Monitoring Amphibians in Great Smoky Mountains National Park by C

Monitoring Amphibians in Great Smoky Mountains National Park By C. Kenneth Dodd, Jr.

— Abstract — a national amphibian monitoring program on Federal lands, to develop the sampling techniques and bio- Amphibian species have inexplicably declined metrical analyses necessary to determine status and or disappeared in many regions of the world, and in trends, and to identify possible causes of amphibian some instances, serious malformations have been declines and malformations. observed. In the United States, amphibian declines The biological importance of Great Smoky frequently have occurred even in protected areas. Mountains National Park has been recognized by its Causes for the declines and malformations probably designation as an International Biosphere Reserve. are varied and may not even be related. The seem- As such, it is clearly the leading region of signifi- ingly sudden declines in widely separated areas, how- cance for amphibian research. Although no other ever, suggests a need to monitor amphibian region shares the wealth of amphibians as found in populations as well as identify the causes when the Great Smokies (31 species of salamanders, and declines or malformations are discovered. 13 of frogs), the entire southern and mid-section of In 2000, the President of the United States and the Appalachian Mountain chain is characterized by Congress directed Department of the Interior (DOI) a high diversity of amphibians, and inventories and agencies to develop a plan to monitor the trends in monitoring protocols developed in the Smokies likely amphibian populations on DOI lands and to conduct will be applicable to other Appalachian National Park research into possible causes of declines. The DOI Service properties. has stewardship responsibilities over vast land hold- From 1998 to 2001, USGS biologists carried ings in the United States, much of it occupied by, or out a pilot inventory and monitoring research project potential habitat for, amphibians. The U.S. Geologi- in Great Smoky Mountains National Park. A variety cal Survey (USGS) was given lead responsibility for of inventory, sampling, and monitoring techniques planning and organizing this program, named the were employed and tested. These included wide-scale Amphibian Research and Monitoring Initiative visual encounter surveys of amphibians at terrestrial (ARMI). Authorization carried the mandate to set up and aquatic sites, intensive monitoring of selected

1 plots, randomly placed small-grid plot sam- critique of what the results tell the observer, and pling, litterbag sampling in streams, monitoring a discussion of the limitations of the techniques nesting females of selected species, call sur- and the data. The report reviews considerations veys, and monitoring specialized habitats, such for site selection, outlines steps for biosecurity as caves. Coupled with information derived and for processing diseased or dying animals, from amphibian surveys on Federal lands using and provides resource managers with a decision various other techniques (automated frog call tree on how to monitor the Park’s amphibians data loggers, PVC pipes, drift fences, terrestrial based on different levels of available resources. and aquatic traps), an amphibian monitoring It concludes with an extensive list of references program was designed to best meet the needs of for inventorying and monitoring amphibians. biologists and natural resource managers after USGS and Great Smoky Mountains National taking into consideration the logistics, terrain, and life histories of the species found within Park biologists need to establish cooperative Great Smoky Mountains National Park. efforts and training to ensure that congression- This report provides an overview of the ally mandated amphibian surveys are per- Park’s amphibians, the factors affecting their formed in a statistically rigorous and distribution, a review of important areas of biologically meaningful manner, and that biodiversity, and a summary of amphibian life amphibian populations on Federal lands are history in the Southern Appalachians. In addi- monitored to ensure their long-term survival. tion, survey techniques are described as well as The research detailed in this report will aid examples of how the techniques are set up, a these cooperative efforts.

Introduction

The Florida Caribbean Science Center constraints. Survey teams sampled more than (now Florida Integrated Science Center) 500 sites (fig. 2) and recorded data on more than received funding in 1997 from the U.S. Geolog- 10,000 amphibians. All parts of the Park were ical Survey (USGS) Inventory and Monitoring visited in all seasons and in all weather (I&M) Program to conduct a pilot project for conditions. amphibians in the southeastern United States. The objectives of the Great Smoky After considering several locations, Great Mountains National Park I&M program were Smoky Mountains National Park (fig. 1) was to: (1) provide a geographically referenced selected for the survey because of its amphibian inventory of the amphibian resources of the diversity and the large number of potential Great Smoky Mountains National Park; threats to its varied ecosystems (Brown, 2000). (2) provide indices of abundance of Park During the course of the next 4 years, a field selected amphibian species, referenced to loca- research team of enthusiastic young biologists tions and habitat types; (3) develop and transfer was assembled to collect information on the to the Great Smoky Mountains National Park species richness and distribution of the Park=s and National Park Service a series of protocols amphibians. Researchers used a variety of sam- suitable for long-term monitoring of amphibian pling techniques, including 10 x 10-meter sur- populations in the “Smokies” and other Appala- vey plots, “permanent” 30 x 40-meter plots, chian parks; (4) evaluate current distributions coverboards, litter-bag surveys, and a great and abundance of amphibian species as possible number of time-constrained litter and stream in the Park with literature reports of past inves- searches. The team looked for previously tigations. This manual fulfills the third objective reported rare species, sampled historic loca- of the I&M program. Additional information on tions, investigated unique habitats (such as amphibian natural history, distribution, land- caves), and examined museum records and pub- scape ecology, trends analysis, and protocol lished literature. Survey activities and tech- development are published in Dodd and others, niques were designed to optimize the use of (2001), Waldron and others, (2003); Dodd, available personnel within budget and logistic (2004), or is under development.

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Figure 2. Location of U.S. Geological Survey sampling sites in Great Smoky Mountains National Park, 1998 to 2001.

Acknowledgments

This manual is the result of collaborative survey National Park, particularly Keith Langdon, Dana efforts during 4 years of hard field work between 1998 Soehn, and Jack Piepenbring. and 2001 in the Great Smoky Mountains. The author For providing funding: Norita Chaney (USGS would like to thank the following individuals for their Inventory and Monitoring Program, Reston, support, assistance, dedication, and companionship: Virginia), Russell Hall (Florida Integrated Science For their field collecting skill, expertise, and gen- Center), and the USGS Amphibian Research and eral good humor during all kinds of weather condi- Monitoring Initiative (ARMI). tions, the USGS field crew from 1998 to 2001 (in For providing research permits: National Park alphabetical order): Jamie Barichivich, Jeff Corser, Service (Great Smoky Mountains National Park), Elizabeth Domingue, Marian Griffey, Kelly Irwin, North Carolina Wildlife Resources Commission, Christy Morgan, Rick >Bubba= Owen, Kevin G. Tennessee Wildlife Resources Agency. Smith, and Jayme Waldron. For figure 7: Todd For providing the beautiful illustrations of larval Campbell. For advice: James Petranka and Chuck amphibians: Jacqualine Grant. Smith. For reading the manuscript and making valuable For their support and assistance: the National Park comments: Jamie Barichivich, Marian Griffey, Service personnel at Great Smoky Mountains Russell Hall, Steve Johnson, Keith Langdon, and Jennifer Staiger.

