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Aquatic Fauna in Peril: The Southeastern Perspective

Edited By George W. Benz And David E. Collins

Design by Lenz, Inc. Decatur, Georgia.



Aquatic Fauna in Peril > Amphibians in Peril: Resource Management in the Southeast

Amphibians in Peril: Resource Management in the SoutheastIllustration by Tom Tarpley.

By Linda V. LaClaire

In this chapter I will review past and current resource management programs that have benefited amphibians in the southeastern United States and management strategies that could be used to conserve the Southeastís imperiled amphibian fauna. Many of the projects and concepts discussed here are relatively new and much of the information provided was obtained from the "gray" literature (e.g., various federal, state or private agency reports) or from unpublished data. Nonetheless, this information provides a significant foundation upon which to build the consensus needed to ensure that the modest strides made in amphibian management will continue. Conservation of our imperiled amphibian fauna will require a commitment from resource managers to include amphibians in management planning at local, regional, national, and global levels.

Amphibian life histories grade from those that are totally aquatic to those that are entirely terrestrial; however, all amphibian species require a moist environment. Management efforts needed to protect the watersheds and wetlands inhabited by aquatic amphibians are often the same ones required to protect the moist woodland environments that are home to many terrestrial amphibians. Moreover, because water quality in wetlands is often dependent on conditions prevailing in surrounding terrestrial habitats, and because many amphibians depend on both aquatic and terrestrial habitats, the management of aquatic and terrestrial habitats can seldom be separated from one another when considering conservation measures aimed at amphibian resources. It is for this reason that this chapter will consider all imperiled amphibians in the southeastern United States, no matter what their degree of connection to aquatic habitats. The southeastern United States is defined as the region composed by the states of Alabama, Arkansas, Florida, Georgia, Kentucky, Louisiana, Mississippi, North Carolina, South Carolina, and Tennessee.

In this chapter, management is considered in broad terms as those techniques and activities which strive to maintain the continuity of ecological and evolutionary processes (OíConnell and Noss, 1992). Because habitat loss is the primary threat to imperiled amphibians in the Southeast, the development of effective habitat management plans is crucial for amphibian conservation. Unfortunately, few management projects have been specifically directed at amphibian species or populations in the Southeast or elsewhere throughout the United States.

Table 1. Imperiled amphibian fauna of the southeastern United States.1 This list contains amphibians listed under the Endangered Species Act and other species of concern (former Category 2 candidate species), as well as those listed by a state, or ranked as G1-G3 by The Nature Conservancy and Natural Heritage Programs (TNC). STATE LIST CATEGORIES: WM = wildlife in need of management; R = rare; SC = species of special concern; T = threatened; E = endangered; NL = not listed; - = not resident in state. FEDERAL RANK: SC = species of concern (former Category 2 candidate species); T = threatened; NR = not ranked. TNC RANK: G3 = rare or uncommon; G2 = imperiled; G1 = critically imperiled; Q = taxonomy questioned; ? = ranking inexact; NR = not ranked.
Amphibian Taxa
State Lists
Federal Rank
TNC Rank
AL
AR
FL
GA
KY
LA
MS
NC
SC
TN
SALAMANDERS
Cryptobranchidae
Cryptobranchus alleganiensis
(eastern hellbender)
NL
NL
-
R
NL
-
NL
SC
NL
WM
SC
G32
Proteidae
Necturus sp.
(Black Warrior waterdog)
NL
-
-
-
-
-
-
-
-
-
SC
G2
N. lewisi
(Neuse River waterdog)
-
-
-
-
-
-
-
SC
-
-
NR
G3
N. maculosus
(mudpuppy)
NL
-
-
NL
NL
NL
NL
SC
-
NL
NR
NR
Amphiumidae
Amphiuma pholeter
(one-toed amphiuma)
NL
-
NL
R
-
-
-
-
-
-
NR
G3
Sirenidae
Pseudobranchus striatus belli
(Everglades dwarf siren)
-
-
NL
-
-
-
-
-
-
-
NR
G3?
P. s. lustricolus
(Gulf hammock dwarf siren)
-
-
NL
-
-
-
-
-
-
-
SC
NR
P. s. spheniscus
(slender dwarf siren)
-
-
NL
NL
-
-
-
-
-
-
NR
G3?
P. s. striatus
(broad-striped dwarf siren)
-
-
NL
NL
-
-
-
-
T
-
NR
G3?
Ambystomatidae
Ambystoma cingulatum
(flatwoods salamander)
NL
-
NL
R
-
-
-
-
E
-
SC
G2/G3
A. talpoideum
(mole salamander)
NL
NL
NL
NL
NL
NL
NL
SC
NL
WM
NR
NR
A. tigrinum tigrinum
(eastern tiger salamander)
NL
NL
NL
NL
NL
NL
NL
T
NL
NL
NR
NR
Salamandridae
Notophthalmus perstriatus
(striped newt)
-
-
NL
R
-
-
-
-
-
-
SC
G2/G3
Plethodontidae
Desmognathinae
Desmognathus aeneus
(seepage salamander)
NL
-
-
NL
-
-
-
NL
-
NL
SC
NR
D. apalachicolae
(Apalachicola dusky salamander)
NL
-
NL
NL
-
-
-
-
-
-
NR
G3
D. brimleyorum
(Ouachita dusky salamander)
-
NL
-
-
-
-
-
-
-
-
SC
NR
D. santeetlah
(Santeetlah dusky salamander)
-
-
-
-
-
-
-
NL
-
NL
NR
G3/Q
D. welteri
(Black Mtn. dusky salamander)
-
-
-
-
NL
-
-
-
-
WM
NR
NR
Phaeognathus hubrichti
(Red Hills salamander)
T
-
-
-
-
-
-
-
-
-
T
NR
Plethodontinae
Aneides aeneus
(green salamander)
NL
-
-
R
NL
-
E
E
NL
WM
SC
NR
Eurycea aquatica
(dark-sided salamander)
NL
-
-
NL
-
-
-
-
-
-
SC
NR
E. junaluska
(Junaluska salamander)
-
-
-
-
-
-
-
SC
-
NL
SC
G2/Q
E. longicauda longicauda
(longtail salamander)
NL
NL
NL
-
NL
-
-
SC
-
NL
NR
NR
E. lucifuga
(cave salamander)
NL
NL
NL
-
NL
-
E
-
-
NL
NR
NR
E. quadridigitata
(dwarf salamander; silver morph)
-
NL
-
-
-
NL
-
SC
-
-
NR
NR
E. tynerensis
(Oklahoma salamander)
-
NL
-
-
-
-
-
-
-
-
NR
G3
Gyrinophilus palleucus
(Tennessee cave salamander)
NL
-
-
NL
-
-
-
-
-
T
SC
G2
G. porphyriticus porphyriticus
(spring salamander)
NL
-
-
NL
NL
-
E
NL
-
NL
NR
NR
Haideotriton wallacei
(Georgia blind salamander)
-
-
SC
T
-
-
-
-
-
-
SC
G2
Hemidactylium scutatum
(four-toed salamander)
NL
NL
NL
NL
NL
NL
NL
SC
NL
WM
NR
NR
Plethodon aureolus
(Tellico salamander)
-
-
-
-
-
-
-
NL
-
NL
NR
G2/G3/Q
P. caddoensis
(Caddo Mountain salamander)
-
NL
-
-
-
-
-
-
-
-
NR
G2
P. dorsalis
(zigzag salamander)
NL
NL
NL
-
NL
-
NL
SC
-
NL
NR
NR
P. fourchensis
(Fourche Mountain salamander)
-
NL
-
-
-
-
-
-
-
-
NR
G2
P. kisatchie
(Louisiana slimy salamander)
-
NL
-
-
-
NL
-
-
-
-
NR
G3/Q
P. ouachitae
(Rich Mountain salamander)
-
NL
-
-
-
-
-
-
-
-
NR
G2/G3
P. petraeus
(Pidgeon Mountain salamander)
-
-
-
R
-
-
-
-
-
-
NR
G1/G2
P. teyahalee
(southern Appalachian salamander)
-
-
-
NL
-
-
-
NL
NL
NL
NR
G3/Q
P. websteri
(Webster's salamander)
NL
-
-
NL
-
NL
NL
-
E
-
NR
NR
P. wehrlei
(Wehrle's salamander)
-
-
-
-
NL
-
-
T
-
NL
NR
NR
P. welleri
(Weller's salamander)
-
-
-
-
-
-
-
SC
-
NL
NR
NR
P. yonahlossee3
(Yonahlossee salamander)
-
-
-
-
-
-
-
SC
-
NL
NR
NR
FROGS
Hylidae
Hyla andersonii
(Pine Barrens treefrog)
NL
-
SC
-
-
-
-
-
T
-
NR
NR
H. gratiosa
(barking treefrog)
NL
-
NL
NL
NL
NL
NL
NL
NL
WM
NR
NR
Pseudacris brachyphona
(mountain chorus frog)
NL
-
NL
-
NL
-
NL
SC
-
NL
NR
NR
P. streckeri illinoensis
(Illinois chorus frog)
-
NL
-
-
NL
-
-
-
-
-
SC
NR
Ranidae
Rana capito aesopus
(Florida gopher frog)
-
-
SC
NL
-
-
-
-
-
-
SC
G3
R. c. capito
(Carolina gopher frog)
-
-
-
NL
-
-
-
SC
NL
-
SC
NR
R. c. sevosa
(dusky gopher frog)
NL
-
SC
-
-
NL
E
-
-
-
SC
G3
R. heckscheri
(river frog)
NL
-
NL
NL
-
-
NL
SC
NL
-
NR
NR
R. okaloosae
(Florida bog frog)
-
-
SC
-
-
-
-
-
-
-
SC
G2
1 Species names and distributions based on Conant and Collins, 1991.
2 Arkansas and Missouri populations only.
3 Listed in North Carolina as Plethodon longicrus, the crevice salamander.

Most amphibian management has involved species listed under the Endangered Species Act or those species listed by individual states. Priorities determining the allocation of management resources are often based on these federal and state lists (Master, 1991). In the past, the federal candidate species list (U.S. Fish and Wildlife Service, 1994b) was used to identify species that may be biologically vulnerable and in need of active management. However, the U.S. Fish and Wildlife Service (1996) has revised its list of candidate taxa to include only those species which have been proposed for listing or which were previously considered Category 1 candidates. Category 1 candidates were those species for which sufficient data were available to support a proposed rule, but issuance of the rule was precluded by other listing activity. Category 2 candidates, those species which were being assessed for biological vulnerability and threat, are no longer included on the candidate list; these taxa are considered species of concern. Another prioritization system has been developed by The Nature Conservancy (TNC) in cooperation with the Natural Heritage Data Center Network (NHDCN) (NHDCN, 1993). Numeric ranks given for species with potential global biological vulnerability include: G1 = critically imperiled; G2 = imperiled; G3 = rare or uncommon; and G4 = widespread, abundant, and apparently secure, but with cause for long-term concern. This system has been used to develop lists of species with ranked levels of imperilment (TNC and the International Network of Natural Heritage Programs and Conservation Data Centers, 1996).

