# Endangered and Threatened Wildlife and Plants; Determination of Endangered Status for Reticulated Flatwoods Salamander; Designation of Critical Habitat for Frosted Flatwoods Salamander and Reticulated Flatwoods Salamander > Briefs, arguments, decisions, and more. URL: https://www.frixlaw.com/law-library/documents/fr%3AE9-2403 ## Record - **Collection:** Federal Register - **Document type:** Rule - **Published:** February 10, 2009 - **Citation:** 74 FR 6700 ## Text DEPARTMENT OF THE INTERIOR Fish and Wildlife Service 50 CFR Part 17 [FWS-R4-ES-2008-0082; MO 9921050083-B2] RIN 1018-AU85 Endangered and Threatened Wildlife and Plants; Determination of Endangered Status for Reticulated Flatwoods Salamander; Designation of Critical Habitat for Frosted Flatwoods Salamander and Reticulated Flatwoods Salamander AGENCY: Fish and Wildlife Service, Interior. ACTION: Final rule. SUMMARY: We, the U.S. Fish and Wildlife Service (Service), finalize the listing under the Endangered Species Act of 1973, as amended (Act), of the currently threatened flatwoods salamander ( Ambystoma cingulatum ) into two distinct species: Frosted flatwoods salamander ( Ambystoma cingulatum ) and reticulated flatwoods salamander ( Ambystoma bishopi ) due to a recognized taxonomic reclassification; determine endangered status for the reticulated flatwoods salamander; retain threatened status for the frosted flatwoods salamander; and designate critical habitat for the frosted flatwoods salamander and the reticulated flatwoods salamander. In total, approximately 27,423 acres (ac) (11,100 hectares (ha)) in 35 units or subunits fall within the boundaries of the critical habitat designation; 22,970 ac (9,297 ha) of critical habitat is designated for the frosted flatwoods salamander and 4,453 ac (1,803 ha) for the reticulated flatwoods salamander. This area is a reduction of 3,205 ac (977 ha) from the proposed designation; 162 ac (66 ha) less for the frosted flatwoods salamander and 3,043 ac (928 ha) less for the reticulated flatwoods salamander. The critical habitat is located in Baker, Calhoun, Franklin, Holmes, Jackson, Jefferson, Liberty, Santa Rosa, Wakulla, Walton, and Washington Counties in Florida; Baker and Miller Counties in Georgia; and Berkeley, Charleston, and Jasper Counties in South Carolina. DATES: This rule becomes effective on March 12, 2009. ADDRESSES: This final rule and final economic analysis are available on the Internet at http://www.regulations.gov. Supporting documentation we used in preparing this final rule is available for public inspection, by appointment, during normal business hours, at U.S. Fish and Wildlife Service, Mississippi Fish and Wildlife Office, 6578 Dogwood View Parkway, Jackson, MS 39213. FOR FURTHER INFORMATION CONTACT: Ray Aycock, Field Supervisor, U.S. Fish and Wildlife Service, Mississippi Field Office, 6578 Dogwood View Parkway, Jackson, MS 39213; telephone: 601-321-1122; facsimile: 601-965-4340. If you use a telecommunications device for the deaf (TDD), call the Federal Information Relay Service (FIRS) at 800-877-8339. SUPPLEMENTARY INFORMATION: This document consists of: (1) A final rule to change the listing of the currently threatened flatwoods salamander ( Ambystoma cingulatum ) to frosted flatwoods salamander ( Ambystoma cingulatum ) and reticulated flatwoods salamander ( Ambystoma bishopi ) (the frosted flatwoods salamander will continue to be listed as threatened and the reticulated flatwoods salamander is listed as endangered); and (2) final critical habitat designations for each species. Previous Federal Actions The flatwoods salamander was listed as threatened on April 1, 1999 (64 FR 15691). At that time, we found that designation of critical habitat for the flatwoods salamander was not prudent because such designation would not be beneficial and may increase threats to the species. On April 1, 2005, Center for Biological Diversity, Wild South, and Florida Biodiversity Project filed a lawsuit against the Secretary of the Interior alleging failure to designate critical habitat for the flatwoods salamander. In a court-approved settlement agreement, we agreed to re-evaluate the need for critical habitat for the species and, if prudent, submit a proposed designation of critical habitat to the Federal Register by January 30, 2007, and submit a final critical habitat rule for publication in the Federal Register by January 30, 2008. We published a proposed rule to designate critical habitat for the flatwoods salamander in the Federal Register on February 7, 2007 (72 FR 5856). After that proposed rule published, new information became available on its taxonomic classification and additional threats to occupied habitat that necessitated a reevaluation of the proposed rule. On January 25, 2008, the court-approved settlement agreement was modified to require that a revised proposed critical habitat designation for the frosted flatwoods salamander and the reticulated flatwoods salamander be submitted for publication in the Federal Register on or before July 30, 2008, with the final critical habitat rule to be submitted for publication in the Federal Register by January 30, 2009. The revised proposed rule was signed on and delivered to the Federal Register on July 30, 2008, and it subsequently published on August 13, 2008 (73 FR 47258). We also published supplemental information on the proposed rule to maintain the status of the frosted flatwoods salamander as threatened (73 FR 54125; September 18, 2008). Public Comments Due to the nature of the proposed rule, we received combined comments from the public on the listing action and the critical habitat designation. Therefore, we have addressed these issues in a single comment section. In this final rule, we have presented the listing analysis first, followed by the analysis for designation of critical habitat. All public comments and our responses to them are presented under the Critical Habitat section. Background It is our intent to discuss only those topics directly relevant to the taxonomic reclassification of the flatwoods salamander into two species, the frosted flatwoods salamander and the reticulated flatwoods salamander, the determination of the status of these two species, and the designation of critical habitat for both species. For more information on the biology and ecology of flatwoods salamanders, refer to the final listing rule published in the Federal Register on April 1, 1999 (64 FR 15691). For information on our proposed determination of endangered status for the reticulated flatwoods salamander, and on the proposed designation of critical habitat for the frosted flatwoods salamander and the reticulated flatwoods salamander, refer to the proposed rule published in the Federal Register on August 13, 2008 (73 FR 47258). Taxonomic Classification The original listing rule (64 FR 15691; April 1, 1999) described the geographic range of the flatwoods salamander as it was known at that time. The range for the species included occurrences across the lower southeastern Coastal Plain in Florida, Georgia, and South Carolina. Taxonomic revision resulted from research done by Pauly et al. (2007, pp. 415-429) that suggested a taxonomic reclassification of the species by splitting the flatwoods salamander into two species—the frosted flatwoods salamander and the reticulated flatwoods salamander. The Apalachicola River drainage forms a geographic barrier between the two species. This drainage is a common site for east-west phylogeographic breaks in many other taxa as well. For this reason, the reclassification of the flatwoods salamander into two species is currently accepted by the scientific community and by the Service. We hereby amend the List of Endangered and Threatened Wildlife at 50 CFR 17.11(h) to reflect this revision to taxonomy. Goin (1950, p. 299) recognized two distinct subspecies of flatwoods salamander based on morphological and color pattern variation. This reclassification between the eastern and western portions of the salamander's range was later discounted in an analysis by Martof and Gerhardt (1965, pp. 342-346) and for the past 40 years the concept of a single undifferentiated species persisted. Pauly et al. (2007, pp. 415-429) conducted molecular and morphological analyses to test whether the flatwoods salamander, as originally described, followed a pattern of east-west disjunction at the Apalachicola River as has been described in many other species. They were able to demonstrate this predicted phylogeographic break. Based on mitochondrial DNA (mtDNA), morphology, and allozymes, they recognize two species of flatwoods salamanders, frosted flatwoods salamander to the east of the Apalachicola drainage and reticulated flatwoods salamander to the west. The Apalachicola River is probably the cause of major disjunctions in species distributions due to the repeated marine embayments during the Pliocene and Pleistocene interglacials that likely caused a barrier to gene flow. In the Pauly et al. (2007, pp. 415-429) analyses, the use of mtDNA splits flatwoods salamander populations into two major clades east and west of the Apalachicola-Flint rivers. Samples from Jackson and Liberty Counties, Florida, are informative because, geographically, they are located on opposite sides of the river but are phylogenetically distant with respect to mtDNA sequence divergence. In contrast, geographically distant populations on the same side of the Apalachicola River are very closely related. Their morphological analyses also support a taxonomic boundary at the Apalachicola-Flint rivers. Salamanders on opposite sides of this boundary significantly differed