4 HOW TO USE THIS GUIDE

Monitoring Amphibians in Great Smoky programs are not considered in detail. However, Mountains National Park is meant to help references are provided at the end of this guide National Park Service natural resource biolo- (see References on Inventorying and Moni- gists, university researchers, nongovernmental toring Amphibians). biologists, and the interested public understand Future amphibian monitoring within and overcome some of the biological and nonbi- Great Smoky Mountains National Park will be ological constraints to setting up a large-scale linked to the U.S. Department of Interior (DOI) inventory and monitoring program for amphibi- Amphibian Research and Monitoring Initiative ans inhabiting the Great Smoky Mountains. (ARMI). Standardized methods of data collec- Some of the information applies only to tion, entry, and analysis currently are being amphibians within the Great Smokies, whereas developed by ARMI researchers for all DOI information on setting up inventory and moni- lands. Pertinent information will be made avail- toring programs may have more broad applica- able to Federal agencies and ARMI partners tions with regard to Appalachian amphibians. through ARMI=s web site: Many persons who use this guide will be famil- edc2.usgs.gov/armi/ iar with basic amphibian biology, but others will require a refresher course or will be unfamiliar A cautionary note: There is always the with amphibian life histories. danger that site information will be misused This guide serves as a companion volume by criminal elements to find amphibians in to Dodd (2004) and, for that reason, information order to collect them. This is true in National in that work has not been duplicated except Parks and on other Federal lands, as well as on when absolutely necessary. There is usually an private lands. None of the amphibians found exception to every generalization discussed in Great Smoky Mountains National Park are below, and biologists should expect to encoun- endangered or threatened under the Federal ter species outside of their “normal” habitat, Endangered Species Act of 1973, as amended, that often do not fit identification information, although several species, such as Hellbenders, or that have unusual behavioral patterns. Exten- are protected by state law. Locations of many sive information is not included on threats to of the Park=s amphibians, including its amphibians (for example, habitat loss and alter- endemic salamanders, are well known via the ation, disease, nonindigenous species, climate published scientific literature and on records change, toxic chemicals, UVB, malformations) attached to museum specimens. Therefore, it or the various reasons why amphibians are vul- seems unlikely that mentioning Park locations nerable to environmental problems (including in this guide will increase the probability of their biphasic life cycle, skin permeability, and collection, especially when these species are the complex morphological and biochemical found readily, and often in greater abundance, transformations which accompany metamor- outside the Park. For example, the Mole Sala- phosis). These topics are dealt with in more mander, Southern Zigzag Salamander, and detail elsewhere (Dodd, 1997, 2004; Alford and Mud Salamander might be considered Arare@ Richards, 1999; Corn, 2000; Houlahan and or Aisolated@ within Great Smoky Mountains others, 2000). National Park, yet very large and widespread All of the potential sampling protocols, populations of these species are found in the techniques, and methods of data analysis that Tennessee Valley and elsewhere. Still, Park may accompany, or be required for, a large- Service employees and research scientists scale amphibian inventory and monitoring program cannot be discussed within one short working within the Park, including field sur- guide. For this reason, many specialized tech- vey teams, must be observant for illegal col- niques are not discussed, instructions are not lectors and immediately report suspicious provided for making traps, and statistical activities to law enforcement personnel.

5 AMPHIBIANS OF THE GREAT SMOKY MOUNTAINS

Species Richness A total of 31 salamanders and 13 frogs have been recorded from the Great Smoky Mountains National Park. Note that common names are capitalized, and that species names (consisting of a genus and specific epithet) are italicized. Species codes allow data to be entered in shorthand for- mat. To minimize data entry errors, species codes should be either all capitalized or all in lower case letters. Capitals and lower-case letters should not be intermixed. Using accepted and standardized common and scientific names (Crother, 2000), the amphibians are: Suggested species Common name Scientific name code Salamanders Spotted Salamander Ambystoma maculatum AMA Marbled Salamander Ambystoma opacum AOP Mole Salamander Ambystoma talpoideum ATA Green Salamander Aneides aeneus AAE Hellbender Cryptobranchus alleganiensis CAL Seepage Salamander Desmognathus aeneus DAE Spotted Dusky Salamander Desmognathus conanti DCO Imitator Salamander Desmognathus imitator DIM Shovel-nosed Salamander Desmognathus marmoratus DMA Seal Salamander Desmognathus monticola DMO Ocoee Salamander Desmognathus ocoee DOC Black-bellied Salamander Desmognathus quadramaculatus DQU Santeetlah Salamander Desmognathus santeetlah DSA Pigmy Salamander Desmognathus wrighti DWR Three-lined Salamander Eurycea guttolineata EGU Junaluska Salamander Eurycea junaluska EJU Long-tailed Salamander Eurycea longicauda ELO Cave Salamander Eurycea lucifuga ELU Blue Ridge Two-lined Salamander Eurycea wilderae EWI Spring Salamander Gyrinophilus porphyriticus GPO Four-toed Salamander Hemidactylium scutatum HSC Common Mudpuppy Necturus maculosus NMA Eastern Red-spotted Newt Notophthalmus viridescens NVI Northern Slimy Salamander Plethodon glutinosus PGL Jordan=s Salamander Plethodon jordani PJO Southern Gray-cheeked Salamander Plethodon metcalfi PME Southern Appalachian Salamander Plethodon oconaluftee POC Southern Red-backed Salamander Plethodon serratus PSE Southern Zigzag Salamander Plethodon ventralis PVE Mud Salamander Pseudotriton montanus PMO Black-chinned Red Salamander Pseudotriton ruber PRU Frogs Northern Cricket Frog Acris crepitans ACR American Toad Bufo americanus BAM Fowler=s Toad Bufo fowleri BFO Eastern Narrow-mouthed Toad Gastrophryne carolinensis GCA Cope=s Gray Treefrog Hyla chrysoscelis HCH Spring Peeper Pseudacris crucifer PCR Upland Chorus Frog Pseudacris feriarum PFE American Bullfrog Rana catesbeiana RCA Northern Green Frog Rana clamitans RCL Pickerel Frog Rana palustris RPA Northern Leopard Frog Rana pipiens RPI Wood Frog Rana sylvatica RSL Eastern Spadefoot Scaphiopus holbrooki SHO

6 Amphibian taxonomy and systematics within the southern Appalachians are topics of intense debate among biologists. Rationale for using the listed names is provided by Dodd Five major forest communities are recog- (2004). nized within the Great Smoky Mountains National Park.... Habitats and Distribution

Five major forest communities are recognized within the Great Smoky Mountains National Park, although 80 percent of the Park The spruce-fir forest (fig. 3) is domi- falls within the Eastern Deciduous Forest Eco- nated by Red Spruce (Picea rubens) and Fraser system (Houk, 1993). Some botanists have fur- Fir (Abies fraseri), and is found generally above ther subdivided the vegetation into as many as 1,676 m (5,500 ft), although the community 67 florally distinct communities. No one spe- descends to 1,372 m (4,500 ft) in some locations cies of amphibian is associated entirely with a and individual Red Spruce are found at even single forest community, although some of the lower elevations. This is the Canadian Zone high-elevation salamanders (Plethodon jordani, boreal forest of high moisture, cool or cold tem- Desmognathus ocoee, D. wrighti) are more often found in the spruce-fir community than in peratures, and high humidity (Houk, 1993). other community types. Habitat structure, par- Ground surface is often dense with fallen tree ticularly one that retains moisture and high branches and trunks, and carpeted by thick lay- humidity, is important in shaping salamander ers of tree needles. Wet, rotten, woody debris distribution. The high-elevation coniferous for- and dense needle mats provide ideal hiding est appears ideal in providing shade, cover (in places for terrestrial salamanders. Streams orig- the form of coarse woody debris), and abundant inate in this habitat, usually beginning as small surfaces for moisture condensation. seeps and springs. As streams trickle through

Figure 3. Spruce-fir forest at Indian Gap.