In the following discussion I will consider "imperiled" amphibians of the Southeast to be those species which are listed under the Endangered Species Act or by a state management agency, former federal Category 2 candidate species, and G1-G3 TNC/NHDCN ranked species (see Table 1). While extensive data are not available to support declines in all of these species, there is general concern among biologists that without effective management these amphibians will truly be in peril.

Under the Endangered Species Act, only one southeastern amphibian is listed, the Red Hills salamander (Phaeognathus hubrichti) (U.S. Fish and Wildlife Service, 1994a). An additional 15 amphibian species (12 salamanders and three frogs) were formerly considered federal Category 2 candidates (U.S. Fish and Wildlife Service, 1994b). Six of the ten states in the Southeast (Florida, Georgia, Mississippi, North Carolina, South Carolina, and Tennessee) have lists of state protected amphibian species. Twenty-six salamanders and six frogs are on these lists. The current list compiled by TNC and the Natural Heritage Program ranks 20 salamanders and two frogs as critically imperiled, imperiled, or rare and uncommon. There is considerable overlap between these lists, as many species appear on more than one list (see Table 1). Counting each species only once, the total number of imperiled amphibians equals 46, including 39 salamanders and seven frogs (see Table 2). The total number of native amphibian species known to occur in the Southeast is 126 (Conant and Collins, 1991). Therefore, the imperiled species represent 37 percent of the total amphibian fauna of the Southeast, with 45 percent of salamander and 18 percent of frog species being imperiled.

Amphibian Resource Management In The Southeast

Over the past several years, herpetologists have begun sharing their observations on declining amphibian populations with the general public (Blaustein and Wake, 1990; Vitt et al., 1990; Livermore, 1992; Yoffe, 1992). The dramatic population declines seen in some areas of the world have not been demonstrated in the Southeast. Granted, range-wide historical data are few and generally not comprehensive enough to provide an adequate baseline with which to compare current distributions. However, given the destruction, degradation, and fragmentation of imperiled amphibian habitats in the Southeast, there can be little doubt that the distribution of many species has probably contracted from historical levels (Vial and Saylor, 1993; Pechmann and Wilbur, 1994).

The management of amphibians as a resource to be protected, in the traditional sense of wildlife management, has only recently been addressed (Szaro et al., 1988). Wildlife management has historically targeted game and other species which have a perceived economic value (Schemnitz, 1980). Amphibians generally do not fall into this category, and thus they have typically been ignored. Additional reasons for the limited management activities focusing on amphibians include a lack of information on the regional fauna; scarce resource dollars for management which are often specifically targeted for birds, mammals, or fish (e. g., Pittman/Robertson funds to states); a perception that amphibians are common and do not require management; and a general attitude that amphibians are not as important a component of ecosystems as other vertebrates. Further neglect may be a consequence of difficulties associated with studying fauna which are inactive for large portions of the day or year (Scott and Seigel, 1992). Lastly, most herpetologists do not have a background in resource management, and they seldom have directly participated in the development of resource management strategies.

Although the implementation of amphibian resource management is in its infancy, resource managers are beginning to show an interest in amphibians. This interest is being supported by researchers who are providing information to assist in planning management strategies (Seehorn, 1982; Dodd and Charest, 1988; Gibbons, 1988; Szaro et al., 1988; Dodd, 1991; Scott and Seigel, 1992; Dodd, 1993; Bookhout, 1994; Dodd and LaClaire, 1995; deMaynadier and Hunter, 1995; Dupuis et al., 1995). This link between managers and scientists is being strengthened in several ways. For example, a manual for wildlife managers that addresses management of both amphibians and reptiles is being developed by the U.S. Forest Service in cooperation with The Nature Conservancy (M. Robertson, U.S. Forest Service, pers. comm.). Scientific assessment methods to reliably determine amphibian rarity or commonness have also recently been standardized and made generally available (Jones, 1986; Corn and Bury, 1990; Heyer et al., 1994). In spite of this progress, the incorporation of amphibians into resource management plans remains rare, and funds for amphibian management projects are usually difficult to obtain from resource management agencies.

Table 2. Summary of numbers of imperiled amphibians in the southeastern United States.
Amphibian Group
Federal Listed Species
Federal Species of Concern
State Listed Species
Species With TNC Global Rank1
Total Number2
Salamanders
1
12
26
20
39
Frogs
0
3
6
2
7
1 This category reflects ranking systems of The Nature Conservancy and Natural Heritage Data Center Network.
2 Total Number will be less than the sum of each row due to overlap in categories (see Table 1).

Although amphibians have generally been overlooked in the past, they have benefited when resource management strategies incidentally protected their habitats. In a few instances, amphibian populations and their habitats have been specifically targeted for protection, usually as a result of federal or state regulations.

Legislation And Regulation As Management Tools

Legal protection afforded by federal and state regulations has been the basis for amphibian resource management. Federal legislation which may afford protection for amphibian resources includes the Clean Water Act, the Endangered Species Act, the Fish and Wildlife Coordination Act, the National Environmental Policy Act, the National Forest Management Act, and the National Wild and Scenic Rivers Act. As mentioned above, six southeastern states have non-game regulations which provide protection for state-listed species, and all ten southeastern states require scientific or commercial permits to collect at least some species of non-game wildlife. These permits can be denied on the basis of species rarity. There are also local zoning and land-use regulations which may offer some protection for amphibian resources.

The implementation of most environmental legislation has been reactive rather than pro-active (see Karr, 1990; Doppelt et al., 1993). For example, the intent of the Endangered Species Act is to provide a means to conserve the ecosystems upon which endangered and threatened species depend and to provide a program for the conservation of those species. Implementation of the Endangered Species Act often results in protecting individuals of a particular species on a case by case basis, but conservation of the habitats and ecosystems in which these species live has rarely been accomplished (Clark et al., 1994). Some legislation, such as the Clean Water Act, can target habitat directly. In general, enforcement of such environmental regulations is difficult and often directed towards only the most blatant violators.

Each of the six acts of federal legislation outlined below are intended, at least in part, to protect the quality of the environment and, as a result, have applicability to amphibian protection and management. The Clean Water, the Fish and Wildlife Coordination, and the National Wild and Scenic Rivers acts, specifically address protection of water resources. The Endangered Species, the National Environmental Protection, and the National Forest Management acts, address protection of species, protection of the human environment, and protection of our national forests and their resources, respectively.

The Clean Water Act (CWA) provides for the restoration and maintenance of the chemical, physical, and biological integrity of the Nationís lakes, rivers, streams, and coastal waters. The CWA specifically regulates the discharge of any pollutant or the discharge of dredge or fill material into all waters of the United States including adjacent wetland ecosystems. Section 404 of the CWA defines a permit program to control dredging or filling activities. The U.S. Army Corps of Engineers (COE) has permit authority over waters of the United States (i.e., all waters affecting interstate and foreign commerce and translated by the U.S. Environmental Protection Agency [EPA] as any wetland habitat used by migratory birds). The EPA has an oversight role in promulgating guidelines for the permit program and has the authority to veto permits. The COE can issue General Permits for activities "similar in nature" and having minimal individual or cumulative adverse effects. One of these General Permits, Nationwide Permit 26, authorizes any dredge or fill activity in headwaters and isolated wetlands provided less than ten acres (approximately four ha) are impacted. In waters other than traditionally navigable waters, states may displace COE permitting (Blumm and Zaleha, 1989) and determine the size of wetlands to be regulated under dredge and fill permits. Many wetlands used for amphibian breeding are smaller than the size designated as jurisdictional by Nationwide Permit 26 or by state agencies.

The Fish and Wildlife Coordination Act (FWCA) was promulgated to allow development of water resources while conserving wildlife resources and environmental quality. The intent of the FWCA is to give wildlife conservation a coequal purpose or objective in federally funded or permitted water resource development proposals or projects. Federal agencies planning projects to develop water resources (e.g., the COE) must consult with the U.S. Fish and Wildlife Service and state game and fish agencies prior to seeking authorization for the action. The planning agency must give full consideration to measures recommended by these agencies to mitigate impacts to wildlife resources. This provides opportunity for amphibian management issues to become part of the decision-making review process. The agency carrying out the development project makes the final decision on wildlife mitigative measures, however.

The National Wild and Scenic Rivers Act (WSRA) created a system for designation and preservation of free-flowing rivers on private, state, or federal lands. A separate federal law is required for each Wild and Scenic River segment protected. Management areas for these congressionally designated rivers generally include adjacent corridors; however, the average acreage per river mile may not exceed 320 acres (about 130 ha) within the designated segment (Doppelt et al., 1993). Federal land management agencies must preserve designated river segments within their jurisdiction in free-flowing condition, protect water quality, and fulfill other national conservation purposes (e.g., protection of wildlife). The federal managing agency decides how best to protect the value of a river, but must consult with the EPA and state water pollution control authorities. Three rivers in the southern Appalachian Mountains have been designated Wild and Scenic Rivers under this Act. Implementation of the WSRA may benefit amphibians living in these watersheds. The three rivers are the Chattooga River in Georgia, North Carolina, and South Carolina, the Obed River in Tennessee, and the headwaters of the New River in North Carolina (Wallace et al., 1992).

The Endangered Species Act (ESA) specifically addresses the conservation of endangered species and the ecosystems on which they depend. Implementation of this Act is initiated by determining if a species meets the criteria to be designated an endangered or threatened species. This determination is made by reviewing a speciesí status based on the following five factors: the present or threatened destruction, modification, or curtailment of its habitat or range; the overutilization for commercial, recreational, scientific, or educational purposes; disease or predation factors; the inadequacy of existing regulatory mechanisms; and other natural or man-made factors affecting the speciesí continued existence. Once a species is placed on the Endangered Species List, the ESA provides a number of methods to protect it and its habitat. Recovery Plans are developed and implemented for listed species. Legal protection provided by the ESA is described under Section 7 (actions of or supported by government agencies shall not jeopardize endangered species), Section 9 ("take" of endangered species is prohibited), and Section 10 (an incidental take permit may be obtained if an approved Habitat Conservation Plan has been developed). Section 6 of the Act describes provisions for states to receive federal funds to perform research on candidate and listed species. Management issues can be addressed through use of these funds, but in most cases research is in response to a need for status surveys or recovery tasks outlined in Recovery Plans. All states in the Southeast have Section 6 agreements with the U. S. Fish and Wildlife Service which require the states to provide protection for federally listed species.