in both body shape and size based on multivariate analyses. The number of costal grooves (grooves along the side body of salamanders used in species identification), snout-vent length, six additional morphometric traits, and sexual dimorphisms in tail length, height, and width are all significantly different between the two taxa. Due to the importance of the tail in ambystomatid courtship and fertilization, tail differences may be particularly important (Duellman and Trueb 1986, pp. 64-66). Allozyme data presented in Shaffer et al. (1991, pp. 290-291, 302) also indicated differences between salamanders on either side of the Apalachicola River. Their results demonstrated these populations have fixed-allele differences, consistent with the mtDNA and morphological results. The frosted and reticulated flatwoods salamanders can be differentiated from each other by the use of several morphological characters (Pauly et al. 2007, pp. 424-425). The frosted flatwoods salamander generally has more costal grooves and tends to be larger than the reticulated flatwoods salamander. For individuals of the same size, the frosted flatwoods salamander has longer forelimbs and hind limbs and a larger head. Male frosted flatwoods salamanders have longer tails than those of the reticulated flatwoods salamander. The belly pattern of the frosted flatwoods salamander consists of discrete white spots on a dark background, while the spots are less distinct in the reticulated flatwoods salamander giving a “salt and pepper” appearance (Goin 1950, pp. 300-314). The back pattern of the reticulated flatwoods salamander has a more net-like appearance than the frosted flatwoods salamander, as the common names imply. In summary, in the Regulation Promulgation section of this document, we present a taxonomic change reflecting the reclassification of flatwoods salamander ( Ambystoma cingulatum ) to frosted flatwoods salamander ( A. cingulatum ) and reticulated flatwoods salamander ( A. bishopi ). Listing of the Reticulated Flatwoods Salamander History of the Action On December 16, 1997, we published a proposed rule to list the flatwoods salamander as a threatened species (62 FR 65787). We published the final rule to list the species on April 1, 1999 (64 FR 15691). On August 13, 2008, we published the proposal to list the reticulated flatwoods salamander, currently known as the flatwoods salamander west of the Apalachicola-Flint Rivers, as a new species (73 FR 47258). Species Information As far as we currently know, the life history traits and habitat use of both the frosted flatwoods salamander and the reticulated flatwoods salamander are similar to those previously described for the flatwoods salamander. Both species of flatwoods salamanders are moderately sized salamanders that are generally black to chocolate-black with fine, irregular, light gray lines and specks that form a cross-banded pattern across their backs (back pattern more net-like in the reticulated flatwoods salamander). The frosted flatwoods salamander generally tends to be larger than the reticulated flatwoods salamander, as described above. Adults are terrestrial and live underground most of the year. They breed in relatively small, isolated ephemeral ponds where the larvae develop until metamorphosis. Post-metamorphic salamanders migrate out of the ponds and into the uplands where they live until they move back to ponds to breed as adults. Both species of flatwoods salamander are endemic to the lower southeastern Coastal Plain and occur in what were historically longleaf pine-wiregrass flatwoods and savannas (Palis and Means 2005, pp. 608-609). The historical range of what is now considered the reticulated flatwoods salamander included parts of the States of Alabama, Florida, and Georgia, which are in the lower Coastal Plain of the southeastern United States west of the Apalachicola-Flint Rivers. We have compiled 26 historical (pre-1990) records for the reticulated flatwoods salamander. In Alabama, there are five historical localities for the reticulated flatwoods salamander, all in the extreme southern portion of the State in Baldwin, Covington, Houston, and Mobile Counties. Surveys have been conducted at numerous sites since 1992; however, no reticulated flatwoods salamanders have been observed in Alabama since 1981 (Jones et al. 1982, p. 51; Godwin 2008). Two historical records for the reticulated flatwoods salamander are known from Georgia, one each in Baker and Early Counties. Site visits to the areas in the vicinity of these two records have indicated that there is no longer suitable habitat for flatwoods salamanders at these localities. The area of the Baker County record has been cleared for agriculture (LaClaire 1994b). The upland habitat surrounding the Early County record has been converted to home sites and agricultural fields (Seyle 1994, p. 4). Four new reticulated flatwoods salamander breeding ponds have been discovered since 1990. One pond is on the Mayhaw Wildlife Management Area owned by the State of Georgia in Miller County. Three ponds are on private property in Baker County. Currently, two reticulated flatwoods salamander populations are supported by these breeding sites in Georgia. Nineteen historical (pre-1990) records for the reticulated flatwoods salamander are known for Florida. Reticulated flatwoods salamander breeding has been documented at only five (26 percent) of these sites since 1990. Extensive surveys throughout the range of the Ambystoma cingulatum, conducted prior to the original listing in 1999, resulted in identifying 39 additional breeding sites. Thirty-one (80 percent) of these sites are located in Okaloosa and Santa Rosa Counties, primarily on Department of Defense lands. Currently, 18 populations of the reticulated flatwoods salamander are known from Florida. The combined data from all survey work completed since 1990 in Florida and Georgia indicate there are 20 populations of the reticulated flatwoods salamander. Some of these populations are inferred from the capture of a single individual. Nine (45 percent) of the known reticulated flatwoods salamander populations occur, at least in part, on public land. Of these, Department of Defense lands in Florida harbor four populations of the reticulated flatwoods salamander at Eglin Air Force Base, Hurlburt Field, and Navy Outlying Landing Field Holley. State and local agencies in Florida and Georgia partially manage habitat for five additional populations and monitor breeding ponds. In Florida, Pine Log State Forest harbors a single population; Northwest Florida Water Management District (NWFLWMD) and Blackwater River State Forest share management of a single population; NWFLWMD and Yellow River Marsh Preserve State Park share management of most of another property supporting an additional population; and the Santa Rosa County School Board owns a portion of the habitat supporting a single population. In Georgia, the Mayhaw Wildlife Management Area supports a single population. Eleven (55 percent) reticulated flatwoods salamander populations are solely on private land. Summary of Factors Affecting the Reticulated Flatwoods Salamander Section 4 of the Act and regulations (50 CFR part 424) promulgated to implement the listing provisions of the Act set forth the procedures for adding species to Federal lists. A species may be determined to be an endangered or threatened species due to one or more of the five factors described in section 4(a)(1). The original listing rule for the flatwoods salamander (64 FR 15691) contained a discussion of these five factors, as did the proposed rule (73 FR 47258; August 13, 2008) and supplemental information (73 FR 54125; September 18, 2008). Only those factors relevant to the proposed reclassification of the reticulated flatwoods salamander ( Ambystoma bishopi Goin, 1950) from threatened to endangered are described below: A. The Present or Threatened Destruction, Modification, or Curtailment of Its Habitat or Range The major threat to the reticulated flatwoods salamander is loss of both its longleaf pine-slash pine flatwoods terrestrial habitat and its isolated, seasonally ponded breeding habitat. The combined pine flatwoods (longleaf pine-wiregrass flatwoods and slash pine flatwoods) historical area was approximately 32 million acres (ac) (12.8 million hectares (ha)) (Outcalt 1997, p. 4). This area has been reduced to 5.6 million ac (2.27 million ha) or approximately 18 percent of its original extent (Outcalt 1997, p. 4). These remaining pine flatwoods (non-plantation forests) areas are typically fragmented, degraded, second-growth forests (Outcalt 1997, p. 6). Conversion of pine flatwoods to intensively managed (use of heavy mechanical site preparation, high stocking rates, and low fire frequencies) slash or loblolly plantations often resulted in degradation of flatwoods salamander habitat by creating well-shaded, closed-canopied forests with an understory dominated by shrubs or pine needles (Outcalt 1997, pp. 4-6; Palis 1997, pp. 61-63). Disturbance-sensitive ground cover species, such as wiregrass ( Aristida stricta [= A. beyrichiana ] Kesler et al. 2003, p. 9), dropseed ( Sporobolus spp.), and perennial forbs were either greatly reduced in extent or were replaced by weedy pioneering species (Moore et al. 1982, p. 216; Outcalt and Lewis 1988, pp. 1-12; Hardin and White 1989, pp. 243-244). In a study conducted by Hedman et al. (2000, p. 233), longleaf pine plots had significantly more herbaceous species and greater herbaceous cover than loblolly or slash pine plots. For example, wiregrass is often lost from a site when habitat is converted from longleaf