7 the dark-green forest, they gather momentum. below 1,372 m (4,500 ft) in sheltered valleys, Even at higher elevations, aquatic salamanders, and is dominated by Tulip Poplar (Liriodendron particularly duskies (Desmognathus) and Blue tulipifera), Dogwood (Cornus florida), Red Ridge Two-lined Salamanders (Eurycea Maple (Acer rubrum), Sweetgum (Liquidamber wilderae), may be plentiful within the head- styraciflua), White Basswood (Tilia americana water streams. var. heterophylla), Yellow Buckeye (Aesculus At somewhat lower elevations (1,067- flava), and Black Birch (Betula lenta). Both 1,524 m; 3,500-5,000 ft), deciduous northern hardwood communities have complex under- hardwoods (fig. 4) predominate, such as Sugar story vegetation, often with much coarse woody Maples (Acer saccharum), American Beech debris, which provides cover for terrestrial sala- (Fagus grandifolia), and Yellow Birch (Betula manders. The streams through these hardwood alleghaniensis). Many terrestrial and aquatic forests are rocky and fast paced, and sala- salamanders reach their lower or upper distribu- manders are very common along streamsides tional range within this community; frogs are and in the water. scarce. Cove hardwoods, the third community, Two somewhat specialized forest com- comprise the most diverse forest community in munities are found in the Smokies. The hemlock the Smokies, one that is endemic to the southern community (fig. 5) is dominated by Eastern Appalachian Mountains. It occurs generally Hemlocks (Tsuga canadensis), commonly

Figure 4. Deciduous forest at Lynn Hollow.

8 Hemlocks are massive with tall, straight trunks. When they fall, they provide excellent habitat for salamanders....

Figure 5. Hemlock forest at Chinquapin Knob.

called “spruce-pines” by natives of the southern straight trunks. When they fall, they provide mountains, and is common between 1,067- excellent habitat for salamanders, both in the 1,524 m (3,500-5,000 ft) in elevation. rotting wood and under exfoliating bark (fig. 6). Hemlocks descend to much lower elevations The pine-oak forest (fig. 7) occupies the along cold mountain stream valleys, and over- drier areas of the Park, particularly the area west lap considerably with both hardwood forests of Cades Cove and at mid-elevations on the and the spruce-fir forest of the higher North Carolina side of the Park. This forest is elevations. Hemlocks are massive with tall, dominated by Southern Red (Quercus falcata), 9 Figure 6. Coarse woody debris in Cove forest at Roaring Fork. Note the pink survey flags marking the position of transects.

Northern Red (Q. rubra), Scarlet (Q. coccinea), Cove. As a result, amphibian species richness is Black (Q. velutina), and Chestnut (Q. prinus) surprisingly high, particularly for frogs. Oaks, and by Pitch (Pinus rigida), White (P. Amphibians are not uniformly distributed strobus), and Table Mountain (P. pungens) throughout the Great Smoky Mountains Pines. Soils are dry, as is the leaf litter. Prior to National Park. There are wide-ranging species, human intervention, this community burned fre- species restricted to specialized habitats, and quently in the western regions of the Park, and species found in only one area of the Park. a fire-adapted vegetation community resulted. Terrestrial salamanders are few, and usually found only during cool, wet times of the year. Aquatic-breeding salamanders and frogs are found along streamsides, where they likely remain close to water. The bottomlands along Cane Creek and Abrams Creek likely formed a corridor from the Tennessee Valley into Cades

10 Monitoring programs will need to take the but not common species; and wide ranging in distribution of species into account to optimize higher or lower elevations, and generally com- time and financial resources. A few generaliza- mon. Because they do not have larvae, they tions can be made about amphibian distribution must be sampled where they carry out their and habitats within the Park. entire life cycle, usually on the forest floor and under leaf litter and other debris. SALAMANDERS Terrestrial salamanders (see Life Amphibians are not History) include species that are: restricted in distribution in the Great Smokies; wide ranging uniformly distributed throughout the Great Smoky Mountains National Park.

Figure 7. Oak-pine forest. Photographer: Todd Campbell, University of Tampa

11 Notes Breeding occurs in occurs Breeding bouts following 2-3 lar-rain; pond type vae larvae; Pond type ventrolat- series of eral light spots a line forming limb inser- below tions larvae; Pond type life histo- variable to ries with regard events timing of In older larvae, spotsblotches or may fuse with slender, more than DQ legs longer snouts round than all much larger lots Desmogs; other of yolk 1-2 mos after hatching Tail attributes Tail tail fin lightly tail fin mottled or stippled; finely tip dark at fin dorsal almost extends to front limbs and yellow black on tail fin spot pattern continues on tail spatulate Belly patternBelly white or light throat stippled; scattered melan- on lat- ophores eral sides on band dark in midline (poor specimens) some Dorsal pattern dull olive green, no dull olive conspicuous mark- ings blackish, drab; older mottling have larvae body on black and yellow blotches along midline of back dorsolat- with sides dorsum stripe; eral variable conspicuous dark, sides on flecks light light brown an] Spots on dorsum 5-8 pairs of even or even pairs of 5-8 or spots alternating blotches light spots 2 rows dor- pairs light 4-5 between sal spots limbs pairs alter- of 4-6 nating light spots on dorsum spots light pairs 6-8 between limbs or even pairs of 4-5 or spots alternating blotches pproximately; <, less than; >, greater th Metamorphosis Size and time of Size ic salamandersMountains of Great National Smoky Park 29-32 mm SVL; 43-60 43-60 SVL; mm 29-32 TL mm 75 (to TL mm mid- if overwinter); June to August 49-58 SVL; mm ~33 mm TL; late March mid-June mm SVL; 32 to 50 May to Sept. to 20 mm SVL, 9-12 SVL; July to earlymm fall? mm SVL; May 26-38 to Oct. June-July mm SVL; May 11-15 to June SVL; mm to 54 35-42, mid-summer to 20 mm SVL, 9-12 SVL; July to earlymm fall? size Hatching 12-17 mm TL 12-17 mm TL 10-14 ~10 mm SVL SVL; mm 8-12 mm TL 12-20 SVL mm 11 mm SVL 11-12 mm TL 13-18 mm SVL 11-16 SVL; mm 8-12 mm TL 12-20 Larval period 2-4 mos 2-4 mos 5-7 but mos, 3-4 variable <1 yr (10-20 3 yrs mos) mos 10-11 mos 9-10 yrs 3-4 <1 yr total length; length; ~, a SVL, snout-vent Hatching April-May winter winter to early spring to earlyJuly fall to mid-Aug mid-Sept. early summer to fall; Sept. Aug to late Sept. to Sept. July to earlyJuly fall Egg deposition Jan. to late Mar. Jan. to late Mar. (mountains-late Feb. 4-7 to early Mar.); weeks incubation (inOct-Nov pond by Sept.); 9-15days must incubation, but days 1-2 flooded be (winter) Sept. to Mar. early May to early perhaps to mid- July, days 45-60 August; incubation late spring to early summer? late spring to early summer?; 10-12 weeks incubation mid-June to mid- incu- 2 mos August; bation to earlyJuly Aug. to days 52-74 Sept.; incubation May to June early May to early perhaps to mid- July, days 45-60 August; incubation Identification and life history of the nonpermanently aquat Species Table 1. year; mm, millimeter; TL, months; yr, [mos, Ambystoma mac- Ambystoma ulatum opacum A. talpoideum A. Desmognathus conanti D. imitator D. marmoratus D. monticola D. ocoee D. quadramaculatus D. santeetlah