The National Environmental Policy Act (NEPA) provides guidelines for federal agencies to follow in the course of performing their various activities. The laws, regulations, and policies of the United States must be administered in accordance with the policies of NEPA. Specifically, federal agencies are required to consult with each other when planning actions that may significantly affect the quality of the human environment. Consultation must occur prior to approval of projects, and a statement of environmental impacts that are likely to result from the actions must be formulated. The federal agency with the lead on the action must prepare an Environmental Impact Statement with input from the public, state and local governments, and all federal agencies with legal jurisdiction or special expertise. The Act offers the opportunity for input from agencies with the lead for natural resource protection, such as the U.S. Fish and Wildlife Service, in the development of mitigative measures which can offset impacts of the proposed action. These measures may provide a direct benefit to imperiled amphibians.

The National Forest Management Act (NFMA) governs stewardship of national forests which is the responsibility of the U.S. Forest Service. The major focus of NFMA is on interdisciplinary land management planning, including procedures for public participation in the decision making process. Forest management plans must provide for multiple uses of national forest lands and must incorporate the values of watersheds and fish and wildlife into planning documents. Guidelines are provided for the following: obtaining inventory data on the various renewable resources in order to maintain the diversity of plant and animal communities; allowing timber harvest only where water bodies are protected from detrimental changes in water temperature, blockage of flow, and deposits of sediment; and, ensuring that harvesting methods are carried out in a manner consistent with the protection of soil, watersheds, and fish and wildlife. As a result of NFMA, amphibian status surveys and research projects are being funded to provide data for improving resource management plans for national forests.

Specific Management Programs Benefiting Imperiled Amphibians

The development of resource management programs for imperiled amphibians has been hindered by the limited data available on speciesí distributions and population sizes. In order to improve this situation, distribution and status surveys of some southeastern amphibians have been initiated by state non-game and heritage programs, The Nature Conservancy, the U.S. Fish and Wildlife Service, the U.S. Forest Service, the U.S. Department of Defense, and individual herpetologists. Funding for surveys has been provided by private landowners, state non-game programs, and by federal agencies. The Alabama Natural Heritage Program and the U.S. Fish and Wildlife Service have initiated the Alabama Herpetological Atlas Project to provide baseline data on reptiles and amphibians in Alabama. These efforts represent the beginnings of a region-wide database on amphibian distribution and abundance.

Specific management projects for imperiled amphibians are discussed below. Since resource management funds are not generally allocated solely for imperiled amphibians, total dollars spent directly for amphibian management could be provided for only a few of the projects discussed. Most of these management programs are ongoing and are being modified as results dictate. Of course, their success or failure cannot be judged until adequate long-term monitoring studies (at least a generation time for each species) have been completed.

Amphibian Management on Federal Lands

Resource management addressing amphibians has generally been tied to timber management on federal lands by the U.S. Forest Service and the Department of Defense. Timber management on these lands has most commonly focused on communities or ecosystems. In some cases, specific actions which benefit amphibians have been included. These include strategies to maintain moist environments and to protect wetland and stream habitats. Streamside management zones and timber harvests, limited to single tree selection, have been used to maintain forest canopy and soil moisture at a level more suitable to imperiled amphibians than would clearcut harvests. Many pond breeding amphibians require temporary wetlands for reproduction. Management actions used to protect these ponds have included the prescribed burning of temporary wetlands, the termination of fish stocking, the curtailment of the use of breeding ponds as fire breaks, and the creation of temporary ponds.

Studies have been undertaken on a number of national forests in different states to examine the effects of different harvest methods and successional impacts on amphibian and reptile populations. Additional studies have focused on surveys for imperiled amphibians on lands managed by the U.S. Forest Service and the Department of Defense. These studies have provided information that will be used in developing timber management plans.

Many federally owned lands in the Southeast have suitable habitat for three imperiled amphibians endemic to areas historically dominated by longleaf pine forests: the gopher frog (Rana capito), the striped newt (Notophthalmus perstriatus), and the flatwoods salamander (Ambystoma cingulatum). Managers of these federal lands have, or plan to, tailor their resource management plans to include measures to protect these amphibians.

A project has been undertaken in the Conecuh National Forest of Alabama to restore Nellie Pond, a breeding site of the dusky gopher frog (Rana capito sevosa). Due to the relatively recent introduction of predaceous fish to the pond by fishermen, tadpoles of the gopher frog, which historically bred in this ephemeral pond, were no longer surviving to metamorphosis. In 1992, U.S. Forest Service personnel drained the pond, removed as many fish as possible, and used rotenone to kill the remaining fish. The pond refilled naturally. Signs were erected to inform the public of the importance of maintaining the pond without a fish community. Public hearings were held to receive public input. Dusky gopher frog eggs were collected from a nearby pond and were raised in a laboratory until they were large tadpoles. They were then introduced into Nellie Pond, and during subsequent monitoring, metamorphic frogs were observed exiting the pond. Continued monitoring will be needed to determine the long-term success of this project. The project cost, approximately $11,000, was supported in part by a timber sale resulting from slash pine thinning of the surrounding habitat (R. Lint, U.S. Forest Service, pers. comm.). In addition to funding the project, the timber thinning improved the habitat for the resident upland-dwelling adult dusky gopher frogs.

Table 3. Imperiled amphibian fauna of the southern Appalachian Mountains. All = endemic, Part = part of range within southern Appalachian Mountains.
Taxon
All
Part
SALAMANDERS
Plethodontidae
Desmognathinae
Desmognathus aeneus, seepage salamander
 
X
D. santeetlah, Santeetlah dusky salamander
X
 
D. welteri, Black Mountain dusky salamander
 
X
Plethodontinae
Aneides aeneus, green salamander
 
X
Eurycea aquatica, dark-sided salamander
X
 
E. junaluska, Junaluska salamander
X
 
E. longicauda longicauda, longtail salamander
 
X
E. lucifuga, cave salamander
X
 
Gyrinophilus palleucus, Tennessee cave salamander
 
X
G. porphyriticus, spring salamander
 
X
Hemidactylium scutatum, four-toed salamander
 
X
Plethodon aureolus, Tellico salamander
 
X
P. dorsalis, zigzag salamander
 
X
P. petraeus, Pidgeon Mountain salamander
X
 
P. teyahalee, southern Appalachian salamander
X
 
P. websteri, Webster's salamander
 
X
P. wehrlei, Wehrle's salamander
 
X
P. welleri, Weller's salamander
X
 
P. yonahlossee, Yonahlossee salamander1
X
 
FROGS
Hylidae
Pseudacris brachyphona, mountain chorus frog
 
X
1 Listed in North Carolina as Plethodon longicrus, the crevice salamander.

The only breeding pond for the dusky gopher frog in Mississippi occurs in the DeSoto National Forest. Projects have been funded by the U.S. Forest Service and the U.S. Fish and Wildlife Service to study gopher frog life history and to improve the pond habitat for this species. Forest management plans since 1994 have incorporated the needs of the frog and will continue to be adapted as new data become available.

In Florida, status surveys have been conducted for the Florida gopher frog (Rana capito aesopus) and the striped newt in the Ocala National Forest (ONF) (Telford, 1993). In addition to these surveys, a study of the effects on amphibians of different timber cutting regimes around selected temporary ponds in the ONF has been initiated (K. Greenberg, U.S. Forest Service, pers. comm.). Results of these studies will be incorporated into resource management plans for the forest. Additional surveys have been funded by the U.S. Forest Service and the U.S. Fish and Wildlife Service to determine the distribution of the flatwoods salamander, formerly considered a Category 2 federal candidate species, in Floridaís Apalachicola National Forest (Palis, 1992, 1993, 1995a) and Osceola National Forest (Palis, 1992; D. Printiss, Florida Natural Areas Inventory, pers. comm.). These distribution data will be used to help formulate resource management plans for the salamander on both of these national forests.

The U.S. Department of Defense (DOD) has funded surveys for rare vertebrates, including the three aforementioned longleaf pine endemic amphibians, on a number of military bases in the Southeast. Surveys on Camp Blanding in Florida have been conducted by biologists with the Florida Natural Heritage Program. Management steps to enhance populations of the Florida gopher frog and striped newt were included in their summary report (Hipes and Jackson, 1994). A faunal survey for rare vertebrates on Fort Benning in Georgia was initiated in 1994 and will continue through 1997 (P. Laumeyer, U.S. Fish and Wildlife Service, pers. comm.). Prior to this study, a breeding site for the imperiled dusky gopher frog had been found on the base. For this reason, biologists conducting the surveys are especially interested in searching for additional localities for the gopher frog and other rare amphibians. Data collected during the survey will be used in future Fort Benning resource management plans. Biologists with The Nature Conservancy, funded by DOD, have surveyed Fort Stewart in Georgia for rare plants and animals including the Carolina gopher frog (Rana capito capito), the striped newt, and the flatwoods salamander (K. Lutz, The Nature Conservancy, pers. comm.). The distribution information generated by this study is being used to help develop a resource management plan (L. Swindell, DOD, pers. comm.). Additional studies on flatwoods salamander and striped newt life histories will continue through the year 2000 to determine how best to manage for these species on Fort Stewart (L. Swindell, DOD, pers. comm.).

A five-year Natural Resources Management Plan has been developed by the DOD for Eglin Air Force Base in Florida (Department of Defense, 1993). The goals of the plan are to protect plant communities and to restore ecosystem function and viability through careful land management. Eglin Air Force Base encompasses what may be the largest remaining, relatively intact longleaf pine/turkey oak forest in the United States. The presence of the federally listed red-cockaded woodpecker has focused planning on the restoration and maintenance of the sandhill community. The longleaf pine sandhills and flatwoods at Eglin Air Force Base are also habitat for the dusky gopher frog and the flatwoods salamander. Population surveys and identification of temporary pond breeding sites used by these two imperiled amphibians have been completed (Palis, 1995b). Habitat restoration, as described in the management plan, will benefit both species.

The U.S. Department of Energy (DOE) has funded long-term studies on reptiles and amphibians at the DOE Savannah River Site (SRS) in South Carolina. A considerable database has been developed for the site by researchers at the Savannah River Ecology Laboratory (SREL) (see Gibbons and Semlitsch, 1991). The U.S. Forest Service manages the forest on the SRS for timber production. Portions of the SRS have been set aside for protection, but at present there are no specific management goals for imperiled amphibians (B. Jarvis, U.S. Forest Service, pers. comm.). A project started at the site in the late 1980s provided potential benefits to imperiled amphibians through the creation of several temporary ponds as replacements for a pond that was destroyed. This attempt to replace the destroyed wetland with newly created ones of equal value was only partially successful (J. Pechmann, SREL, pers. comm.). The new ponds were dug near the periphery of the construction site and the location of the original wetland. Re-creating the original hydrologic conditions was problematic. Initially, the created ponds drained more frequently and remained dry for longer periods of time than the original temporary pond. Later, when liners were used to create an impervious bottom layer, the ponds became too permanent. As a result, species composition differed in the created ponds from the original wetland. Uncommon species, formerly present in the destroyed pond, were not recorded in the created ponds.