pine forest to other habitat types using common mechanical site preparation methods (Outcalt and Lewis 1988, p. 2). Loss of wiregrass is considered an indicator of site degradation from fire suppression or soil disturbance (Clewell 1989; pp. 226, 230-232). Flatwoods salamanders are unlikely to persist in uplands with a disturbed, wiregrass-depauperate ground cover (Palis 1997, p. 63). Forest management that includes intensive site preparation may adversely affect flatwoods salamanders directly and indirectly (Means et al. 1996, p. 426). Bedding (a technique in which a small ridge of surface soil is elevated as a planting bed) alters the surface soil layers, disrupts the site hydrology, and often eliminates the native herbaceous ground cover. This can have a cascading effect of reducing the invertebrate community that serves as a food source for flatwoods salamander adults. Post-larval and adult flatwoods salamanders occupy upland flatwoods sites where they live underground in crayfish burrows, root channels, or burrows of their own making (Goin 1950, p. 311; Neill 1951, p. 765; Mount 1975, pp. 98-99; Ashton and Ashton 2005, pp. 63, 65, 68-71). The occurrence of these underground habitats is dependent upon protection of the soil structure. Intensive site preparation destroys the subterranean voids and may result in entombing, injuring, or crushing individuals. Ecologists consider fire suppression the primary reason for the degradation of remaining longleaf pine forest habitat. The disruption of the natural fire cycle has resulted in an increase in slash and loblolly pine on sites formerly dominated by longleaf pine, an increase in hardwood understory, and a decrease in herbaceous ground cover (Wolfe et al. 1988, p. 132). Although reticulated flatwoods salamanders have been found at sites with predominately loblolly or slash pine, the long-term viability of populations at these sites is unknown. On public lands, prescribed burning is a significant part of habitat management plans. However, implementation of prescribed burning has been inconsistent due to financial constraints and limitations of weather (drought, wind direction, etc.) that restrict the number of opportunities to burn. These alterations of the longleaf pine ecosystem, as a result of incompatible forest practices, have caused historic losses of reticulated flatwoods salamander habitat. Conversion of native pine flatwoods to plantation forests is not considered a significant threat at this time. Forecasts indicate that most new plantation forests will come from converting agricultural fields (Wear and Greis 2002, p. 47). Nevertheless, we have documented the historic extirpation of at least one previously known population each from Gulf and Jackson Counties in Florida, over the last 4 decades because of habitat degradation on lands currently managed as pine plantations. In addition, ponds surrounded by pine plantations and protected from the natural fire regime may become unsuitable as reticulated flatwoods salamander breeding sites due to canopy closure and the resultant reduction in emergent herbaceous vegetation needed for egg deposition and larval development sites (Palis 1997, p. 62). In addition, lack of fire within the pond during periods of dry-down may result in chemical and physical (vegetative) changes that are unsuitable for the salamander (Palis 1997, p. 62). Lack of fire in the ecotone may result in the development of a thick shrub zone making it physically difficult or impossible for adult salamanders to enter the breeding ponds (Ripley and Printiss 2005, pp. 1-2, 11). Land use conversions to urban development and agriculture eliminated large areas of pine flatwoods in the past (Schultz 1983, pp. 24-47; Stout and Marion 1993, pp. 422-429; Outcalt and Sheffield 1996, pp. 1-5; Outcalt 1997, pp. 1-6). Urbanization and agriculture have resulted in the loss of one reticulated flatwoods salamander population from each of the following counties: Mobile and Baldwin Counties, Alabama; Escambia, Jackson, and Washington Counties, Florida; and Early County, Georgia. Two known populations have been extirpated from Santa Rosa County, Florida. State forest inventories completed between 1989 and 1995 indicated that flatwoods losses through land use conversion were still occurring (Outcalt 1997, pp. 3-6). Urbanization in the panhandle of Florida and around major cities is reducing the available pine forest habitat. Wear and Greis (2002, pp. 47, 92) identify conversion of forests to urban land uses as the most significant threat to southern forests. They predict that the South could lose about 12 million ac (4.9 million ha) of pine forest habitat to urbanization between 1992 and 2020. Several relatively recent discoveries of previously unknown reticulated flatwoods salamander breeding sites in Santa Rosa County, Florida, have been made in conjunction with wetland surveys associated with development projects (Cooper 2008a). No reticulated flatwoods salamanders have been observed at these degraded sites since completion of the projects (Cooper 2008a). In addition to the loss of upland forested habitat, the number and diversity of small wetlands where reticulated flatwoods salamanders breed have been substantially reduced. Threats to breeding sites include alterations in hydrology, agricultural and urban development, road construction, incompatible silvicultural practices, shrub encroachment, dumping in or filling of ponds, conversion of wetlands to fish ponds, domestic animal grazing, soil disturbance, and fire suppression (Vickers et al. 1985, pp. 22-26; Palis 1997, p. 58; Ashton and Ashton 2005, p. 72). Hydrological alterations, such as those resulting from ditches created to drain flatwoods sites or fire breaks and plow lines, represent one of the most serious threats to reticulated flatwoods salamander breeding sites. Lowered water levels and shortened hydroperiods at these sites may prevent successful flatwoods salamander recruitment because larval salamanders require 11 to 18 weeks to reach metamorphosis and leave the ponds (Palis 1995, p. 352). Drought conditions exacerbate other threats and, although they represent a natural phenomenon, can lower the resiliency of populations to withstand other man-made threats. The U.S. Geological Survey (USGS) has documented multiple drought periods in the southeastern United States since the 1890s (USGS 2000, p. 1). Significant drought periods documented in the last three decades are: 1980-1982, 1984-1989, 1998-2002, 2005-2008 (USGS 1991, p. 163; USGS 2000, p. 1; Seager et al. 2008, pp. 2, 22). Although a naturally occurring condition, drought presents additional complications for a species, like reticulated flatwoods salamander, which has been extirpated from most of its historic range and for which populations are represented by single ponds. Palis et al. (2006, p. 5-6) conducted a study in Florida on a population of the closely related frosted flatwoods salamander during a drought from 1999-2002. This study found three consecutive years of reproductive failure and a steadily declining adult immigration to breed at the site as the drought progressed. Taylor et al. (2005, p. 792) noted that wide variation in reproductive success is common among pond-breeding amphibians that depend on seasonal filling of these areas, but that adult persistence may buffer against fluctuations in that success, particularly for species that are long-lived. Although Palis et al. (2006, p. 6) suggested that the flatwoods salamander may only live about 4 years (based on captive animals), we are currently unsure of the exact lifespan of wild individuals. Other sources have suggested 10 years may represent a maximum lifespan (Jensen 2008). As a result, it is difficult to predict how long adults could persist in the landscape without a successful breeding event to replenish the population. However, Taylor et al. (2005, pp. 792, 796) constructed a model, based on extensive population data available for the marbled salamander ( Ambystoma opacum ), to look at how many years of reproductive failure would be required to result in local extinction of pond-breeding salamanders (with varying lifespans) and found that even without total reproductive failure, populations required moderate to high upland post-metamorphic survival to persist. Catastrophic failure in this study created fluctuations in the population, raised the threshold of survival required to achieve persistence, and imposed the possibility of extinction even under otherwise favorable environmental conditions. Reproductive failure was closely tied to hydrologic conditions; insufficient or short hydroperiod was the primary cause for complete failure. In addition, early filling of the ponds could also facilitate the establishment of invertebrate or vertebrate predators before hatching of the eggs (Taylor et al. 2005, p. 796). Palis et al. (2006, p. 6-7) discussed the necessity of protecting clusters of flatwoods salamander breeding sites, especially those with different hydrologic regimes, to guard against population declines at any one breeding site resulting from random events, such as droughts (Palis 2006, p. 7). A cluster of breeding sites represents a metapopulation, which is defined as neighboring local populations close enough to one another that dispersing individuals could be exchanged (gene flow) at least once per generation. Currently, the only place where a metapopulation exists for the reticulated flatwoods salamander is on Eglin Air Force Base. Habitat fragmentation of the longleaf pine ecosystem resulting from habitat conversion threatens the survival of the