12 Notes stream type dense, well-defined cheek patches; of margin lower dark pigmentation straight stream type; tail fin near stops insertion of rear limbs; red- dish gills; square snouts truncated long smallsnouts with eyes larvae; pond type occurs; joint nesting brooding stream type; over- occurs; wintering can be very larvae large type; no stream black chinsor dorsal spots Tail attributes Tail dorsal does fin for- not extend ofward rear legs Belly pattern immaculate no iridophores immaculate light with iridophores immaculate dull white Dorsal pattern cream; uniformly stippled; then dark dorsolateral broad stripe; mid- narrow dorsal stripe green to deep olive brown cream colored; then uniformly dark,similar to adults; no spots paired sparse pigmentation with 3 longitudinal the spots on series of side dusky to brown light yellow fleck- withgray fine ing yellow nondescript, fin dorsal brown; to head extends older light brown; with widely scattered spots weakly light brown; mottled or streaked an] Spots on dorsum no paired light spots 6-9 pairs light dorsolateral proximately; <, less than; >, greater th Metamorphosis Size and time of Size salamanders of Great Smoky Mountains National Park (Continued) 22-27 mm to mm SVL, 22-27 32 mm SVL; June to August 34-42mm SVL; mid- to August May mm SVL; 23-28 > 50over- mm TL if mid-June- wintering; July to SVL; mm 31-37 70 spring mm TL; in SVL mm 18-19 1to 32 mm SVL in yr, April to July 2 yr; to mm SVL, 55-65 70 mm high elevation; late June to August 11-15mm SVL; 17- 25 mm TL; July? 35-44mm SVL; mid- May to Sept. 34-46mm SVL; 62- 86 mm TL; May to July size Hatching 11-12.5 mm mm 11-12.5 SVL 7-9SVL; mm TL mm 11-13 mm 18-21 mm SVL; 40 TL; 9-12 mm SVL; to 17.5 mm TL 7-9SVL; mm TL mm 11-14 TL mm 18-22 mm SVL <13 TL mm 11-14 Larval period 3.5-5.5 mos (< 1 yr), but may overwinter 1-2 yr normally <1 yr (4-7 mos) 6-18 mos; most 12- 15 mos 1-2 yr to 4 yr to 61 21 days mos 15-17 29-30 to mos 1.5 to 3.5yr (27-31 mos) tal length; SVL, snout-vent length; ~, ap length; SVL, snout-vent tal length; ory of the nonpermanentlyory of aquatic Hatching early to mid- Mar? June? early Nov-March after 4-12 weeks to May August late summer to autumn May-June? winter mid-Dec to mid-Feb Egg deposition winter at mid-May least by late autumn to early spring Sept. to Feb. to May; 4-10 Feb. weeks incubation summer Feb to May autumn to early winter autumn to early winter; 3 mos incubation Identificationand life hist Species Table 1. [mos, months; yr, year; mm, millimeter; TL, to [mos, months; yr, Eurycea Eurycea guttolineata junaluska E. longicauda E. E. lucifuga E. wilderae Gyrinophilus porphyriticus Hemidactylium scutatum Pseudotriton montanus P. r u b e r

13 Monitoring programs can target each type of Species that are common and wide distributional pattern or habitat listed below, ranging at higher elevations depending upon the objectives of the researchers and the funds and personnel available. For Pigmy Salamander (Desmognathus wrighti); example, whereas a few people can easily mon- Jordan=s Salamander (Plethodon jordani); itor the status of the Southern Zigzag Sala- Southern Gray-cheeked Salamander (Pleth- mander, a much more elaborate protocol will be odon metcalfi). necessary to monitor populations of the South- Species that are common and wide ern Red-backed Salamander. A number of these ranging at lower elevations species are syntopic, making multispecies monitoring a realistic objective. As much as possi- Northern Slimy Salamander (Plethodon glu- ble, single species sampling and monitoring tinosus); Southern Red-backed Salamander should be avoided in favor of multispecies (Plethodon serratus). sampling and data recording. Some examples of River-dwelling salamanders inhabit only typical distribution patterns follow: the largest of the Smokies’ rivers (fig. 8), Species restricted in distribution including Little River, Middle Prong, Ocon- aluftee River, Little Pigeon River, Abrams Southern Zigzag Salamander (Plethodon Creek, the lower reaches of Deep Creek and, ventralis). perhaps, Hazel Creek. There are only two true Wide ranging, but not common, species river-dwelling salamanders in the Great Smok- ies, the Hellbender (Cryptobranchus alle- Southern Appalachian Salamander (Pleth- ganiensis), known presently only from Little odon oconaluftee). River, Oconaluftee River, and Deep Creek

Figure 8. Middle Prong at Tremont.

14 Figure 9. Ideal habitat for Hellbenders in Lower Abrams Creek.

(Nickerson and others, 2002; Dodd, 2004) Larvae are found near the shore, and the adults (fig. 9), and the Common Mudpuppy (Necturus inhabit streambanks for at least part of the year. maculosus), known only from Little River and However, this species also inhabits some Abrams Creek. One additional salamander, the smaller streams, and it is by no means a Ariver- Junaluska Salamander (Eurycea junaluska), dwelling@ species. tends to be associated with some of the Park=s Creek and stream salamanders have lar- larger western and northwestern streams and vae that develop in the creeks and streams of the rivers on the Tennessee side of the Smokies. Park (figs. 10-12), whereas the adults may be

Figure 10. Small stream in unnamed tributary to Falls Branch.

15 aquatic, semi-aquatic, or even terrestrial to a greater or lesser degree. Many of these species are widespread in the Park because of the large number of creeks and streams available for colonization. A few species are found only at higher mountain elevations (for example, the Ocoee and Imitator Salamanders), whereas others are lowland species (Spotted Dusky, Three- lined, and Long-tailed Salamanders). Instead of a circumscribed area, their habitat is often lin- ear, following the streams and streamsides. The dusky salamanders (Desmognathus) are prominent in this group, but there are many exceptions to each habitat categorization listed below. Even Black-bellied Salamanders have been found well above the forest floor in rock crevices among boulders at considerable distances from water. Monitoring adults and larvae of these species requires very different techniques, and may require sampling very different types of habitats. Nearly aquatic species Shovel-nosed Salamander (Desmognathus marmoratus). Predominantly aquatic and streamside species Spotted Dusky Salamander (Desmognathus conanti); Seal Salamander (Desmognathus monticola); Black-bellied Salamander Figure 11. Medium-sized stream in normal flow at Roaring (Desmognathus quadramaculatus); Santeet- Fork. lah Salamander (Desmognathus santeetlah);

Many of these species are widespread in the Park because of the large number of creeks and streams available for colonization.

Figure 12. Medium-sized stream in high water at Whiteoak Flats Branch.

16 Junaluska Salamander (Eurycea junaluska). stage, and the adults are only found in wet The extremely high elevation areas where seeps. some streams first appear may be devoid of Cave inhabitants (fig. 13) salamanders if the water emanates from the Anakeesta rock formation (Dodd, 2004). Cave Salamander (Eurycea lucifuga). Species with aquatic larvae but are largely Known only from Stupkas Cave, the Calf terrestrial as adults caves, and one record from Whiteoak Sink. Imitator Salamander (Desmognathus imita- Other salamanders in the Smokies may live in tor); Ocoee Salamander (Desmognathus caves, especially around the entrances (Dodd ocoee); Three-lined Salamander (Eurycea and others, 2001). The larvae of some sala- guttolineata); Long-tailed Salamander manders (for example, E. longicauda in (Eurycea longicauda); Blue Ridge Two- Gregorys Cave) develop in pools well inside lined Salamander (Eurycea wilderae). cave passages (fig. 14). A few salamanders require very specialized habitats in the Great Smokies, or at least Rock face inhabitants (fig. 15) are usually found in these habitats. Some of Spotted Dusky Salamander (Desmognathus these species have larvae which are found in the conanti); Seal Salamander (Desmognathus same streams and creeks as the preceding species, although the adults prefer to leave the monticola). streams. Whereas the larvae may be relatively Permanent to near permanent wet rock easy to survey, adults often can be quite difficult walls with hiding places, particularly along to find with any regularity. One species, the trails, road cuts, and in the vicinity of waterfalls, Seepage Salamander, does not have a larval especially at lower elevations.