Projects on national forest lands in the southern Appalachian Mountains, which are habitat for a number of imperiled amphibians (see Table 3), are being carried out in the Daniel Boone National Forest in Kentucky, and the Nantahala and Pisgah national forests in North Carolina. In the Daniel Boone National Forest, traditional wildlife ponds have been created for many years for use by the resident deer population. U.S. Forest Service biologists, interested in managing for amphibians, modified the design of these ponds by varying their depth and shape, and by adding logs and branches to encourage colonization by amphibians. Eight of these newly created ponds were monitored during the first year (1994) after construction. Six species of frogs and two species of salamanders were found in the ponds, including an imperiled species, the mountain chorus frog (Pseudacris brachyphona) (J. MacGregor, U.S. Forest Service, pers. comm.).

On the Nantahala and Pisgah national forests, a study of the distribution and habitat of the Junaluska salamander (Eurycea junaluska) was completed in 1995. The results of this study will be used in the development of a conservation strategy for this imperiled species. Discussion of a multi-agency conservation agreement was initiated in May 1996 between the U.S. Forest Service, the U.S. Fish and Wildlife Service, the North Carolina Department of Transportation, and a regional power company (R. McClanahan, U.S. Forest Service, pers. comm.). Additionally, a database on the distribution of another imperiled species, the green salamander (Aneides aeneus), has been compiled for these two national forests. Using this database, timber operations are modified when needed, on a case by case basis, to ensure that green salamanders and their habitat are protected (R. McClanahan, U.S. Forest Service, pers. comm.).

The Ouachita Mountains of Arkansas and Oklahoma are also habitat for a number of imperiled amphibians. Two imperiled species, the Fourche Mountain salamander (Plethodon fourchensis) and the Caddo Mountain salamander (P. caddoensis), are endemic to Arkansas and virtually all of their habitat occurs within the Ouachita National Forest. Both species occur on steep wooded slopes and are vulnerable to reductions in soil moisture. The U.S. Fish and Wildlife Service, the U.S. Forest Service, and the Arkansas Game and Fish Commission have developed a memorandum of understanding for each species to benefit their conservation. Since 1993, these three agencies have been working together to determine the distribution of these salamanders and to monitor their populations. In addition, timber harvest and selective forest thinning have been restricted in streamside management zones and on slopes greater than 35 degrees to protect salamander habitat. The management of these salamanders has been incorporated into the larger, ecosystem-level Ouachita National Forest Management Plan.

Additional research in the Ouachita National Forest, part of the Ouachita Mountains Management Research Project, examines the distribution of amphibians and reptiles in the forest. Cooperators on this project, which was initiated in 1992 and continues through 1997, include the U.S. Forest Service, Weyerhaeuser Company, Oklahoma State University, the University of Arkansas, and the National Council of the Paper Industry for Air and Stream Improvement, Inc. (NCASI) (B. Wigley, NCASI, pers. comm.). Data on amphibian and reptile species abundance and community composition are being collected at both the forest-stand level and landscape-level in four watersheds with different histories of silviculture and intensity of timber management. A study of forest stands compares surveys on sites managed under three different regimes: single-tree selection (not clearcut), even-age management (clearcut, burn, replant), and eighty-year mature forest. Results of the surveys will be used to examine the effects of forest succession on the distribution of amphibians and reptiles (D. Crosswhite, Oklahoma State University, pers. comm.). A landscape-level study used satellite imagery to classify habitats from which different size study plots were chosen. The study plots are being censused to determine amphibian and reptile community composition. These data will be used to compare amphibian population diversity in different size patches and to determine if certain patches have higher value than others for amphibians. Geographic Information System (GIS) methods are being tested as a technique to make predictions about species composition on forest patches. The results of these studies will be used in preparing a management plan for the Ouachita National Forest in 1997 or 1998. This plan will encompass management for imperiled amphibians as well as other rare vertebrates (R. Perry, U.S. Forest Service, pers. comm.).

The U.S. Department of the Army, Corps of Engineers is planning a study at Haines Island Park in Alabama on the federally threatened Red Hills salamander (Phaeognathus hubrichti) (B. Peck, COE, pers. comm.). The study will focus on the development of a management plan to promote the recovery of the Red Hills salamander. An additional study at the site, initiated in 1995 and funded by the U.S. Fish and Wildlife Service, is testing the feasibility of using implanted passive integrated transponder (PIT) tags to monitor salamanders. This technique shows promise for use in mark and recapture studies and as a method to study burrow use by the animals. Data from this study will be valuable in determining management strategies for the Red Hills salamander.

Amphibian Management on Private and State Lands

While most amphibian management has been the result of forest management on public lands, some management has been directed at imperiled amphibians on private lands. Principles of conservation biology have rarely been used as guidance for private land-use planners. However, some private landowners are beginning to incorporate ecological principles into economic land-use decisions. OíConnell and Noss (1992) outlined ecological criteria for use in developing appropriate management plans and general methods for monitoring the effectiveness of those programs. Private land management plans that have accommodated both land-use objectives and the needs of rare species often use mitigation, restoration, and compensation to meet these goals. Projects directed at protection or enhancement of imperiled amphibian populations on private lands have been undertaken by the International Paper Timberlands Operating Company, Ltd. in Alabama; The Nature Conservancy in North Carolina; the National Council of the Paper Industry for Air and Stream Improvement, Inc. in Alabama, Florida, and Georgia; and a development company in Florida.

The International Paper Timberlands Operating Company, Ltd. (IPTOC) has developed a Habitat Conservation Plan (HCP) for the federally threatened Red Hills salamander in Conecuh and Monroe counties in Alabama (International Paper Timberlands Operating Company, Ltd., 1993). The plan was developed in cooperation with the U.S. Fish and Wildlife Service as part of an application for an incidental take permit under Section 10 of the ESA. The ESA defines the term "take" as meaning to harass, harm, pursue, hunt, shoot, wound, kill, trap, capture, collect, or to attempt to engage in any such conduct. Section 10 of the ESA allows issuance of permits if a taking is incidental to, and not the purpose of, the carrying out of an otherwise lawful activity. The underlying philosophy of an HCP is that resource management on private lands (e.g., silviculture) and endangered species management objectives can be integrated for the benefit of both. The cost for developing the IPTOC plan was between 50,000 and 75,000 dollars, and included costs for species surveys, plan preparation, and training of field personnel (J. McGlincy, IPTOC, pers. comm.).

The primary form of incidental take under the aforementioned Section 10 permit is habitat modification resulting from timber harvesting activities. To address this issue, results of habitat and population surveys were used to classify Red Hills salamander habitat as optimal, sub-optimal, or marginal. The HCP includes provisions to minimize take by only allowing forest management activities in marginal salamander habitat and to avoid and mitigate the impacts of take by establishing high-value habitat areas totaling 4,500 acres (about 1,821 ha) where no timber harvesting will occur. These habitats represent 92 percent of all occupied Red Hills salamander habitat on IPTOC lands. In forested buffer zones, adjacent to the areas where no harvest will occur, timber harvesting may occur, but a 50 percent canopy cover will be maintained. The plan includes methods for ongoing habitat and species monitoring and procedures for implementing contingency plans and amending the HCP. The permit allows incidental take for a 30-year period, with a comprehensive review conducted every ten years.

The North Carolina field office of The Nature Conservancy, in cooperation with the North Carolina Herpetological Society and the North Carolina State Museum of Natural Sciences (NCSMNS), has worked to improve breeding habitat for the Carolina gopher frog on properties they manage in the state. The imperiled Carolina gopher frog is listed as a species of special concern in North Carolina. The project involved altering the hydrology of one pond and it required creating two ponds on TNC properties, and creating one pond on North Carolina Department of Agriculture property (A. Braswell, NCSMNS, pers. comm.). The five-year project was directed by the NCSMNS and funded by the U.S. Fish and Wildlife Service at a cost of approximately 10,000 dollars.

The existing pond was converted from a permanent pond to a temporary pond by removing the resident fish and adjusting the contour to allow the pond to dry completely. The three created ponds were dug in areas of suitable habitat. They were inoculated with aquatic vegetation, a nutrient source, and gopher frog tadpoles from local populations. Metamorphic frogs were observed leaving two of the created ponds, but thus far, no adults have returned to the ponds to breed (A. Braswell, NCSMNS, pers. comm.).

Difficulties were encountered in this project when trying to establish ponds of a temporary nature. The converted fish pond remains too permanent and may require more modifications to make it shallower and to allow it to dry completely on a seasonal basis. One of the created ponds may be too ephemeral. Although no gopher frog breeding has been observed in any of the four ponds, the eastern tiger salamander (Ambystoma tigrinum tigrinum), listed as threatened in North Carolina, has bred in the converted pond. Many other amphibian species have also used the ponds for breeding.

The National Council of the Paper Industry for Air and Stream Improvement, Inc., a non-profit research organization associated with the forest products industry, is conducting several amphibian research projects which may provide benefits to southeastern imperiled amphibians. One of these projects involves studies on the effects of various timber practices on amphibians and wetlands at a site in Alachua County in Florida. Imperiled amphibians are not specifically targeted in this project; however, researchers hope the results of the study will provide a means to integrate the needs of sustainable industrial forestry with those of maintaining biodiversity (B. Wigley, NCASI, pers. comm.).

A second research project, funded by participating American Forest and Paper Association member timber companies and conducted by NCASI, targets three imperiled longleaf pine endemics, the gopher frog, the striped newt, and the flatwoods salamander, on lands owned or managed by the timber companies. Potential breeding sites for these three species are being surveyed within selected counties of Alabama, Georgia, and Florida. The three-year project was initiated in 1995. Once the surveys are completed, it is hoped that the distribution data from the study will be used by the timber companies to develop resource management plans that protect these three species.