reticulated flatwoods salamander. Large tracts of intact longleaf pine flatwoods habitat are fragmented by pine plantations, roads, and unsuitable habitat. Most reticulated flatwoods salamander populations are widely separated from each other by unsuitable habitat. This has been verified through recent reviews of aerial photography and site visits to localities of historical and current records for the species. Studies have shown that the loss of fragmented populations is common, and recolonization is critical for their regional survival (Fahrig and Merriam 1994, pp. 50-56; Burkey 1995, pp. 527-540). Amphibian populations may be unable to recolonize areas after local extirpations due to their physiological constraints, relatively low mobility, and site fidelity (Blaustein et al. 1994, pp. 60, 67-68). In the case of the reticulated flatwoods salamander, 70 percent of populations only have one breeding pond and if the habitat at that one site is destroyed, recolonization would be impossible (see further discussion of metapopulation dynamics under Factor E). Roads contribute to habitat fragmentation by isolating blocks of remaining contiguous habitat. They may disrupt migration routes and dispersal of individuals to and from breeding sites. Road construction can result in changes in hydrology and destruction of breeding ponds, as described above. In addition, vehicles may also cause the death of reticulated flatwoods salamanders when they are attempting to cross roads (Means 1996, p. 2). Road construction resulted in the destruction of a historic reticulated flatwoods salamander breeding pond in Escambia County, Florida (Palis 1997, p. 62). A road through Eglin Air Force Base (Eglin) and Hurlburt Field has been proposed by the Northwest Florida Transportation Corridor Authority (NWFTCA) (NWFTCA 2007). We are currently in consultation regarding this bypass project. The conceptually approved route for the project, as currently proposed, places the road adjacent to or through 22 breeding sites that support the largest reticulated flatwoods salamander population (Mittiga 2007). However, the Service has been assured by Eglin that they will not allow negative impacts to the salamander's habitat and that they will continue to ensure the conservation of the reticulated flatwoods salamander (Department of the Air Force (DoAF) 2008a, p. 1; 2008b, p. 1). The Service will work with Eglin to protect these breeding sites which represent the only population of this species supported by more than three breeding ponds and functioning as a metapopulation. In summary, the loss of habitat is a significant threat to the reticulated flatwoods salamander. This threat is compounded by current drought conditions and the nature of pond-breeding salamanders to undergo periodic reproductive failure. We consider this threat to be imminent and of high magnitude because of this species' narrow range and the rapid rate of habitat loss that is currently occurring within the range of this species. Thirteen (65 percent) of the reticulated flatwoods salamander populations are partly or completely on private land where habitat continues to be degraded by management that frequently includes fire suppression and intensive site preparation that alters surface soil layers, disrupts site hydrology, disturbs the ground cover, and which has the potential to entomb, injure, or crush individual salamanders. Forest management conducted in this way is considered incompatible for maintaining flatwoods salamander populations. Range-wide historic losses of both upland and wetland habitat have occurred due to conversion of flatwoods sites to agriculture, urban development, and intensively managed pine plantations. The remaining flatwoods habitat continues to be threatened by fire suppression and other incompatible forest management practices, road construction, and habitat fragmentation across the range of the species. Localized threats to existing wetland breeding sites include alterations in hydrology from agriculture, urban development, road construction, and incompatible forest management; and fire suppression. As a result, we have determined that the present or threatened destruction, modification, or curtailment of the reticulated flatwoods salamander's habitat and range represents an imminent and significant threat to the species. B. Overutilization for Commercial, Recreational, Scientific, or Educational Purposes Overcollecting does not appear to be a threat to the reticulated flatwoods salamander at this time. There is no evidence of a past or current problem with collection of this species. Consequently, we have determined that the factor of overutilization for commercial, recreational, scientific, or educational purposes is not a threat to the reticulated flatwoods salamander at this time. C. Disease or Predation Although disease has not been specifically documented in the reticulated flatwoods salamander thus far, disease outbreaks with mass mortality in other species of salamanders indicate that disease may be a threat for this species as well (Daszak et al. 1999, p. 736). “Red-leg” disease ( Aeromonas hydrophila ), a pathogen bacterium, caused mortality of the mole salamander ( A. talpoideum ) at the breeding pond of the reticulated flatwoods salamander in Miller County, Georgia (Maerz 2006), and reticulated flatwoods salamanders have not been observed at this site since the disease was reported. In addition, Whiles et al. (2004, p. 211) found a parasitic nematode ( Hedruris siredonis , family Hedruridae) in larvae of the closely related frosted flatwoods salamander from South Carolina and Florida. This parasite has been found in other ambystomatids and can cause individuals to become undersized and thin, thus reducing their fitness (Whiles et al. 2004, p. 212). The infestations were not considered heavy and were probably not having a negative impact on the larvae studied; however, environmental degradation may change the dynamics between salamander populations and normally innocuous parasites (Whiles et al. 2004, p. 212). Ranaviruses in the family Iridoviridae and the amphibian chytrid fungus ( Batrachochytrium dendrobatidis ) may be other potential threats, although the susceptibility of the reticulated flatwoods salamander to these diseases is unknown. Ranaviruses have been responsible for die-offs of tiger salamanders throughout western North America and spotted salamanders ( A. maculatum ) in Maine (Daszak et al. 1999, p. 736). Chytrid fungus has been discovered and associated with mass mortality in tiger salamanders in southern Arizona and California, and the Santa Cruz long-toed salamander ( A. macrodactylum croceum ) (Vredenburg and Summers 2001, p. 151; Davidson et al. 2003, p. 601; Padgett-Flohr and Longcore 2005, p. 50). This discussion of disease in other species of closely related salamanders indicates the potential existence of similar threats to reticulated flatwoods salamander populations. Exposure to increased predation by fish is a threat to the reticulated flatwoods salamander when isolated, seasonally ponded wetland breeding sites are changed to or connected to more permanent wetlands inhabited by fish species not typically found in temporary ponds. Studies of other ambystomatid species have demonstrated a decline in larval survival in the presence of predatory fish (Semlitsch 1987, p. 481). Ponds may be modified specifically to serve as fish ponds or sites may be altered because of drainage ditches, firebreaks, or vehicle tracks that can all provide avenues for fish to enter the wetlands. Red imported fire ants ( Solenopsis invicta ) are potential predators of flatwoods salamanders, especially in disturbed areas. They have been seen in areas disturbed by the installation of drift fences at known breeding sites of the closely related frosted flatwoods salamander (Palis 2008). The severity and magnitude, as well as the long-term effect, of fire ants on reticulated flatwoods salamander populations are currently unknown. In summary, diseases of amphibians in the southeastern United States remain largely unstudied. However, given the incidence of disease in species that could be considered surrogates for flatwoods salamanders, the probability exists for similar infections to occur in reticulated flatwoods salamander populations. We consider this to be an imminent threat of moderate magnitude. Predation by fish is a historic threat that continues to be a localized problem when ditches, firebreaks, or vehicle ruts provide connections allowing the movement of fish from permanent water bodies into reticulated flatwoods salamander breeding sites. Sixty-five percent of reticulated flatwoods salamander breeding ponds are partly or completely on private land. This situation increases the probability of fish being introduced to a breeding site, which would then cause the breeding habitat to become unsuitable and result in the extinction of the population. Fire ants also have the potential of being a localized threat, particularly in disturbed areas. In addition, we believe that the threats described here would also act to exacerbate other threats to the species. Overall, we consider the threat within this factor to be imminent and of moderate magnitude because 70 percent of reticulated flatwoods salamander populations are supported by a single breeding pond; diseases and fish and invertebrate predators have been found at ponds within the species' range; and these diseases and predators are known to cause mortality or reproductive failure in related species. D. The Inadequacy of Existing Regulatory Mechanisms There are no existing regulatory