Figure 13. Entrance to Gregorys Cave.

17 few salamanders require Figure 14. Rimstone pools and cave pool at A Gregorys Cave. Salamander larvae develop very specialized habitats in in the pools, although they are unlikely to the Great Smokies, or at complete metamorphosis. least are usually found in these habitats.

Figure 15. Rock face near Double Gap.

18 Spring Inhabitants Spring Salamander (Gyrinophilus porphyriticus); Black-chinned Red Salamander (Pseudotriton ruber).

Inhabitants of swampy and mucky habitats (fig. 16) Mud Salamander (Pseudotriton montanus). Known only from a few scattered locations in the lowlands of the northern side of the Park.

Inhabitants of wet seepages (fig. 17) Seepage Salamander (Desmognathus aeneus). Known only from drainages on the southwestern side of the Park. Finally, there are salamanders that breed in ponds, and it is virtually only at this time that these species can be censused. Five species fall into this category: the Spotted Salamander (Ambystoma maculatum); Mar- bled Salamander (A. opacum); the rare Mole Salamander (A. talpoideum); Four-toed Salamander (Hemidactylium scutatum); and Eastern Red-spotted Newt (Notophthalmus viridescens). Breeding ponds are limited within the Park, being concentrated in Cades Figure 17. Seep at Big Spring Cove. Cove and nearby Big Spring Cove (the four Finley-Cane sinkhole ponds), the Cane Creek drainage, and at scattered localities between (at the Sinks and ditches along the road to Sugarlands and Cades Cove along Little River Tremont). These locations are on the Tennessee side of the Park. Although beaver ponds are found in Bone Valley and Big Cove in North Carolina, and small scattered ditches and wetlands occur in Cataloochee Valley, no pond salamanders are known to breed in them.

Mud Salamanders are known only from a few scattered locations in the lowlands of the Figure 16. Former trout pond mucky habitat in northern side of the Park. Cataloochee.

19 FROGS are used by frogs and toads: ponds (natural, as well as of beaver or human origin); woodland Frogs in the Great Smoky Mountains pools; grassy ditches, pools, and rivulets; and National Park require water for breeding and larger streams and rivers. for tadpole development. As such, the diversity and distribution of frogs are not as great in the Ponds mountains as in the adjacent lowlands of the Pond distribution is limited in the Tennessee Valley and Atlantic Coastal Plain. In Smokies, being confined mostly to Cades Cove, the Smokies, four major types of breeding sites Big Spring Cove, and two beaver ponds. The

Figure 18. Gum Swamp at Cades Cove in high water.

Figure 19. Gum Swamp at Cades Cove when dry.

20 most important frog-breeding ponds are Gum Northern Green Frog (Rana clamitans), Swamp (figs. 18, 19), Gourley Pond (figs. 20, Pickerel Frog (R. palustris), Wood Frog (R. 21), Methodist Church Pond (fig. 22), and the sylvatica), and Eastern Spadefoot (Scaphiopus sewage-treatment pond (all in Cades Cove); the holbrooki), known only from Gum Swamp. four sinkhole ponds in Big Spring Cove (also American Bullfrogs (R. catesbeiana) also have known as the Finley-Cane ponds); and the bea- been heard at the beaver pond in Big Cove. ver ponds in Bone Valley and Big Cove Some of these ponds dry completely as the (fig. 23). Species that commonly use these summer progresses, particularly Gum Swamp ponds are the American Toad (Bufo america- (fig. 19), Gourley Pond (fig. 21), and the nus), Cope=s Gray Treefrog (Hyla chrysoscelis), Finley-Cane sinkhole ponds.

Figure 20. Gourley Pond at Cades Cove in high water.

Figure 21. Gourley Pond at Cades Cove when dry.

21 Figure 22. Methodist Church Pond at Cades Cove.

Figure 23. Beaver pond at Big Cove.

22 Woodland Pools Woodland pools are scattered at various areas within the Park. They range from a few centimeters deep to about 0.5 m, and they usually dry as summer progresses. Woodland pools are located in level ground at Cosby, Sugar- lands, Metcalf Bottoms, Big Spring Cove, Little Cataloochee Valley, throughout the Cane Creek drainage (fig. 24), Cades Cove (especially along Abrams Creek at the western edge of the cove), and doubtless in other areas of the Park. Amphibians that use these small pools for breeding include the Eastern Red-spotted Newt (Notophthalmus viridescens), American Toad (Bufo americanus), Cope=s Gray Treefrog (Hyla chrysoscelis), Northern Green Frog (Rana clamitans), Pickerel Frog (R. palustris), and Wood Frog (R. sylvatica).

Grassy Ditches, Pools, and Rivulets Grassy ditches, pools, and rivulets are generally shallow, open-canopied habitats, with a grassy vegetation where concealment and breeding sites are available (fig. 25). Only two places in the Park contain much of this habitat: Cades Cove and Cataloochee Valley. Frogs found here include the American Toad (B. americanus); Eastern Narrow- mouthed Toad (Gastrophryne carolinensis), known only from grassy pools at the Abrams Creek Ranger Station and at Shields Pond in Cades Cove; Spring Figure 24. Woodland drainage pool at Cane Creek. Peeper (Pseudacris crucifer); Upland Chorus Frog (Pseudacris feriarum); and the ubiquitous Wood Frog (R. sylvatica). These habitats normally dry rapidly with Smokies, the American Bullfrog=s (R. catesbei- the warm weather, although the rivulets and ana) large tadpoles are conspicuous in Abrams some pools in Cades Cove may persist well into Creek near the Abrams Creek Ranger Station. summer. Additional species, such as Fowler=s Toad (B. fowleri), breed in the backwaters formed Streams and Rivers from flooding along streams and rivers. Other frogs, such as Northern Green Frogs (R. clami- A few species of frogs breed in the shal- tans), are found along streambanks during the lows of rivers and larger streams. In the Great nonbreeding seasons.