In Lake and Sumter counties in Florida, the development plan for a 3,200 acre (about 1,295 ha) residential area, The Villages of Lake Sumter, included on-site protection for the Florida gopher frog. The protection was achieved through mitigation and compensation resulting from the Development of Regional Impact (DRI) review process required by the state of Florida for large scale development projects (R. McCann, Florida Game and Fresh Water Fish Commission, pers. comm.). Within the development, a 126.2 acre (about 51 ha) preserve was created to protect a number of vertebrate species including the gopher tortoise, the burrowing owl, and the southeastern kestrel. Part of the preserve (33.2 acres or about 13.4 ha), including a 0.25 acre (about 0.1 ha) sinkhole pond breeding site, has been set aside to protect the Florida gopher frog. Habitat restoration on the preserve, which was formerly part of a cattle ranch, occurred as development of the site progressed. This included exotic vegetation removal, mowing, and burning. A wildlife habitat management plan written for the preserve, as part of a conservation easement, contains restrictions on land-use in the preserve and habitat management requirements, including ones specifically addressing the gopher frog. The Florida Game and Fresh Water Fish Commission (FGFWFC) holds the conservation easement on the property which gives the FGFWFC interest in the title. The development company must adhere to the management plan as outlined in the easement. Management activities are intended to be financially supported in perpetuity through a homeownersí association fee. Parts of the management plan addressing the gopher frog include monitoring the water quality and hydrology of the gopher frog breeding pond, protecting the pond from residential and golf course drainage, monitoring frog activity at the breeding pond, managing the upland areas used by the gopher frog with prescribed burning, and providing information to the public on the frog and the importance of protecting its habitat (R. Ashton, Applied Technology and Management, Inc., pers. comm.).

Future Of Amphibian Management In The Southeast

In the future, successful management of amphibian resources in the Southeast will require a coordinated effort between land managers on both public and private lands. Fortunately, there is a growing awareness within federal and state agencies and private organizations about the importance of protecting our native amphibian fauna. The development of quality management plans will require the expertise of herpetologists, the skill of resource managers, the support of public citizens, and the creativity and cooperation of all parties involved.

Ideally, the development of most amphibian management plans should begin with an inventory and threat assessment of the amphibian community of concern. High priority species and habitats, as well as research needs should be identified during this inventory and threat assessment. Site-specific, single-species or ecosystem conservation strategies must be based on the threats to these species and habitats and should consider changes in species distributions and population status. Resource management plans should include criteria to determine effectiveness and measures of progress, as well as provisions to allow for modifications as new management information becomes available. Once a plan has been formulated, funding should be provided at a level sufficient to fully implement the management plan and to complete long-term monitoring studies.

Determining the distribution of imperiled amphibians and the threats they face can be difficult and time consuming. State non-game and heritage programs, The Nature Conservancy, the U.S. Fish and Wildlife Service, and interested herpetologists have begun surveys to determine the distribution and status of many imperiled amphibians in the Southeast. Additional data on amphibian status and distribution can be found by searching museum records and by gathering anecdotal information from experienced herpetologists. The Declining Amphibian Populations Task Force of the International Union for the Conservation of Nature/Species Survival Commission (IUCN/SSC) has developed a database called FROGLOG to summarize available information. Data from all of these sources have been used to prioritize species and communities that are imperiled (Natural Heritage Data Center Network, 1993). More rigorous methodologies, such as that developed by Millsap et al. (1990) in Florida, are needed for assessing priorities for research and monitoring based on species imperilment.

The difficulty in assessing priorities becomes apparent when looking at the literature published on the status of amphibian populations in the Southeast. There is a growing body of work documenting population declines on sites where habitats have been degraded or destroyed (Vickers et al., 1985; Enge and Marion, 1986; Ash, 1988; Dodd, 1991; Raymond and Hardy, 1991; Petranka et al., 1993; Phelps and Lancia, 1995; Means et al., 1996). The long-term effects of these declines to the species involved are unclear (Ash and Bruce, 1994). Clearly, when the habitat of a given population is destroyed, that population has gone or will shortly go extinct. Many species may be unable to recolonize areas after local extinctions, especially when unsuitable habitat exists between the extinct population and extant populations. In cases where the habitat is degraded but population stability is documented, the stability may simply reflect the persistence of long-lived individuals (Blaustein et al., 1994). On the other hand, several long-term studies have demonstrated that wide fluctuations in population numbers may be characteristic of amphibians (Pechmann et al., 1991; Hairston and Wiley, 1993). Also, many amphibians can survive prolonged drought or lack of food, and they may persist for long periods if the essential aspects of their habitat can be protected (Scott and Seigel, 1992). Long-term population studies, for at least one complete generation turnover of the monitored population, will be needed to determine the significance of declines and to critically assess the imperilment of many southeastern amphibians (Franklin, 1989; Blaustein et al., 1994). In the case of some amphibians, these studies may need to be extended to 15 or more years.

Other research needs will become evident after inventories and threat assessments are completed. Herpetologists from the public and private sectors need to become actively involved in studies that involve resource management of imperiled amphibians. Data are needed for use in the development of population and habitat viability models. For example, habitat requirements (both adult and larval), spatial use, and population structure are relatively unknown for many species. Research is also needed on the taxonomy and ecology of imperiled amphibians. Groups such as the plethodontids, Necturus spp., Pseudobranchus spp., and the ranid complexes may contain more species than presently described. Virtually nothing is known of the genetic diversity of rare amphibian populations (Stiven and Bruce, 1988). Without basic information about a speciesí biology, conservation programs will be based on assumptions that may not ensure the protection of amphibian populations (see Scott and Seigel, 1992; Underwood, 1995). The best available scientific information is needed to develop sound conservation strategies. This information should be continually updated to identify and investigate new management techniques and to critically assess the value of current conservation activities.

Species with specialized habitats or very limited distributions, such as the Red Hills salamander or the Fourche and Caddo mountain salamanders discussed above, can be protected by developing site specific management plans with a narrow focus. Other wide-ranging species may occur in multiple states and will require a management plan directed at ecosystem or watershed protection. The following discussion outlines management plan development for three areas which could improve conservation of a large portion (30 of 46 species) of the imperiled amphibians in the Southeast, namely, the southern Appalachian Mountains, the Mobile River Basin, and the longleaf pine forest ecosystem. A watershed management approach can be used to develop a conservation plan for both the southern Appalachian Mountains and the Mobile River Basin. A plan developed for the forests of the longleaf pine ecosystem will involve land-use management across watersheds, as well as wetlands management within geographically separated forests.

Southern Appalachian Mountains

The southern Appalachian region encompasses portions of Alabama, Georgia, Kentucky, North and South Carolina, and Tennessee. The imperiled amphibian fauna in this region, within the Blue Ridge and Ridge and Valley physiographic provinces, includes one frog and 19 salamanders (see Table 3). The frog is the mountain chorus frog and the salamanders are species within six genera of the family Plethodontidae. Eight of the salamanders are endemic to the southern Appalachian Mountains. These 20 amphibians inhabit areas adjacent to or in association with high-gradient streams. The streams are the headwaters of many major river systems, and the water originating from these watersheds is an important resource for major metropolitan areas (Wallace et al., 1992). Since rivers are open, directional systems, protection of any segment requires protection of the entire upstream network along with the surrounding landscape (Sedell et al., 1990). Within the southern Appalachian Mountains, watershed impacts occur as a result of forestry and surface mining activities, and industrial, agricultural, and municipal waste-water inputs.

A management plan for imperiled amphibians in watersheds throughout the Southern Appalachian Mountains should begin with a forest management plan. Aquatic amphibians and terrestrial species with aquatic larval stages are sensitive to degraded stream and wetland quality. Completely terrestrial southern Appalachian salamanders are sensitive to modifications in prevailing temperature, humidity, or soil moisture because adults lack lungs and gas exchange must occur through moist skin (Duellman and Trueb, 1986). Clearcutting and large-scale canopy removal can have negative effects on the quality of both aquatic and terrestrial habitats. Vegetation removal increases stream flow and water temperatures and reduces the input of coarse woody debris and other organic matter after cutting. Reductions in the quality of aquatic habitats can also occur through sedimentation. Loss of soil stability and sedimentation results from logging and associated practices such as road building (Anderson et al., 1976). In addition, where the forest has been removed, streams are decoupled from self-regulatory mechanisms within the watershed and they become more hydrologically unstable (Sedell et al., 1990). The quality of terrestrial habitats is reduced when canopy removal eliminates shading and increases temperatures and moisture loss at the soil surface.

Habitat destruction and degradation resulting from timbering operations may create problems for long-term survival of imperiled amphibians (Kramer et al., 1993; Petranka et al., 1993). Several studies in the southern Appalachian Mountains have demonstrated declines in salamander numbers after clearcutting (Ash, 1988; Petranka et al., 1993; Ash, 1994). Petranka et al. (1993) found that numbers of salamanders on study plots in the Pisgah National Forest in North Carolina were five times higher in mature forest stands than in clearcuts. Species richness also was higher in the mature stands. Clearcuts contained, on average, about 50 percent of the species found in mature forest sites. Ash (1988, 1994) studied a site in Nantahala National Forest near Highlands, North Carolina, for over 15 years. Salamanders were surveyed on two clearcut and two control plots. Numbers of Jordanís salamanders (Plethodon jordani) collected on clearcut plots ranged from 40 percent of the number on forested plots during the first summer after cutting to zero the fourth summer. By the sixth year after cutting, salamanders had returned to clearcut plots and by the fifteenth year, salamander numbers exceeded 50 percent of those found on forested plots.

Additional threats to imperiled amphibians can occur through impacts to the watershed resulting from surface mining and associated acid mine drainage, industrial and municipal wastes, and acid precipitation (Wallace et al., 1992). Numerous conflicts exist between commercial and other economic interests, private citizens, and the federal government over land and stream use in the southern Appalachian Mountains. Watershed protection in this sensitive area will require compromise and cooperation between these groups. There are strong, vested economic considerations in continuing practices that contribute to environmental degradation. In many cases, adequate knowledge exists to sharply curtail much of the stream degradation; however, it has been difficult to transfer technology from researchers to those engaged in forestry, agriculture, and development (Wallace et al., 1992). Public education is needed to provide alternatives to harmful practices and to demonstrate the long-term benefits of quality water resources. Public involvement at the local level is imperative. Cooperation is needed between the private sector and government agencies to share responsibility for both water quality degradation and improvement. Adequate laws must be enacted to protect water quality. In addition to regulation, government could support water quality improvement practices undertaken by the private sector through tax breaks or other economic incentives.

To protect imperiled amphibians in the Blue Ridge and Ridge and Valley physiographic provinces of the southern Appalachian Mountains, a region-wide conservation plan is needed. Core areas within the region, and connecting linkages or corridors of habitat for exchange of individuals and their genes, should be defined for each species (Mitchell, 1994; Dupuis et al., 1995). Streamside management zones may act as refugia and centers of dispersal on sites where timber has been harvested (Foley, 1994). Recommendations to preserve cool, moist habitats within harvested areas include maintaining logs and snags as moist microhabitats and retaining some understory as a source of shade (Dupuis et al., 1995). The core areas should be protected from further degradation by the maintenance of natural riparian and hardwood forest areas and reduction of fragmentation, isolation, and edge effects (Mitchell, 1994). A plan of this scope will also require public education on the importance of conservation efforts to protect the habitat and its imperiled species.