mechanisms for the protection of the upland habitats where reticulated flatwoods salamanders spend most of their lives. Section 404 of the Clean Water Act is the primary Federal law that has the potential to provide some protection for the wetland breeding sites of the reticulated flatwoods salamander. However, due to recent case law ( Solid Waste Agency of Northern Cook County ( SWANCC ) v. U.S. Army Corps of Engineers 531 U.S. 159 (2001) ; Rapanos v. United States 547 U.S. 715 (2006)) , isolated wetlands are no longer considered to be under Federal jurisdiction (not regulatory wetlands). Wetlands are only considered to be under the jurisdiction of the U.S. Army Corps of Engineers (Corps) if a “significant nexus” exists to a navigable waterway or its tributaries. Currently, some Corps Districts do not coordinate with us on flatwoods salamanders and, since isolated wetlands are not considered under their jurisdiction, they are often not included on maps in permit applications (Brooks 2008). We are aware of two isolated wetlands that supported reticulated flatwoods salamander populations that have been lost since 2006 under this scenario. Longleaf pine habitat management plans have been written for public lands occupied by the reticulated flatwoods salamander. They include management plans for State-owned lands and integrated natural resource management plans (INRMPs) for Department of Defense lands. Most of the plans contain specific goals and objectives regarding habitat management that would benefit reticulated flatwoods salamanders including prescribed burning. However, because multiple-use is the guiding principle on most public land, protection of the flatwoods salamander may be just one of many management goals including timber production and military and recreational use. At the State and local levels, regulatory mechanisms are limited. Although not listed as threatened or endangered in Alabama, the reticulated flatwoods salamander is listed among those nongame species for which it is “unlawful to take, capture, kill, or attempt to take, capture or kill; possess, sell, trade for anything of monetary value, or offer to sell or trade for anything of monetary value” (Alabama Department of Conservation and Natural Resources 2008, p. 1). The flatwoods salamander is listed as a threatened species in the State of Georgia (Jensen 1999, pp. 92-93). This designation protects the species by preventing its sale, purchase, or possession in Georgia and by prohibiting actions that cause direct mortality or the destruction of its habitat on lands owned by the State of Georgia (Ozier 2008). There is only one known flatwoods salamander population on lands owned by the State of Georgia, and that is Mayhaw Wildlife Management Area. In 2001, the Florida Fish and Wildlife Conservation Commission (FFWCC) listed the flatwoods salamander (which would include the reticulated flatwoods salamander) as a species of special concern (FFWCC 2007, p. 2) and prohibited direct take except through permit. As part of the listing process, a statewide management plan was developed for the salamander in Florida (FFWCC 2001, p. 1-60). This plan sets an ambitious conservation goal of maintaining at least 129 self-sustaining populations of flatwoods salamanders (which would include both frosted and reticulated flatwoods salamander species) in Florida. The plan also outlines a monitoring plan for population status assessment, an implementation strategy for the management of populations, and areas for future research. The Alabama and Florida regulations offer no protection against the most significant threat to the reticulated flatwoods salamander, loss of habitat. In summary, existing regulatory mechanisms provide little direct protection of reticulated flatwoods salamander habitat, the loss of which is the most significant threat to the species. Reticulated flatwoods salamander breeding sites may in some instances come under the jurisdiction of the Corps, but most often they are provided little regulatory protection. These inadequacies represent range-wide historic and known threats to the reticulated flatwoods salamander on private lands within the range. We consider this threat as imminent because the existing regulations are not protecting against the other imminent threats to the species. Also, this threat is of high magnitude because of the small range of the species, and because 65 percent of populations are not protected from further development because they are located partially or completely on private lands. E. Other Natural or Manmade Factors Affecting Its Continued Existence Metapopulations are important to the long-term survival of temporary pond breeding amphibians. In these species, such as the reticulated flatwoods salamander, breeding ponds may differ in the frequency of their ability to support amphibian reproduction. As a result, extirpation and colonization rates can be a function of pond spatial arrangement as well as local habitat quality (Marsh and Trenham 2001, p. 41). Of the 20 known reticulated flatwoods salamanders populations, only 6 (30 percent) are supported by more than one breeding pond and only one (5 percent) population (on Eglin-Hurlburt Field) is supported by more than three breeding ponds. For 70 percent (14 out of 20) of the known reticulated flatwoods salamander populations, any one of the many threats that may render a breeding pond unsuitable could cause the extirpation of the affected population. Invasive plant species, such as cogongrass ( Imperata cylindrica ), threaten to further degrade existing flatwoods habitat. Cogongrass, a perennial grass native to Southeast Asia, is one of the leading threats to the ecological integrity of native herbaceous flora, including that in the longleaf pine ecosystem (Jose et al. 2002, p. 43). Cogongrass can displace most of the existing vegetation except large trees. Especially threatening to the reticulated flatwoods salamander is the ability of cogongrass to outcompete wiregrass, a key vegetative component of flatwoods salamander habitat. Changing the species composition in this way can alter the soil chemistry, nutrient cycling, and hydrology of an infested site (Jose et al. 2002, p. 43). Reticulated flatwoods salamander habitat management plans will need to address threats posed by cogongrass and other invasive plant species and include strategies to control them. An integrated management approach to controlling cogongrass is outlined in Jose et al. (2002, p. 42). Pesticides (including herbicides) may pose a threat to amphibians, such as the reticulated flatwoods salamander, because their permeable eggs and skin readily absorb substances from the surrounding aquatic or terrestrial environment (Duellman and Trueb 1986, pp. 199-200). Negative effects that commonly used pesticides and herbicides may have on amphibians include delayed metamorphosis, paralysis, reduced growth rate, and mortality (Bishop 1992, pp. 67-69). In addition, herbicides used near reticulated flatwoods salamander breeding ponds may alter the density and species composition of vegetation surrounding a breeding site and reduce the number of potential sites for egg deposition, larval development, or shelter for migrating salamanders. However, if application by aerial spraying is avoided, the potential for negative effects from pesticide and herbicide use in areas adjacent to breeding ponds would be reduced (Tatum 2004, p. 1047). Herbicides may be a necessary tool to reduce or eliminate woody vegetation or invasive plants when the use of prescribed fire is not possible or effective (Jensen 2007, Wigley 2008). Nevertheless, pesticides should not be used in flatwoods salamander habitat unless no other habitat management tool is available; herbicide label directions should be followed closely; and aerial spraying should never be used as an application technique. Under these conditions, we consider this threat to be of moderate magnitude. Studies of other ambystomatid species have demonstrated a decline in larval survival in the presence of predatory fish, as mentioned above under Factor C. One of the potential reasons for this decline may be the negative effect resulting from these fish competing with salamander larvae for invertebrate prey. The invertebrates found by Whiles et al. (2004, p. 212) in a study of larval frosted and reticulated flatwoods salamander gut contents are typical of freshwater habitats in the Southeast that do not contain predatory fish on a regular basis. The presence of predatory fish has a marked effect on invertebrate communities and alters prey availability for larval salamanders with the potential for negative effects on larval fitness and survival (Semlitsch 1987, p. 481). Wherever connections have been created between permanent water and flatwoods salamander ponds, such as through installation of firebreaks or ditches, this threat from predatory fish exists. Studies of reticulated flatwoods salamander populations, since the original species listing of flatwoods salamander as threatened (64 FR 15691; April 1, 1999), have been limited due to drought. Data on the numbers of adults within existing populations do not exist. However, given the low number of individuals encountered even when breeding is verified, populations are likely to be very small at any given breeding site. Small populations are at increased threat of extirpation from natural processes (genetic isolation, inbreeding depression, and drought), as well as the manmade threats listed above. In summary, a variety of other natural or manmade factors historically or currently threaten, or have the potential to