23 Table 2. Identification and Life History of the Frogs of Great Smoky Mountains National Park [<, less than; >, greater than; mm, millimeter; cm, centimeter; m, meter; m2, square meters] Breeding Metamorph Species Eggs Tadpole description Larval period times size Acris eggs deposited singly; 1 gelati- a medium-sized light to medium-gray tadpole; April to June, 35-70 days, 10-15 mm crepitans nous envelope, >2.3 mm in throat light; tail musculature mottled or reticu- possibly into based on Acris diameter; deposited in shallow lated; usually a very distinctive "black flag" on July crepitans blan- water among stems of grass or the tail tip; tail long and narrow; anus dextral (to chardi on bottom; 250 eggs per com- the right); oral disk emarginate; most 30-36 mm plement total length, rarely to 46 mm Bufo eggs in strings with gelatinous body round or oval in dorsal view; eyes dorsal spring 50-65 days 7-12 mm americanus casings; 2 envelopes present; (looks cross-eyed); nostrils large; color dark (March-April) strings long, to 60 m; 15-17 brown to black; dorsal portion of the body uni- eggs per 25 mm; 4,000-12,000 colored; venter with aggregate silvery or cop- eggs on bottom of quiet pools per spots; snout sloping in lateral view; tail musculature distinctly bicolored; anus medial (in the center); spiracle is distinctly on left side of body B. fowleri eggs in strings with gelatinous body round or oval in dorsal view; eyes dorsal April to July 40-60 days 7.5-11.5 mm casings; 1 envelope present and (looks cross-eyed); nostrils large; color dark; <5 mm in diameter; strings 2.4- dorsal portion of body slightly mottled; snout 3 m with 17-25 eggs per 25 rounded in lateral view; tail musculature often mm; 5,000-10,000 eggs; in tan- not distinctly bicolored; anus medial (in the gled mass around vegetation center); spiracle is distinctly on left side of body Gastrophryne eggs in small surface film that a small jet-black tadpole with lateral white to mid-May to 20-70 days 8.5-12 mm carolinensis has a mosaic structure; enve- pink stripes on posterior portion of body mid-August lope a truncated sphere; mass extending to the tail musculature. Viewed from round or square; 10-150 eggs the side, the head comes to a point; body round per mass; in any depression in dorsal view; eyes wide set and lateral; anus with water, but not deep pools median; jaws do not have keratinized sheaths, and the oral disc and labial teeth are absent Hyla eggs in small surface film, but small to medium-sized grayish tadpole with a April to June, 45-65 days 13-20 mm chrysoscelis envelope not in truncated high dorsal tail fin; dorsal tail fin height equal but calls occa- sphere; no mosaic structure; 5- to or greater than musculature height; tail long, sionally heard 40 eggs per mass; in shallow with black blotches; background color of at other times ponds attached loosely to vege- mature tail orange to scarlet; throat rarely pig- of the year tation, or free. Air bubbles mented; dorsal fin never extends anterior to present. midway between the spiracle and eye; anus dextral (to the right); oral disk not emarginate Pseudacris eggs deposited singly in shal- a small-sized deep-bodied tadpole with a late winter to 90-100 days 9-14 mm crucifer low water near bottom among medium-sized tail; tail musculature mottled; early spring vegetation; one gelatinous enve- fins clear or with blotches; no dots on body; (February to lope. snout square when viewed dorsally; anus dex- April); calls tral (to the right); oral disk not emarginate occasionally heard at other times of the year P. feriarum egg mass in lump, but loose small olive to black tadpole with a bronze February to 50-60 days 8-12 mm irregular cluster; 1 envelope, belly; tail medium; anus dextral (to the right); April. 3.6-4.0 mm; deposited in oral disk not emarginate; tadpoles develop marshy areas and pools in mat- rapidly ted vegetation

24 Table 2. Identification and Life History of the Frogs of Great Smoky Mountains National Park (Continued) [<, less than; >, greater than; mm, millimeter; cm, centimeter; m, meter; m2, square meters] Breeding Metamorph Species Eggs Tadpole description Larval period times size Rana eggs in large surface film in large olive to grayish green tadpole with small late spring 1-2 years 31-59 mm catesbeiana form of a disc; 10,000-12,000 widely spaced small spots (dots) covering the and through- eggs per disc; deposited among body and tail; venter straw; eyes bronze; body out the sum- water plants or brush; 1 gelati- oval and round in dorsal view; eyes dorsal or mer. Calls nous envelope dorsolateral; nostrils small compared with may be heard eyes; lower jaw wide; anus dextral (to the at other times right); oral disk emarginate of the year R. clamitans eggs in surface film; mass large (but not deep bodied) olive green tadpole late April to to 1 year 23-38 mm <0.09 m2; 1,000-5,000 per with large dark spots, generally with a white late July or mass; attached to vegetation or throat; belly deep cream without iridescence; even early free; 2 gelatinous envelopes body oval and round in dorsal view; eyes dor- August. Calls sal or dorsolateral; nostrils small compared may be heard with eyes; tail green mottled with brown; at other times lower jaw wide; anus dextral (to the right); oral of the year disk emarginate R. palustris eggs in firm regular cluster; large, full, deep-bodied tadpole; olive green late winter to 70-80 days 19-27 mm brown above and yellow below; shading through yellow on sides; venter cream, spring (mid- mass a sphere 38-100 mm in back marked with fine black and yellow spots; March-April) diameter; 2 envelopes present; belly with blotches of white; venter iridescent, 2,000-4,000 eggs; mass depos- viscera visible; tail very dark, black blotches ited 75-100 mm to 91 cm under can aggregate to purple-black; body oval and water; attached to debris and round in dorsal view; eyes dorsal or dorsolat- vegetation eral; nostrils small compared with eyes; lower jaw narrow; anus dextral (to the right); oral disk emarginate R. pipiens mass a firm regular cluster; large, deep-bodied tadpole; dorsally dark probably early 60-80 days 18-31 mm 3,500-6,500 eggs close together brown, covered with small gold spots; belly March to in mass; 2 envelopes present; deep cream, with bronze iridescence; viscera early May outer envelope 5 mm; eggs visible; throat translucent and more extensive black above and white below; than Pickerel Frog; similar in appearance to deposited near surface, usually Green Frog, but darker; body oval and round in attached to grasses and vegeta- dorsal view; eyes dorsal or dorsolateral; nos- tion, sometimes free trils small compared with eyes; lower jaw narrow; anus dextral (to the right); oral disk emarginate R. sylvatica eggs in firm regular cluster; medium-sized tadpole with usually very dark winter and 45-85 days 16-18 mm black above and white below; to gray coloration, and with a faint light stripe early spring mass a sphere 38-100 mm in of cream, white or gold along the upper jaw (mid-Decem- diameter; 2 envelopes present; (like a mustache); venter cream with belly ber to March) 2,000-4,000 eggs; mass depos- slightly pigmented at sides; body oval and ited 75-100 mm to 91 cm under round in dorsal view; eyes dorsal or dorsolat- water; attached to debris and eral; nostrils small compared with eyes; anus vegetation dextral (to the right); oral disk emarginate; tail quite long; dorsal crest high extending on to body Scaphiopus eggs in loose irregular cylinder a small dark tadpole, bronze to brown with only heard 14-60 days 8.5-12 mm holbrooki or band; mass 25-75 mm wide close-set tiny orange spots; body round or oval calling once and 25-305 mm long; deposited in dorsal view; eyes close-set and dorsal, iris (July 12, on stems of plants/grass; black; head wide relative to body width; tail 1999). Proba- 1 gelatinous envelope; 200 per short, with tip blunt and rounded; anus medial bly any time packet (in the center); spiracle is ventrolateral. Often from March to found in "schools" of hundreds of tadpoles October

25 Figure 25. Grassy pool at Cades Cove.

Other Breeding Sites Life History

Four minor types of wetlands and aquatic Terrestrial Salamanders (Plethodon- sites are used occasionally by frogs for breeding tidae). The life cycle of terrestrial plethodontids in the Great Smokies. American toads (B. amer- takes place in a multidimensional space. Natu- icanus) breed in the backwaters along the north ralists tend to think of salamanders as surface- shore of Fontana Reservoir, although reser- dwelling, but surface activity is only a small voirs (fig. 26) are generally depauperate of part of the life cycle of a terrestrial salamander. amphibians. Small, usually closed-canopied, Most terrestrial species probably do not have a swampy and mucky wetlands (for example, very large home range on the ground surface, including beneath debris and litter. They spend those found along Indian Creek, at Smokemont, a considerable part of their lives underground, and at the old trout pond in Cataloochee; see and biologists really know very little about their fig. 16) are used by Wood Frogs (R. sylvatica). life history, especially their time spent under- Wood Frogs are quite variable in their choice of ground and the depth and range of underground breeding sites, even to depositing eggs in lateral movement. In addition, terrestrial spe- human-enlarged spring pools and roadside cies occasionally become arboreal during the ditches. Indeed, virtually any pool in late winter night or under rainy conditions; salamanders to early spring is likely to be colonized by often take refuge under loose bark. Salamanders breeding Wood Frogs. at different life stages may remain nearly