The U.S. Fish and Wildlife Service (1995) has completed an ecosystem management plan of this scope for the Southern Appalachian Ecosystem (SAE). Many of the components described above have been incorporated into the plan. The purpose of the plan is to outline goals, objectives, and strategies to protect and restore U.S. Fish and Wildlife Service (USFWS) trust resources and ecological integrity within the SAE. A number of initiatives are described in the plan which, if implemented, would benefit imperiled amphibians. These include strategies for projects on aquatic systems, riparian restoration, declining amphibian populations, education and outreach, and wetland restoration. The USFWS is just one of many partners whom share responsibility for ecosystem health. Additional partners include other federal agencies (e.g., U.S. Forest Service, U.S. National Park Service), state and local agencies, communities, organizations, and corporate and other private landowners. The four major goals and objectives in the SAE plan are as follows:

  1. Protect, restore, and enhance habitats and essential processes necessary to maintain healthy biological diversity in SAE. The objective is to reverse the decline of communities and species groups most "at risk" through an inventory and assessment of threats, then initiate the development and implementation of protection, management, and monitoring strategies for high priority species and habitats.
  2. Promote and support compatible and sustainable uses of ecosystem resources. The objective is to reduce the overuse and/or exploitation of biological and cultural resources and emphasize cooperation through partnerships to develop strategies to increase compliance with federal and state regulations.
  3. Increase public knowledge and support for ecosystem resources and their management. The objective is to promote stewardship of ecosystem resources, emphasizing cooperation built via partnerships.
  4. Increase the coordination and cooperation among agencies and organizations. The objective is to enhance the effective and efficient management of our shared natural resources.

Mobile River Basin

The Mobile Basin includes the Mobile, Tombigbee-Black Warrior, and Alabama-Coosa-Tallapossa river systems. Total drainage area prior to the construction of the Tennessee Tombigbee Waterway included more than half of the state of Alabama and portions of Georgia, Mississippi, and Tennessee (Livingston, 1992). Two imperiled amphibians occur in this drainage, the Black Warrior waterdog (Necturus sp.), and the one-toed amphiuma (Amphiuma pholeter). Water quality and the habitat of these two aquatic species have been degraded in the Mobile River Basin due to impoundment; channelization; dredging; coal, sand, and gravel mining; industrial and municipal discharge; and nonpoint discharge and run-off. All streams surveyed by Bailey (1992, 1995) for the waterdog, which occurs in the Black Warrior River system, showed some degree of degradation from sedimentation, and many appeared to be biologically depauperate. Bailey (1995) felt that surface mining was probably the greatest threat to the integrity of waterdog habitat, and that prospects for the long-term survival of the species were poor unless conditions in the watershed improved.

The Mobile River Basin ecosystem has been severely degraded. As a result, at least 18 species of mussels and 32 species of aquatic snails are presumed to have gone extinct, mostly within the past few decades. The Jackson, Mississippi field office of the U.S. Fish and Wildlife Service has initiated development of a recovery plan draft to address water quality improvement measures for the benefit of all federally listed aquatic flora and fauna in the Mobile River Basin. There are currently 32 aquatic animal and plant species in the basin that are listed under the ESA (U.S. Fish and Wildlife Service, 1994a). The primary objective of the plan under development is to protect the Basinís aquatic flora and fauna by achieving ecosystem stabilization. Actions presented in the draft plan to achieve this objective include the following:

  1. Protection of habitat integrity and water quality by full and appropriate implementation of federal and state regulations.
  2. Development and implementation of Best Management Practices and Water Quality Plans for construction, agriculture, urban, and suburban activities that affect aquatic ecosystems.
  3. Development and implementation of Watershed Management Plans that address problems specific to watersheds occupied by listed species.
  4. Development, funding, and implementation of programs that educate and directly involve the general public in ecosystem recovery.
  5. Basic research on life history, ecology, anatomy, taxonomy, contaminant sensitivities, and propagation of listed and candidate species.
  6. Reintroduction of listed and endemic species into restored habitats, as appropriate.
  7. Monitoring of listed species populations.
  8. Coordination of ecosystem and species recovery efforts.

Many of the needs and objectives of the Mobile River Basin Draft Recovery Plan are similar to those previously discussed for the southern Appalachian Mountains, including stressing the need for public education and cooperation between government agencies and the private sector. Improvements in water quality have been made in the Mobile River Basin, in part due to government regulation. These regulations alone, however, canít provide adequate protection to the imperiled species of the Mobile River Basin. It has become obvious that the public will have to become directly involved if the flora and fauna of the Basin, including its imperiled amphibians, are to be protected. Government agencies can provide support through conservation agreements, on-site guidance and assistance, and funds for stream protection and restoration. Water quality problems can be addressed through federal government funded research and the development of alternative practices.

In addition to this cooperation, both private and public entities need to formulate strategies to improve water quality through local actions. Individuals and organizations will need to enlist support for programs at the local level and to encourage individual- and community-based responsibility for the protection of water resources. Organizations already in existence, such as The Nature Conservancy, Alabama Natural Heritage Program, River Watch, Adopt-a-Stream, and others, could take the lead in encouraging public involvement. Government sponsored projects such as the construction of impoundments, channelization, and dredging should require assessments of need and alternatives, and economic analyses including assessments of costs associated with environmental impacts. Federal agencies such as the COE and the EPA need to fulfill their obligations to maintain and improve water quality within the Basin.

Conservation and management strategies focused on imperiled amphibians should prioritize streams within the Mobile River Basin based on occurrences of rare fauna and potential for restoration. Drainage Management Plans could then be designed to implement stream restoration and species specific management. State and federal regulatory agencies should be encouraged to enforce current water quality and mining regulations within these drainages. Through a cooperative effort between the public and private sector, Best Management Practices should be developed to address nonpoint discharge and run-off within an ecosystem context.

Longleaf Pine Forest Ecosystem

The longleaf pine forest ecosystem was once the predominant vegetative community of the southeastern Coastal Plain and stretched from Virginia to Texas (Means, 1988). Longleaf pine communities in pre-settlement times covered 60-80 million acres (roughly 24.3-32.4 million ha), over 60 percent of the upland area of the coastal plain (Croker, 1979; Ware et al., 1993). Today these communities cover less than 2 percent of their original distribution (Ware et al., 1993). The original longleaf pine stands ranged across a wide spectrum of environmental conditions from wet, poorly drained flatwoods to well-drained high pine or dry sandhills (Boyce and Martin, 1993). Urban development and conversion to agriculture have eliminated large portions of this ecosystem. Wetlands, integrated within the longleaf pine ecosystem, have been lost through ditching, draining, filling, and lowering of the water table. Intensive forestry has changed the once open longleaf pine stands to dense loblolly and slash pine plantations and second-growth hardwoods. Most of the longleaf pine ecosystem that remains is second-growth and degraded by logging, turpentining, grazing, intensive site preparation, and fire suppression (Noss, 1988).

The gopher frog, the striped newt, the flatwoods salamander, and two subspecies of dwarf siren (Pseudobranchus striatus) are imperiled amphibians endemic to longleaf pine forests. An additional five imperiled amphibians, three salamander species and two frog species, would also benefit from protection of the longleaf pine habitat (Table 4).

Habitat changes in the longleaf pine ecosystem could have significant impacts on the distribution and abundance of imperiled amphibians. Grant et al. (1994) compared amphibian communities on even-aged pine plantations in South Carolina in four age-classes (one, three, eight, and 26 years). First-year pine plantations had the lowest species richness and the 26-year-old plantations had the highest. Enge and Marion (1986) examined the effects of clearcutting and site preparation on amphibian community dynamics in pine flatwoods forest in Florida. Clearcutting decreased reproductive success, reducing species abundance by a factor of ten. Means and Moler (1979) studied the imperiled Pine Barrens treefrog (Hyla andersonii) in seepage bogs in the Florida panhandle. They found that the seepage habitat could be destroyed by woody shrub succession resulting from fire suppression. Another study in Florida implicated the conversion of longleaf pine savannas to bedded slash pine plantations in the decline of a local flatwoods salamander population (Means et al., 1996). In a few cases, however, forestry practices such as clearcutting may mimic some natural processes. Campbell and Christman (1982) suggested that these techniques may substitute for the hot, lightning-ignited crown fires to which sand pine stands are adapted. Researchers studying amphibian populations within the longleaf pine ecosystem need to become familiar with current and past forest management practices (site preparation, regeneration method, rotation time, etc.) in different longleaf pine habitats (deMaynadier and Hunter, 1995). Studies can then be conducted to isolate and evaluate the impacts of various practices on imperiled amphibians and to provide information to land managers for use in development of resource management plans.

At present, imperiled amphibians are benefiting from protection afforded other species with which they share their terrestrial habitat (e.g., the federally threatened gopher tortoise and eastern indigo snake, and the endangered red-cockaded woodpecker). Maintenance of longleaf pine habitat for amphibians will also require wetlands conservation. The imperiled dwarf sirens are totally aquatic. The other imperiled longleaf pine species, except for the pine barrens tree frog, require temporary ponds in which to breed. The ponds required by these species are typically devoid of large predaceous fish and dry completely on a cyclic basis, generally during the summer months (Moler and Franz, 1987; LaClaire and Franz, 1991). These temporary ponds are very sensitive to changes in hydrology (LaClaire, 1992). In order to protect these sites, the practice of using the pond basins as recipient sites for road run-off or for slash disposal during timbering operations should be terminated. Bedding, ditching, and harvesting of hardwoods in pond basins should be curtailed whenever possible to maintain pond function (Vickers et al., 1985; Dodd and LaClaire, 1995). The hydrology of the area surrounding these ponds should not be altered, or the capacity of the basin to hold water may be compromised. In addition, fires should be allowed to burn completely across dry pond basins in order to maintain the wetland plant community necessary for nutrient cycling and amphibian egg attachment and larval development (LaClaire, 1995).

Most of the remaining acres of longleaf pine habitat are on private lands and are rapidly being converted to slash and loblolly pine or developed for residential or industrial use (Means and Grow, 1985). Efforts are being made by the U.S. Fish and Wildlife Service to assist the private timber industry in the formation of Habitat Conservation Plans which will allow for the needs of industrial forestry and for conservation of endangered species on industry lands. Public education programs are also underway in several states to demonstrate the benefits of planting and managing for longleaf pine to the small private landowner.