threaten, the reticulated flatwoods salamander. The loss of metapopulation structure in the distribution of reticulated flatwoods salamander populations was a range-wide threat that caused historic losses of this species. It continues to be a current threat for 70 percent of the remaining reticulated flatwoods salamander populations. Fire suppression and inadequate habitat management continue to cause the degradation of occupied sites, primarily on private land. Invasive plant species probably did not have much of a historic impact on salamander populations, but they are a range-wide current threat, and they are likely to become more widespread and difficult to control. Range-wide, low densities of individuals in a given population have been a historic threat and continue to be a threat for most reticulated flatwoods salamander populations, particularly due to past and current drought conditions, habitat loss, population fragmentation, and periodic reproductive failures that occur naturally in pond-breeding amphibians. The impact that competing predators may have on the salamander's prey base, and the threat of pesticide and herbicide use, are less clear as historic threats but remain potential localized threats for the species. Therefore, while we have determined that other natural and manmade factors, such as invasive species, pesticides, and competition for the species' prey base may threaten the reticulated flatwoods salamander, the severity and magnitude of these threats are not currently known. Acting in coordination with threats listed above under Factors A through D, the threats under Factor E could increase the severity of the other threats. In addition, small population size is particularly detrimental when combined with habitat loss, the ongoing drought, and the nature of this pond-breeding amphibian to experience periodic reproductive failure. Determination We have carefully assessed the best scientific and commercial information available regarding the past, present, and future threats to the reticulated flatwoods salamander. In summary, the most significant historical threat to the reticulated flatwoods salamander, as listed above in Factor A, is loss of its habitat. However, a variety of localized threats described under Factors A, C, D, and E continue to impact the remaining reticulated flatwoods salamander populations and their habitat. These include alterations in the hydrology of existing wetland breeding sites (including “ditching,” which can result in the introduction of predatory fish), urban development, road construction, incompatible forest management, fire suppression, and disease. The severity and magnitude of threats under Factor E are not currently known. Nevertheless, we have determined that threats under this factor will exacerbate the effects of threats due to habitat loss and drought. As described in Factor E above, small populations are at increased threat of extirpation from natural processes (genetic isolation, inbreeding depression, and drought), as well as the manmade threats listed above. Furthermore, as described in Factor D (above), existing regulatory mechanisms provide little direct protection of reticulated flatwoods salamander habitat, the loss of which is the most significant threat to the species. Reticulated flatwoods salamander breeding sites may in some instances come under the jurisdiction of the Corps, but most often they are provided little regulatory protection. This is likely the reason that two populations were lost recently to development. These inadequacies of existing regulatory mechanisms addressing habitat loss represent range-wide historic and potential threats to the reticulated flatwoods salamander. Finally, there are potential localized threats from fire ants, pesticides, and invasive plants for which the extent of impact is yet undeterminable, but we believe they are legitimate threats due to both their impact on surrogate species and their prevalence in the types of habitats used by this species. Only 20 reticulated flatwoods salamander populations are known. Fourteen (70 percent) of these populations are supported by only one breeding site. A population with only one breeding site has a tenuous future just given randomly varying environmental factors without considering the additional threats of habitat destruction and degradation that further threaten these populations. As noted previously, the habitat within the range of the reticulated flatwoods salamander is currently experiencing drought conditions. Palis et al. (2006, p. 5-6) studied a frosted flatwoods salamander population in Florida during a drought from 1999-2002. This study documented three consecutive years of reproductive failure and a steady decline in adult immigration to the site for breeding as the drought progressed. Catastrophic reproductive failure occurs even in healthy populations of pond-breeding amphibians. When it does occur, the modeling efforts of Taylor et al. (2005, p. 796) showed that each year of reproductive failure raises the threshold of survival required to achieve persistence and imposes the possibility of extirpation even under otherwise favorable environmental conditions. Taylor et al. (2005, p. 799) reminds us that, particularly with small populations or low population growth rates (as exists with the reticulated flatwoods salamander), the effects of reproductive failure are made worse by demographic stochasticity. Even in populations with multiple breeding ponds, amphibian populations may be unable to recolonize areas after local extirpations due to their physiological constraints, relatively low mobility, and site fidelity (Blaustein et al. 1994, pp. 60, 67-68). In the case of the reticulated flatwoods salamander, 70 percent of populations have only one breeding pond. If the habitat at that site is destroyed, recolonization would be impossible and the population supported by that breeding pond would be extirpated. Since the early 1990s, four reticulated flatwoods salamander populations have been lost, two populations due to urbanization and two populations due to incompatible forest management (Palis 2006, Cooper and LaClaire 2007, Cooper 2008b). The most robust reticulated flatwoods salamander population remaining is located on Eglin. Continued conservation of this locality is imperative because it represents habitat for the only population that is supported by more than three breeding ponds and functions as a metapopulation. In other words, this population has the best chance of surviving demographic and environmental stochasticity given that the distribution of breeding sites is within the dispersal distance of adult reticulated flatwoods salamanders. Based on the best available scientific and commercial information, we have determined that the reticulated flatwoods salamander is in danger of extinction throughout all or a significant portion of its range. Endangered status reflects the vulnerability of this species to factors that negatively affect the species and its limited and restricted habitat. Habitat loss on private lands is an imminent threat that is compounded by a variety of other factors. Fire suppression on private lands occupied by the reticulated flatwoods salamander represents one of the biggest threats to the species' habitat and the continued existence of the species on these sites. In addition, since 1999 we have lost at least two reticulated flatwoods salamander breeding ponds due to the threat of inadequate existing regulatory mechanisms. We believe the destruction of these ponds was a result of the continuing threat that isolated wetlands are rarely, if ever, under the jurisdiction of the Corps. We believe that, combined, the effect of the historical and ongoing drought; historical, current, and projected habitat loss and degradation; and the exacerbating effects of disease, predation, small population size, and isolation result in the reticulated flatwoods salamander being in danger of extinction throughout all of its range. We believe these threats, in particular the threats from habitat loss and drought, to be imminent and are projected to continue at the current rate or increase in the future. Further, we have determined that these threats are operating on the species and its habitat with a high degree of magnitude in that they affect the species throughout all of its range and with a high degree of severity, as discussed above. Listing of the Frosted Flatwoods Salamander History of the Action The final rule to list the flatwoods salamander as threatened was published on April 1, 1999 (64 FR 15691). On August 13, 2008, we published a proposed rule to reclassify the listing of the species into two distinct species: Frosted flatwoods salamander and reticulated flatwoods salamander due to new taxonomic information (73 FR 47258). In that proposed rule, we provided the analysis of the threats for the reticulated flatwoods salamander and our determination of its endangered status. On September 18, 2008, we published a notice providing supplemental information to the proposed rule that included our analysis and determination to retain threatened status for the frosted flatwoods salamander (73 FR 54125). Species Information Taxonomic revision resulting from research done by Pauly et al. (2007, pp. 415-429) split the flatwoods salamander into two species—the frosted flatwoods salamander and the reticulated flatwoods salamander. Background information on flatwoods salamanders, a discussion of their taxonomic status, and the five-factor analysis and associated determination of endangered status for the reticulated flatwoods salamander