26 Figure 26. Chilhowee Lake at mouth of Abrams Creek.

entirely underground (tiny juveniles perhaps; (ponds, ditches) free of fishes. For semi-aquatic adults during egg deposition and mating) or on plethodontids, breeding sites include seeps and the surface (adult feeding and territoriality, streams from little trickle trails to sizeable environmental conditions permitting). It is by streams or rivers. Adults (mole salamanders and no means clear that space is used similarly by newts) may migrate synchronously to breeding different life stages. Thus, detection probabili- sites in a quite orderly fashion, although tempo- ties may change with life stage within a habitat. rally constrained to one or a few nights during The eggs of some terrestrial species have never the breeding season. Breeding adults and egg been seen, and nests have been located only masses can be censused, but herpetologists with extreme infrequency. Some plethodontids know little about what proportion of a popula- may be long-lived (5-10 years). tion breeds annually, and from what area they Semi-Aquatic Salamanders (Ambysto- are drawn. Males and females may not stay for matidae, Plethodontidae, Salamandridae). All equal amounts of time during the breeding sea- attributes that apply to terrestrial salamanders son, even when the breeding season is extended. apply to semi-aquatic salamanders in terms of Stream-breeding species may live perma- surface and underground habitat use. nently in the streams (Desmognathus marmora- Semi-aquatic salamanders, however, require tus), streamsides (many other Desmognathus), water for reproduction. For mole salamanders or at various distances from the stream (D. imi- (Ambystoma) and newts (Notophthalmus), tator, Gyrinophilus, Pseudotriton). Distances breeding sites are usually standing water may range from a few meters to hundreds of

27 meters away, and breeding migrations are not because the larvae are little known and, for the synchronized. Little is known about spatial dis- most part, rarely seen. tribution during terrestrial nonbreeding times. Frogs. All of the frogs in the southern For some species (for example, Hemidactylium Appalachians have a “typical” amphibian life scutatum) virtually nothing is known about their cycle. Adults move to a breeding site, deposit lives away from woodland pools and streams/ eggs that hatch into larvae (tadpoles), metamor- ditches outside of the breeding season. For cer- phose to juveniles, disperse, and grow until they tain species (D. quadramaculatus) adults can be are ready to repeat the cycle. For most species, censused streamside, whereas adults of other however, many questions about the life cycle species (D. imitator) can be readily found in ter- remain unanswered (what percentage is breed- restrial habitats; some species (Pseudotriton) ing in any one year, where do juveniles go, how can be found terrestrially as adults usually only far do adults disperse). Larval periods may be by luck, and the adults of a few species extremely brief (days in Scaphiopus) to (egg-brooding adult female Hemidactylium) are extremely long (years in some Rana). Breeding observed only during the breeding season. may be synchronous (spadefoots, many ranids) All eggs of semi-aquatic salamanders are or extended (Rana catesbeiana). Even when deposited in water, and the egg masses of some synchronous and explosive (Rana sylvatica), species (Ambystoma) can be cen- the actual breeding date may extend over a sused easily. All period of months (December to semi-aquatic species have March) as adults wait for the larvae which remain in a right combination of environ- larval stage from a few mental conditions. Adults months to as long as 2-3 years. (and perhaps juveniles) of Paedomorphosis (the ability to many frog species spend breed while maintain- most of their lives away ing a larval appear- from the breeding sites. ance) occurs in a few Individuals have been species (Ambystoma found hundreds (or even talpoideum) under favorable con- thousands) of meters ditions, but no salamanders from from the nearest breeding the Park are known to be paedo- sites. Frogs are often exceptionally morphic. Larvae metamorphose and pre- hard to locate outside the breeding season, sumably take up adult habits, but nothing is much less to sample them. However, the terres- known concerning dispersion for most species. trial sites are extremely important to survival Maturation can range from one to many years, since individuals spend most of their lives as depending on species. Individuals of some spe- terrestrial predators. cies (Ambystoma, Notophthalmus, large Des- mognathus) may live 10-15 or more years. Although most species of frogs call dur- Aquatic salamanders (Cryptobranchidae, ing the breeding season, some species do not or Proteidae). Little is known about the life history they have only weak voices that do not carry far. of most of these species, except for Cryptobran- Calling times are variable among species; some chus. Species within these families are entirely call during the day, some call at dusk and during aquatic. The spatial use of habitat is largely the early evening, and some call only between unstudied except for Hellbenders, which are midnight and early dawn. Some species call known to have home ranges and to guard nest- only during rains, whereas others will call most ing sites. Fully aquatic species (Cryptobran- evenings of the breeding season. Some frogs chus, Necturus) inhabit medium to large breed in winter (even in the mountains of the streams and rivers in the southern Appala- South), others breed in the spring or summer, chians. Hellbenders may live 25 or more years. whereas others call during an extended breeding Nothing, however, is known about longevity of season. Calling times and seasons also vary lat- the Common Mudpuppy (Proteidae: Necturus), itudinally and perhaps with elevation.