Table 4. Imperiled amphibian fauna of the longleaf pine ecosystem. All = endemic, Part = part of range within longleaf pine ecosystem.
Taxon
All
Part
SALAMANDERS
Ambystomatidae
Ambystoma cingulatum, flatwoods salamander
X
 
A. talpoideum, mole salamander
 
X
A. tigrinum tigrinum, tiger salamander
 
X
Salmandridae
Notophthalmus perstriatus, striped newt
X
 
Sirenidae
Pseudobranchus striatus spheniscus, slender dwarf siren
X
 
P. s. striatus, broad-striped dwarf siren
X
 
FROGS
Hylidae
Hyla andersonii, Pine Barrens treefrog
 
X
H. gratiosa, barking treefrog
 
X
Ranidae
Rana capito, gopher frog
X
 

Primary obligation for protection of longleaf pine communities in public ownership rests with the U.S. Forest Service and the U.S. Department of Defense. The national forests of the Southeast contain about 700,000 acres (some 283,290 ha) of longleaf pine, representing one percent of the original longleaf pine community (Means and Grow, 1985). The DOD also has considerable acreage of longleaf pine habitat. For example, Eglin Air Force Base has 400,641 forested acres (about 162,140 ha) of which 81 percent are, or were historically, longleaf pine dominated sandhill communities (Department of Defense, 1993). The U.S. Forest Service and the DOD have re-oriented resource management programs toward an ecosystem-based approach.

To benefit imperiled amphibians, a longleaf pine ecosystem plan should include strategies for the following: re-establishing longleaf pine on appropriate sites; single tree selection timber harvesting to retain soil moisture; reducing soil disturbance during site preparation; maintaining the existing hydrology of temporary pond breeding sites; protecting streamside management zones; retaining downed logs, stumps, and snags as microhabitats; and using summer controlled burning (Means and Grow, 1985; Means, 1988; LaClaire, 1992; Dodd and LaClaire, 1995; Dupuis et al., 1995).

A region-wide plan is needed to protect the remaining longleaf pine ecosystem. A recent meeting hosted in 1994 by the U.S. Forest Service brought together a group of resource managers and researchers working in longleaf pine communities. This meeting highlighted the importance of a group effort to address protection and management of the remaining longleaf pine ecosystem. Development and coordination of a plan could be organized by working groups in each state. The working groups could then network with The Nature Conservancy, the U.S. Forest Service, the U.S. Fish and Wildlife Service, state heritage programs, other state and federal agencies, and private timber companies which have an interest in the status of longleaf pine communities and rare and endangered species.

Conclusions

There are 46 species of imperiled amphibians in the southeastern United States. Historically, resource management on public and private lands has not targeted amphibians, and their protection has been incidental to management for other resources. Recently, however, more emphasis has been placed on managing habitats and ecosystems which include imperiled amphibians.

There are many state and federal agencies and private conservation organizations that need to work together to implement resource management plans to conserve imperiled amphibians. Amphibian management in the Southeast will require status assessments, research, ecosystem management strategies (including long-term monitoring plans), and public participation to protect imperiled southeastern amphibians. Public involvement is crucial. Without the commitment of private landowners to conservation, amphibian management efforts can realize only minimal success. Legislation and regulations, no matter how worthwhile, will be effective only when the public perceives the good associated with them.

Most land available to imperiled amphibians in the Southeast is in private ownership. For example, timber companies own millions of acres that are often the only undeveloped sites large enough to maintain viable populations of amphibians and other wildlife. Increased efforts are needed to encourage conservation through mutual cooperation between the public and private sectors. As federal agencies, such as the U.S. Forest Service, the U.S. Fish and Wildlife Service, and others, shift from resource management to ecosystem management, rich opportunities for effective partnerships should develop. If these partnerships are nourished, they will provide tremendous opportunities to grow broad-based conservation support for imperiled amphibians as well as other plants and animals.

