the quantity and quality of their habitat. These studies are needed to better determine the current range distribution, vulnerability to extinction and population viability.

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16. APPENDIX * none

17. LIST OF CONTACTS Jeff Corser, Tennessee Department of Environment and Conservation Andrew Johnson, Marshall University, WV James Petranka, University of North Carolina, Ashville Anne Timm, Hoosier National Forest, IN Jayme Waldron, Clemson University, SC Jennifer Wykle, West Virginia Division of Natural Resources