28 Areas of Particular Amphibian Species genus, species complex, or as “unknown” in Richness field notes. One of the best ways to identify sala- Three areas within Great Smoky Moun- mander and frog larvae, in addition to color and tains National Park are particularly rich in morphology, is to examine their habitats and the amphibians. Two (Cades Cove, Cane Creek times of year they are found. This can most eas- drainage) are lowland sites, whereas the third is ily be done through a comparative table. Mor- the high-elevation spruce-fir forest. The low- phological and life history characteristics are land sites are similar in amphibian species com- listed in tables 1 and 2 to help field biologists position; they are rich in species because they identify the species that are being examined. are the only two sizeable lowland areas within These data can be used in conjunction with the the Park with a large variety of wetlands. As information in Dodd (2004). such, they contain most of the frogs and pond- breeding salamanders. Both areas share species SALAMANDERS affinities with the herpetofauna of the Tennes- see Valley, from whence lowland amphibians All salamanders in the Great Smoky colonized Cane Creek and Cades Cove (via Mountains have four limbs with four (Necturus, Abrams Creek). On the other hand, the high- Hemidactylium) or five (all others) toes on each elevation amphibians are composed entirely of hind foot. They all have tails, lack dry scales salamanders, and two species (Plethodon jor- covering the body (lizards have dry scales), and have skins that are moist or wet to the touch. dani, Desmognathus imitator) are virtually The skins of a few species, such as Jordan=s endemic to the Park (D. imitator is found also in Salamander (Plethodon jordani), are sticky the Plott Balsams). Other high-elevation species because of glandular secretions, but only the in the spruce-fir forest (for example, D. ocoee, Hellbender and Common Mudpuppy are truly D. wrighti, P. metcalfi) are found in other slimy. restricted regions of the Southern Appalachians. These three areas should be the special focus of Biologists take two standard measure- amphibian monitoring activities. ments with regard to length. The total length (TL) is the length of an animal from the tip of the snout to the tip of the tail. Because some Identification salamanders lose their tails (or parts thereof) to predation, another common measurement Most biologists working at Great Smoky recorded is the snout-vent length (SVL). SVL is Mountains National Park should be able to measured from the tip of the snout to the poste- identify the majority of the amphibians that they rior portion of the vent (the opening of the clo- observe by using a combination of the color aca, the common receptacle for the digestive, photographs, species descriptions, and identifi- excretory, and reproductive tracts). All scien- cation/life history tables found in this manual tific measurements are recorded in metric units, and in Dodd (2004). Some individual animals usually millimeters. may be impossible to identify with certainty. Salamander larvae sometimes are divided Larvae, especially small salamander larvae and into two general groups, depending on mor- tadpoles, often cannot be distinguished without phology and the type of wetland in which they microscopic examination. Adult salamanders, develop. The pond form (fig. 27A) is stout bod- especially the duskies (Desmognathus), are ied, with long filamentous gills and a wide dor- notoriously variable with overlapping pheno- sal fin which extends well onto the body. Mole typic and genotypic characters. Field biologists salamanders (Ambystoma), for example, have have found it increasingly difficult to place this type of larva. Pond larvae develop in still some individual animals into a species category water, and use the extra surface area of the body because of the range of genetic and color varia- and fin as aids in swimming. Stream larvae tion observed in natural populations. As a (Eurycea, Pseudotriton) are slimmer than pond result, sometimes an animal must be recorded to larvae, with more streamlined bodies, shorter 29 gills, and a narrower tail fin that does not extend white line bordered by black lines, that runs from onto the body (fig. 27B). These larvae usually in front of each eye to the nostrils. Salamanders live in swift flowing water, where extra surface of the genus Pseudotriton do not have this line. area on the body would be a distinct disadvan- Spring Salamanders use the canthus rostralis as a tage. “gunsight” to zero in on prey. A number of useful characters are avail- Plethodontidae versus all other sala- able which can be used to identify salamanders mander families: All lungless salamanders have to genus or family. A few illustrative examples a nasolabial groove that extends from each nos- are provided, but more detailed comparisons are tril to the upper jaw. The nasolabial groove found in Dodd (2004) under the heading “Sim- transmits chemicals to the salamander from the ilar Species.” substrate; no other salamander family has this Desmognathus: All dusky salamanders groove. have a light line which extends from the back of the eye to the angle of the jaw. The duskies also FROGS have well-developed muscles on the sides of Like most salamanders, frogs have four their heads. They need these muscles to raise legs with four toes on the front limbs and five the upper jaw in order to open their mouths, toes on the rear limbs. The hind limbs are much since the lower jaw is fused to the skull. larger than the front limbs, and are used to pro- Gyrinophilus versus Pseudotriton: pel the body when walking, hopping, or jump- Although these colorful salamanders are superfi- ing. Frogs are measured in TL, that is, from the cially similar in appearance, Spring Salamanders tip of the snout to the end of the body between (Gyrinophilus) have a canthus rostralis, a large the hind limbs (that is, at the end of the

gill fimbriae dorsal fin gill rachis

gular flap costal grooves (A) Pond form antlerlike gills tail fin

(B) Stream form

Figure 27. Body morphology of a salamander larva: (A) pond form; (B) stream form.

30 BL TAL

Narial aperature

MTH TMH

Anus Oral disc Spiracle TL (A) Lateral view EXPLANATION BL body length IOD IND inter narial distance IOD inter orbidal distance MTH maximum tail height TAL tail length TMW TL total length TMH tail muscle height TMW tail muscle width

IND (B) Dorsal view

Figure 28. Body morphology of a tadpole.

urostyle). Of course, there are many other oral disk consists of the mouth parts; the narial measurements which could be made, such aperture is the opening to the nostrils; the as the length of the various sections of the spiracle is the opening from the gills (water is hind limb, but these data generally are not taken in through the mouth, passes over the important in amphibian monitoring-programs gills, and is expelled via the spiracle); the anus except in studies of fluctuating asymmetry is the opening from the digestive tract. The total (Alford and others, 1999, but see McCoy and length (TL) consists of the body length (BL) Harris, 2003). and tail length (TAL). Sometimes additional Tadpoles are morphologically complex. morphological measurements are taken, such as As with salamander larvae, there are two gen- the maximum width of the tail musculature eral tadpole body types, the pond type and the (TMH) or the maximum tail depth (MTH). The stream type. Pond-type tadpoles have deeper location and size of these characters, or their bodies and higher tail fins than do stream-type ratios in relation to one another, are useful in tadpoles. Structures important in the identifica- identifying what otherwise appears to be just tion of tadpoles are labeled on figure 28. The another drab, olive-green, or black tadpole.

31 Labial teeth The tadpoles of different A-1 species of frogs often appear iden- A-2 tical to one another, but the structure of their mouthparts readily separate them. Biologists may SM need to examine mouthparts to determine which species is in hand. For this reason, a diagram Not MP has been included of tadpole Emarginate mouthparts is provided in figure 29. The nomenclature fol- lows Altig and McDiarmid (1999). PL LP The location, number, and degree of separation among labial teeth and papillae are important charac- LJ ters for identifying tadpoles. Examining tadpole oral disks A-2 Gap (sometimes incorrectly termed “teeth”) also gives researchers an opportunity to check the health of AL the tadpole. For example, the horny jaw sheaths drop out when the tadpole is exposed to certain Emarginate toxic compounds and to the dan- gerous disease, chytridiomycosis. However, tadpoles should not be

UJ cuspate P-1 P-3 P-2 EXPLANATION AL anterior (upper) labium A-1 and A-2 first and second upper tooth rows A-2 GAP medial gap in second anterior tooth row LJ lower jaw sheath LP lateral process of upper jaw sheath M mouth MP marginal papilla OD oral disk PL posterior (lower) labium P-1, P-2, P-3 first, second, and third posterior (lower) tooth rows SM submarginal papilla UJ upper jaw sheath

Figure 29. Oral disc (mouthparts) of a tadpole.

32 held too long before examination or preserva- Rana palustris versus R. pipiens: these tion, since some tadpoles may shed denticles in very similar frogs are both green and spotted. In the laboratory. R. palustris, the spots are squarish, paired and As with salamanders, there are certain of nearly equal size, whereas in R. pipiens they useful defining characteristics that help to iden- are smaller, rounded, and more randomly scat- tify certain superficially similar animals. Some tered on the frog=s back. of these are listed below (also see “Similar Spe- cies” in Dodd, 2004). Bufonidae versus Pelobatidae: Frogs in Additional Information both of these families are terrestrial. However, Information on the etymology, identifica- the true toads (Bufo) are dry-skinned and tion of adults, larvae, and eggs, similar species “warty,” and have prominent cranial crests and and how to differentiate them, taxonomic prob- parotoid glands. The spadefoot toads (Scaphio- lems, distribution both within the Park and else- pus) are smooth-skinned, lack cranial crests and where in North America, life history, abundance parotoids, and have a sharp digging spade on their hind feet. and status, and remarks on interesting aspects of the biology of the species are found in Dodd Hylidae versus other frog families: all hylid frogs (Acris, Pseudacris, Hyla) in the (2004). Data on 44 amphibians are presented, Great Smokies have slightly to completely including information on species no longer expanded toepads, but only in the treefrogs thought present (for example, Aneides aeneus) (Hyla) are these greatly expanded for climbing; or which were reported historically from the the other hylids are mostly ground-dwelling Park, but whose actual occurrence may be (however, note that Spring Peepers, Pseudacris doubtful (Acris crepitans). Distribution maps, crucifer, often call from the trees from late fall color photographs of amphibians from the Park, to early spring before descending to breeding and original color illustrations accompany each ponds). account.