References

Anderson, H. W., M. D. Hoover, and K. G. Reinhart. 1976. Forests and water: Effects of forest management on floods, sedimentation and water supply. General Technical Report PSW-GTR-18, Pacific Southwest Forest and Range Experiment Station, Forest Service, U.S. Department of Agriculture, Berkeley, CA, 118 p.
Ash, A. N. 1988. Disappearance of salamanders from clearcut plots. Journal of the Elisha Mitchell Science Society 104:116-122.
Ash, A. N. 1994. Disappearance and return of salamanders to clearcut plots in the southern Blue Ridge Mountains. In Proceedings of the Fifth Annual Southern Appalachian Man and the Biosphere Conference. (Abstracts). Hendersonville, NC, p. 13.
Ash, A. N., and R. C. Bruce. 1994. Impacts of timber harvesting on salamanders. Conservation Biology 8:300-301.
Bailey, M. A. 1992. Black Warrior waterdog status survey: Final report 1991-92. Unpublished Report submitted to the U.S. Fish and Wildlife Service, Atlanta, GA, 27 p.
Bailey, M. A. 1995. Black Warrior waterdog survey, 1994-95. Unpublished Report submitted to the U.S. Fish and Wildlife Service, Atlanta, GA, 27 p.
Blaustein, A. R., and D. B. Wake. 1990. Declining amphibian populations: A global phenomenon? Trends in Ecology and Evolution 5:203-204.
Blaustein, A. R., D. B. Wake, and W. P. Sousa. 1994. Amphibian declines: Judging stability, persistence, and susceptibility of populations to local and global extinctions. Conservation Biology 8:60-71.
Blumm, M. C., and D. B. Zaleha. 1989. Federal wetlands protection under the Clean Water Act: Regulatory ambivalence, intergovernmental tension, and a call for reform. University of Colorado Law Review 60:695-772.
Bookhout, T. A. (ed.). 1994. Research and Management Techniques for Wildlife and Habitat, 5th Edition. The Wildlife Society, Bethesda, MD.
Boyce, S. G., and W. H. Martin. 1993. The future of the terrestrial communities of the southeastern United States. In Biodiversity of the Southeastern United States: Upland Terrestrial Communities. W. H. Martin, S. G. Boyce, and A. D. Echternacht (eds.). John Wiley and Sons, Inc., New York, NY, p. 339-366.
Campbell, H. W., and S. P. Christman. 1982. The herpetological components of Florida sandhill and sand pine scrub association. In Herpetological Communities. N. J. Scott, Jr. (ed.). Wildlife Research Report 13, Fish and Wildlife Service, U.S. Department of the Interior, Washington, D.C., p. 163-171.
Clark, T. W., R. P. Reading, and A. L. Clarke. 1994. Endangered Species Recovery: Finding the Lessons, Improving the Process. Island Press, Washington, D.C.
Conant, R., and J. T. Collins. 1991. A Field Guide to Reptiles and Amphibians of Eastern and Central North America. Houghton Mifflin Company, Boston, MA.
Corn, P. S., and R. B. Bury. 1990. Sampling methods for terrestrial amphibians and reptiles. General Technical Report PNW-GTR-256, Forest Service, U.S. Department of Agriculture, Portland, OR, 34 p.
Croker, T. C., Jr. 1979. The longleaf pine story. Journal of Forestry History 1979 (January):32-43.
deMaynadier, P. G., and M. L. Hunter. 1995. The relationship between forest management and amphibian ecology: A review of the North American literature. Environmental Review 3:230-261.
Department of Defense. 1993. Natural resources management plan, Eglin Air Force Base. U.S. Department of the Air Force, Eglin Air Force Base, FL, 235 p.
Dodd, C. K., Jr. 1991. The status of the Red Hills salamander Phaeognathus hubrichti, Alabama, USA, 1976-1988. Biological Conservation 55:57-75.
Dodd, C. K., Jr. 1993. Strategies for snake conservation. In Snakes: Ecology and Behavior. R. A. Seigel, and J. T. Collins (eds.). McGraw-Hill, Inc., New York, NY, p. 363-393.
Dodd, C. K., Jr., and B. G. Charest. 1988. The herpetological community of temporary ponds in north Florida sandhills: Species composition, temporal use, and management implications. In Management of Amphibians, Reptiles, and Small Mammals in North America. R. E. Szaro, K. E. Severson, and D. R. Patton (eds.). General Technical Report RM-166, Forest Service, U.S. Department of Agriculture, Fort Collins, CO, p. 87-97.
Dodd, C. K., Jr., and L. V. LaClaire. 1995. Biogeography and status of the striped newt (Notophthalmus perstriatus) in Georgia, USA. Herpetological Natural History 3:37-46.
Doppelt, B., M. Scurlock, C. Frissell, and J. Karr. 1993. Entering the Watershed. Island Press, Washington, D.C.
Duellman, W. E., and L. Trueb. 1986. Biology of Amphibians. McGraw-Hill Book Company, New York, NY.
Dupuis, L. A., J. J. M. Smith, and F. Bunnell. 1995. Relation of terrestrial-breeding amphibian abundance to tree-stand age. Conservation Biology 9:645-653.
Enge, K. M., and W. R. Marion. 1986. Effects of clearcutting and site preparation on herpetofauna of a north Florida flatwoods. Forest Ecology Management 14:177-192.
Foley, D. H. 1994. Short-term response of herpetofauna to timber harvesting in conjunction with streamside management zones in seasonally-flooded bottomland-hardwood forests of southeast Texas. M.S. Thesis, Texas A&M University, College Station, TX.
Franklin, J. E. 1989. Importance and justification of long-term studies in ecology. In Long-term Studies in Ecology. G. E. Likens (ed.). Springer Publishing Company, New York, NY, p. 3-19.
Gibbons, J. W. 1988. The management of amphibians, reptiles and small mammals in North America: The need for an environmental attitude adjustment. In Management of Amphibians, Reptiles, and Small Mammals in North America. R. C. Szaro, K. E. Severson, and D. R. Patton (eds.). General Technical Report RM-166, Forest Service, U.S. Department of Agriculture, Fort Collins, CO, p. 4-10.
Gibbons, J. W., and R. D. Semlitsch. 1991. Guide to the Reptiles and Amphibians of the Savannah River Site. The University of Georgia Press, Athens, GA.
Grant, B. W., Brown, K. L., Ferguson, G. W., and J. W. Gibbons. 1994. Changes in amphibian biodiversity associated with 25 years of pine forest regeneration: Implications for biodiversity management. In Biological Diversity: Problems and Challenges. S. K. Majumdar, F. J. Brenner, J. E. Lovich, J. F. Schalles, and E. W. Miller (eds.). The Pennsylvania Academy of Science, Philadelphia, PA, p. 355-367.
Hairston, M. G., Sr., and R. H. Wiley. 1993. No decline in salamander (Amphibia: Caudata) populations: A twenty-year study in the southern Appalachians. Brimleyana 18:59-64.
Heyer, W. R., M. A. Donnelly, R. W. McDiarmid, L.-A. C. Hayek, and M. S. Foster. 1994. Measuring and Monitoring Biological Diversity: Standard Methods for Amphibians. Smithsonian Institution Press, Washington, D.C.
Hipes, D., and D. Jackson. 1994. Rare vertebrate survey of Camp Blanding Training Site. Florida Natural Areas Inventory and The Nature Conservancy, Tallahassee, FL, 66 p.
International Paper Timberlands Operating Company, Ltd. 1993. Red Hills salamander habitat conservation plan. U.S. Fish and Wildlife Service, Jackson, MS, 32 p.
Jones, K. B. 1986. Amphibians and reptiles. In Inventory and Monitoring of Wildlife Habitat. A. Y. Cooperrider, R. J. Boyd, and H. R. Stuart (eds.). Bureau of Land Management Service Center, U.S. Department of the Interior, Denver, CO, p. 267-290.
Karr, J. R. 1990. Biological integrity and the goal of environmental legislation: Lessons for conservation biology. Conservation Biology 4:244-250.
Kramer, P., N. Reichenbach, M. Hayslett, and P. Sattler. 1993. Population dynamics and conservation of the Peaks of Otter salamander, Plethodon hubrichti. Journal of Herpetology 27:431-435.
LaClaire, L. V. 1992. Ecology of temporary ponds in north-central Florida. M.S. Thesis, University of Florida, Gainesville, FL.
LaClaire, L. V. 1995. Vegetation of selected upland temporary ponds in north and north-central Florida. Bulletin of the Florida Museum of Natural History 38:69-96.
LaClaire, L. V., and R. Franz. 1991. Importance of isolated wetlands in upland landscapes. In Proceedings of the Second Annual Meeting. M. Kelly (ed.). Florida Lake Management Society, Winter Haven, FL, p. 9-15.
Livermore, B. 1992. Amphibian alarm: Just where have all the frogs gone? Smithsonian Magazine 1992(October):113-120.
Livingston, R. J. 1992. Medium-sized rivers of the gulf coastal plain. In Biodiversity of the Southeastern United States: Aquatic Communities. C. T. Hackney, S. M. Adams, and W. H. Martin (eds.). John Wiley and Sons, New York, NY, p. 351-386.
Master, L. L. 1991. Assessing threats and setting priorities for conservation. Conservation Biology 5:559-563.
Means, D. B. 1988. Management recommendations for the gopher tortoise in the longleaf pine ecosystem. In Gopher Tortoise Habitat Management: Strategies and Options. C. K. Dodd, Jr. (ed.). Proceedings of the Sixth Annual Meeting of the Gopher Tortoise Council, Florida Museum of Natural History, Gainesville, FL, p. 41-56.
Means, D. B., and G. Grow. 1985. The endangered longleaf pine community. ENFO Report, Environmental Information Center of the Florida Conservation Foundation, Inc., Winter Park, FL, 12 p.
Means, D. B., and P. E. Moler. 1979. The pine barrens treefrog: Fire, seepage bogs, and management implications. In Proceedings of the Rare and Endangered Wildlife Symposium. R. R. Odum, and L. Langers (eds.). Technical Bulletin WL-4, Game and Fish Division, Georgia Department of Natural Resources, Atlanta, GA, 77-83.
Means, D. B., J. G. Palis, and Mary Baggett. 1996. Effects of slash pine silviculture on a Florida population of flatwoods salamander. Conservation Biology 10:426-437.
Millsap, B. A., J. A. Gore, D. E. Runde, and S. I. Cerulean. 1990. Setting priorities for the conservation of fish and wildlife species in Florida. Wildlife Monographs 111:1-57.
Mitchell, J. C. 1994. Habitat conservation assessment for the Cow Knob salamander (Plethodon punctatus) in the George Washington National Forest. Report submitted to the U.S. Fish and Wildlife Service, Annapolis, MD, and the George Washington National Forest, Harrisonburg, VA, 16 p.
Moler, P. E., and R. Franz. 1987. Wildlife values of small, isolated wetlands in the Southeastern Coastal Plain. In Proceedings of the Third Southeastern Nongame and Endangered Wildlife Symposium. R. R. Odom, K. A. Riddleberger, and J. C. Ozier (eds.). Georgia Department of Natural Resources, Atlanta, GA, p. 234-241.
Natural Heritage Data Center Network. 1993. Perspectives on species imperilment, Revised printing. The Nature Conservancy, Arlington, VA, 40 p.
Noss, R. F. 1988. The longleaf pine landscape of the Southeast: Almost gone and almost forgotten. Endangered Species Update 5:1-6.
OíConnell, M. A., and R. F. Noss. 1992. Private land management for biodiversity conservation. Environmental Management 16:435-450.
Palis, J. G. 1992. Distribution of the flatwoods salamander, Ambystoma cingulatum, on the Apalachicola and Osceola National Forests, Florida. Unpublished Report submitted to the Florida Natural Areas Inventory, Tallahassee, FL, 19 p.
Palis, J. G. 1993. A status survey of the flatwoods salamander, Ambystoma cingulatum, in Florida. Unpublished Report submitted to the U.S. Fish and Wildlife Service, Jackson, MS, 36 p.
Palis, J. G. 1995a. A survey of the flatwoods salamander (Ambystoma cingulatum) breeding sites east of the Apalachicola River, Florida. Unpublished Report submitted to the U.S. Fish and Wildlife Service, Jackson, MS, 40 p.
Palis, J. G. 1995b. Distribution and breeding biology of the flatwoods salamander (Ambystoma cingulatum) and gopher frog (Rana capito) on Eglin Air Force Base, Florida. Florida Natural Areas Inventory, Tallahassee, FL, 46 p. + appendices.
Pechmann, J. H. K., D. E. Scott, R. D. Semlitsch, J. P. Caldwell, L. J. Vitt, and J. W. Gibbons. 1991. Declining amphibian populations: The problem of separating human impacts from natural fluctuations. Science 253:892-895.
Pechmann, J. H. K., and H. M. Wilbur. 1994. Putting declining amphibian populations in perspective: Natural fluctuations and human impacts. Herpetologica 50:65-84
Petranka, J. W., M. E. Elridge, and K. E. Haley. 1993. Effects of timber harvesting on southern Appalachian salamanders. Conservation Biology 7:363-370.
Phelps, J. P., and R. A. Lancia. 1995. Effects of a clearcut on the herpetofauna of a South Carolina bottomland swamp. Brimleyana 22:31-45.
Raymond, L. R., and L. M. Hardy. 1991. Effects of a clearcut on a population of the mole salamander, Ambystoma talpoideum, in an adjacent unaltered forest. Journal of Herpetology 25:509-512.
Schemnitz, S. D. (ed.). 1980. Wildlife Management Techniques Manual. The Wildlife Society, Washington, D.C.
Scott, N. J., Jr., and R. A. Seigel. 1992. The management of amphibian and reptile populations: Species priorities and methodological and theoretical constraints. In Wildlife 2001: Populations. D. R. McCullough, and R. H. Barrett (eds.). Elsevier Applied Science, New York, NY, p. 343-367.
Sedell, J. R., G. H. Reeves, R. R. Hauer, J. A. Stanford, and C. P. Hawkins. 1990. Role of refugia in recovery from disturbances: Modern fragmented and disconnected river systems. Environmental Management 14:711-724.
Seehorn, M. E. 1982. Reptiles and amphibians of southeastern national forests. Forest Service, U.S. Department of Agriculture, Atlanta, GA, 84 p.
Stiven, A. E., and R. C. Bruce. 1988. Ecological genetics of the salamander Desmognathus quadramaculatus from disturbed watersheds in the Southern Appalachian Biosphere Reserve Cluster. Conservation Biology 2:194-205.
Szaro, R. E., K. E. Severson, and D. R. Patton (eds.). 1988. Management of Amphibians, Reptiles, and Small Mammals in North America. General Technical Report RM-166, Forest Service, U.S. Department of Agriculture, Fort Collins, CO, 458 p.
Telford, S. 1993. Breeding sites for the gopher frog and the striped newt in the Ocala National Forest, and the discovery of a possibly undescribed species of frog apparently restricted to the forest. Unpublished Report submitted to staff at Ocala National Forest, Ocala, FL, 29 p.
The Nature Conservancy and the International Network of Natural Heritage Programs and Conservation Data Centers. 1996. Biological and conservation databases. The Nature Conservancy, Arlington, VA, 58 p.
Underwood, A. J. 1995. Ecological research and (research into) environmental management. Ecological Applications 5:232-247.
U.S. Fish and Wildlife Service. 1994a. Endangered and threatened wildlife and plants. 50 CFR 17.11 and 17.12. U.S. Government Printing Office, Washington, D.C., 42 p.
U.S. Fish and Wildlife Service. 1994b. Endangered and threatened wildlife and plants: Animal candidate review for listing as endangered or threatened species, proposed rule. U.S. Federal Register 50 CFR, Part 17, 59(219):58982-59028.
U.S. Fish and Wildlife Service. 1995. Ecosystem management plan for the southern Appalachian ecosystem. Asheville Field Office, U.S. Fish and Wildlife Service, Asheville, NC, 15 p.
U.S. Fish and Wildlife Service. 1996. Endangered and threatened species, plant and animal taxa; proposed rule. U.S. Federal Register 50 CFR, Part 17, 61(40):7596-7613.
Vial, J. L., and L. Saylor. 1993. The status of amphibian populations: A compilation and analysis. Working Document Number 1, IUCN/SSC Declining Amphibian Populations Task Force, Corvallis, OR, 98 p.
Vickers, C. R., L. D. Harris, and B. F. Swindel. 1985. Changes in herpetofauna resulting from ditching of cypress ponds in coastal plains. Forest Ecology Management 11:17-29.
Vitt, L. J., J. P. Caldwell, H. M. Wilbur, and D. C. Smith. 1990. Amphibians as harbingers of decay. BioScience 40:418.
Wallace, J. B., J. R. Webster, and R. L. Lowe. 1992. High-gradient streams of the Appalachians. In Biodiversity of the Southeastern United States: Aquatic Communities. C. T. Hackney, S. M. Adams, and W. H. Martin (eds.). John Wiley and Sons, New York, NY, p. 133-192.
Ware, S., C. Frost, and P. D. Doerr. 1993. Southern mixed hardwood forest: The former longleaf pine forest. In Biodiversity of the Southeastern United States: Lowland Terrestrial Communities. W. H. Martin, S. G. Boyce, and A. C. Echternacht (eds.). John Wiley and Sons, New York, NY, p. 447-493.
Yoffe, E. 1992. Silence of the frogs. New York Times Magazine 1992(December 13):36-39, 64, 66, 76.

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