are provided above. Information provided here, and in the analysis that follows, will only address issues specific to the frosted flatwoods salamander. Based on the best available information, the life-history traits and habitat use of both the frosted flatwoods salamander and the reticulated flatwoods salamander are similar to those previously described for the flatwoods salamander (64 FR 15691, April 1, 1999; 73 FR 47258, August 13, 2008). However, most of our references predate Pauly et al. (2007, p. 415) and, therefore, do not distinguish between the two species. Flatwoods salamanders are endemic to the lower southeastern Coastal Plain and occur in what were historically longleaf pine-wiregrass flatwoods and savannas. The historical range of what is now considered the frosted flatwoods salamander included parts of the States of Florida, Georgia, and South Carolina. This area encompassed the lower Coastal Plain of the southeastern United States along the Gulf Coast east of the Apalachicola-Flint Rivers, across north Florida, south into north-central Florida, and north along the Atlantic Coast through coastal Georgia and South Carolina. We have compiled 84 historical (pre-1990) records for the frosted flatwoods salamander. Twenty historical records (with supporting locality information) for the frosted flatwoods salamander are known from eight counties in Florida. Frosted flatwoods salamander breeding has been documented at only four (20 percent) of these sites since 1990. Surveys conducted since 1990 by Federal and State agency personnel, as well as private parties, have resulted in the identification of more than 50 additional frosted flatwoods salamander breeding sites, including two sites in Jefferson County, a county that previously was not known to be occupied by the salamander. Most of these new breeding sites are located on the Apalachicola and Osceola National Forests, and on St. Marks National Wildlife Refuge. One site, discovered in 1998 on Tate's Hell State Forest, has been degraded as a result of habitat modification efforts that created a more permanently flooded wetland and flooded the ecotone at the historic breeding pond. The upland habitat is degraded as well with the result that the primary constituent elements (PCEs) on the site are no longer present (Enge 2008). Fifteen populations of the frosted flatwoods salamander are known from Baker, Franklin, Jefferson, Liberty, and Wakulla Counties in Florida. Thirty-four historical records for the frosted flatwoods salamander are known from 20 counties in Georgia. Frosted flatwoods salamanders have not been seen again at any of these sites in recent years; however, surveys conducted since 1990 have resulted in the discovery of 23 new breeding sites. All but one of these new sites are located on the Fort Stewart Military Installation. The one additional pond was discovered on the Townsend Bombing Range. Currently, these breeding sites support six frosted flatwoods salamander populations in Bryan, Evans, Liberty, and McIntosh Counties, Georgia, all on Department of Defense lands. The frosted flatwoods salamander is assumed to be extirpated from 16 other counties in Georgia where it previously occurred. However, some suitable habitat still remains on the Okefenokee National Wildlife Refuge and the potential exists for the species to occur there. Thirty historical records for the frosted flatwoods salamander are known from five counties in South Carolina. Since 1990, metamorphic frosted flatwoods salamanders have been documented at six (21 percent) of these sites, and one new breeding site has been discovered. Currently, four populations of the frosted flatwoods salamander are known from Berkeley, Charleston, and Jasper Counties in South Carolina. Two populations are on private land in Jasper County: One population occurs on the Francis Marion National Forest in Berkeley County, and one population occurs on the Santee Coastal Preserve (state-owned and managed) in Charleston County. The combined data from all survey work completed since 1990 in Florida, Georgia, and South Carolina indicate there are 25 populations of the frosted flatwoods salamander. Some of these populations are inferred from the capture of a single individual. Twenty-two (88 percent) of the known frosted flatwoods salamander populations occur primarily on public land. Sixteen of the populations (64 percent of total populations of the species) on public land represent metapopulations supported by more than one breeding site. A single population occurs on each of the following publicly owned sites: Osceola National Forest in Florida; Townsend Bombing Range in Georgia; and Francis Marion National Forest and Santee Coastal Reserve in South Carolina. In Florida, habitat supports 10 populations on Apalachicola National Forest and 2 populations on St. Marks National Wildlife Refuge. In Georgia, five populations occur on Fort Stewart Military Installation. Three (12 percent) frosted flatwoods salamander populations are solely on private land. Summary of Factors Affecting the Species (Frosted Flatwoods Salamander) Section 4 of the Act (16 U.S.C. 1531 et seq. ) and regulations (50 CFR part 424) promulgated to implement the listing provisions of the Act set forth the procedures for adding species to the Federal Lists of Endangered and Threatened Wildlife and Plants. A species may be determined to be endangered or threatened due to one or more of the five factors described in section 4(a)(1) of the Act. The original listing rule for the flatwoods salamander (64 FR 15691; April 1, 1999), and the supplement to the August 2008 proposed rule (73 FR 54125; September 18, 2008), contain a discussion of these five factors. Only those factors relevant to the frosted flatwoods salamander ( Ambystoma cingulatum Cope, 1867) are described below: A. The Present or Threatened Destruction, Modification, or Curtailment of Its Habitat or Range The major historical threat to the frosted flatwoods salamander was loss of habitat, both its longleaf pine-slash pine flatwoods terrestrial habitat and its isolated, seasonally ponded breeding habitat. Refer above to Factor A under “Summary of Factors Affecting the Species (Reticulated Flatwoods Salamander)” for general information on threats to pine flatwoods habitat that also applies to the frosted flatwoods salamander. Degradation of the remaining frosted flatwoods salamander habitat in Florida, Georgia, and South Carolina is a current, ongoing threat, primarily on private land. Ecologists consider fire suppression the primary reason for the degradation of remaining longleaf pine forest habitat. The disruption affects both the upland forested habitat of adult salamanders and their ponded breeding habitat also required for development of larval salamanders. Alterations of the longleaf pine ecosystem, as a result of incompatible forest practices, have caused the historic loss of most of the original frosted flatwoods salamander habitat. Conversion of native pine flatwoods to plantation forests is not considered a significant threat at this time. However, much of the historic extirpation of frosted flatwoods populations in Florida, Georgia, and South Carolina over the last six decades resulted from habitat degradation on lands managed for timber extraction. Land use conversions to housing, other development projects, and agriculture eliminated large areas of pine flatwoods in the past (Schultz 1983, pp. 24-47; Stout and Marion 1993, pp. 422-429; Outcalt and Sheffield 1996, pp. 1-5; Outcalt 1997, pp. 1-6). Residential development and conversion to agriculture have resulted in the historical loss of one frosted flatwoods salamander population each from Ben Hill, Berrien, Brooks, Effingham, Emanuel, and Irwin Counties, Georgia (Seyle 1994, pp. 4-5); an additional site has been degraded in Orangeburg County, South Carolina, and the population at this site is also considered extirpated (LaClaire 1994a). State forest inventories completed between 1989 and 1995 indicated that flatwoods losses through land use conversion were still occurring (Outcalt 1997, pp. 3-6); however, further conversions are only likely to impact three of the populations occurring in large part on private lands or only 12 percent of the total frosted flatwoods salamander populations. In addition to the loss of upland forested habitat, the number and diversity of small wetlands where frosted flatwoods salamanders breed have been substantially reduced. Threats to breeding sites include alterations in hydrology, agricultural and urban development, road construction, incompatible silvicultural practices, shrub encroachment, dumping in or filling of ponds, conversion of wetlands to fish ponds, domestic animal grazing, soil disturbance, and fire suppression (Vickers et al. 1985, pp. 22-26; Palis 1997, p. 58; Ashton and Ashton 2005, p. 72). As described above (see Species Information), the unintentional result of hydrological restoration on Tate's Hell State Forest was the destruction of the ephemeral nature of a reticulated flatwoods salamander breeding site and the extinction of the salamander population on that site. Drought conditions exacerbate other threats, and although they represent a natural phenomenon, can lower the resiliency of populations to withstand other man-made threats. The U.S. Geological Survey (USGS) has documented multiple drought periods in the southeastern United States since the 1890s (USGS 2000, p. 1). Among significant periods documented in the last three decades are: 1980-1982, 1984-1989, 1998-2002, 2005-2008 (USGS 1991, p. 163; USGS 2000, p. 1; Seager et al. 2008, pp. 2, 22). Although drought is a naturally occurring condition, it presents additional complications for a species like the frosted flatwoods salamander, which has been extirpated from most of its historic range. Palis et al. (2006, p. 5-6) conducted a study in Florida on a population of the frosted flatwoods salamander during a drought from 1999-2002. This study found 3 consecutive years of reproductive failure and a steadily declining adult immigration to breed at the site as the drought progressed. Palis et al. (2006, p. 6-7) discussed the necessity of protecting clusters of flatwoods salamander breeding sites, especially those with different hydrologic regimes, to guard against population declines at any one breeding site resulting from random events, such as droughts. Currently, 15 populations of the frosted flatwoods salamander, occurring on public land, are supported by multiple breeding sites. Habitat fragmentation of the longleaf pine ecosystem resulting from habitat conversion is primarily a historical threat to the frosted flatwoods salamander. Large tracts of intact longleaf pine flatwoods habitat are fragmented by pine plantations, roads, and unsuitable habitat. Although the threat of ongoing habitat fragmentation has slowed, the effect of past habitat loss is that many frosted flatwoods salamander populations are widely separated from each other by unsuitable habitat. This has been verified through recent reviews of aerial photography and site visits to localities of historical and current records for the species. Studies have shown that the loss of fragmented populations is common, and recolonization is critical for their regional survival (Fahrig and Merriam 1994, pp. 50-56; Burkey 1995, pp. 527-540). Amphibian populations may be unable to recolonize areas after local extirpations due to their physiological constraints, relatively low mobility, and site fidelity (Blaustein et al. 1994, pp. 60, 67-68). In the case of the frosted flatwoods salamander, 36 percent of populations have only one breeding pond. If the habitat at that site is destroyed, recolonization would be impossible (see further discussion of metapopulation dynamics under Factor E). Roads have contributed to habitat fragmentation by isolating blocks of remaining contiguous habitat. Roads disrupt migration routes and dispersal of individuals to and from breeding sites. Road construction can result in changes in hydrology and destruction of breeding ponds. Highway construction and associated development resulted in the destruction of a historic frosted flatwoods salamander breeding pond in Chatham County, Georgia (Seyle 1994, pp. 3-4). In addition, vehicles may also cause the death of frosted flatwoods salamanders when they are attempting to cross roads (Means 1996, p. 2). Off-road vehicle (ORV) use within frosted flatwoods salamander breeding ponds and their margins severely degrades the wetland habitat. In the Southeast, ORV use impacts habitat used by frosted flatwoods salamanders, has the potential to cause direct mortality of individual salamanders, and is a threat on both public and private land. On public lands, areas may be designated as off-limits to ORV use (U.S. Forest Service 2007, p. 19), but these restrictions are difficult to enforce. Even a single afternoon of individuals riding their ORVs in a pond can completely destroy the integrity of breeding sites by damaging or killing the herbaceous vegetation and rutting the substrate (Ripley and Printiss 2005, pp. 11-12). There is also the potential for direct injury or mortality of salamanders by ORVs at breeding sites (Ripley and Printiss 2005, p. 12). In summary, the loss of habitat was a significant historical threat to the frosted flatwoods salamander. This range-wide loss of both upland and wetland habitat occurred primarily due to conversion of flatwoods sites to agriculture, residential development, and intensively managed pine plantations. This historic loss of habitat is presently compounded by current environmental conditions (drought), proposed projects on private land that do not require Corps permits under the Clean Water Act (33 U.S.C. 1251 et seq. ), and the nature of pond-breeding salamanders to undergo periodic reproductive failure. We consider this threat to be primarily a past and future threat of moderate magnitude because most of the remaining occupied habitat of this species occurs on public lands that are managed to support the native longleaf pine ecosystem. However, 12 percent of frosted flatwoods salamander populations are on private land where habitat continues to be degraded by fire suppression and incompatible management (defined above under summary discussion for reticulated flatwoods salamander). If the remaining frosted flatwoods salamander habitat on public land continues to be protected from fire suppression and other incompatible forest management practices, road construction, and additional habitat fragmentation, the threat of habitat loss should be limited. Localized threats on private lands would include loss or alteration of habitat from agriculture, residential development, road construction, incompatible forest management, ORVs, fire suppression, and ditching or draining wetland breeding sites. As a result, we have determined that the present or threatened destruction, modification, or curtailment of frosted flatwoods salamander habitat and range represents a moderate but significant threat to the species. B. Overutilization for Commercial, Recreational, Scientific, or Educational Purposes Overutilization does not appear to be a threat to the frosted flatwoods salamander at this time. There is no evidence of a past or current problem with collection of this species. Consequently, we have determined that the factor of overutilization for commercial, recreational, scientific, or educational purposes is not a threat to the frosted flatwoods salamander at this time. C. Disease or Predation Although disease has not been specifically documented in the frosted flatwoods salamander thus far, disease outbreaks with mass mortality in other species of salamanders indicate that disease may be a threat for this species as well (Daszak et al. 1999, p. 736). Whiles et al. (2004, p. 211) found a parasitic nematode ( Hedruris siredonis , family Hedruridae) in larvae of the frosted flatwoods salamander from South Carolina and Florida. This parasite has been found in other ambystomatids and can cause individuals to become undersized and thin, thus reducing their fitness (Whiles et al. 2004, p. 212). The infestations were not considered heavy and were probably not having a negative impact on the larvae studied; however, environmental degradation may change the dynamics between salamander populations and normally innocuous parasites (Whiles et al. 2004, p. 212). Ranaviruses in the family Iridoviridae and the amphibian chytrid fungus ( Batrachochytrium dendrobatidis ) may be other potential threats, although the susceptibility of the frosted flatwoods salamander to these diseases is unknown. Ranaviruses have been responsible for die-offs of tiger salamanders throughout western North America and spotted salamanders ( A. maculatum ) in Maine (Daszak et al. 1999, p. 736). The amphibian chytrid fungus has been discovered and associated with mass mortality in tiger salamanders in southern Arizona and California, and the Santa Cruz long-toed salamander ( A. macrodactylum croceum ) (Vredenburg and Summers 2001, p. 151; Davidson et al. 2003, p. 601; Padgett-Flohr and Longcore 2005, p. 50). This fungus has been found at Fort Stewart Military Installation in Georgia, a locality where the frosted flatwoods salamander occurs (Mitchell 2002, p. 191-202). This disease has negatively impacted populations of other ambystomatid salamanders ( A. macrodactylum croceum ) (Vredenburg and Summers 2001, p. 151; Davidson et al. 2003, p. 601; Padgett-Flohr and Longcore 2005, p. 50), and it is likely to negatively impact frosted flatwoods salamander populations as well. This discussion of disease in other species of closely related salamanders indicates the potential existence of similar threats to frosted flatwoods salamander populations. Exposure to increased predation by fish is a threat to the frosted flatwoods salamander when isolated, seasonally ponded wetland breeding sites are changed to or connected to more permanent wetlands inhabited by fish species not typically found in temporary ponds. Red imported fire ants ( Solenopsis invicta ) are also potential predators of flatwoods salamanders, especially in disturbed areas. They have been seen in areas disturbed by the installation of drift fences at known frosted flatwoods salamander breeding sites (Palis 2008). Mortality of amphibians trapped at drift fences has occurred when fire ants were present and traps were not monitored with sufficient frequency (Palis et al. 2002, p. 6). The severity and magnitude of effects, as well as the long-term effect, of fire ants on frosted flatwoods salamander populations are currently unknown. In summary, diseases of amphibians in the southeastern United States remain largely unstudied. However, given the incidence of disease in species in the western United States that could be considered surrogates for flatwoods salamanders, the probability exists for similar infections to occur in frosted flatwoods salamander populations. We consider this to be a potential threat of moderate magnitude. Predation by fish is a historic threat that continues to be a localized problem when ditches, firebreaks, or vehicle ruts provide connections allowing the movement of fish from permanent water bodies into frosted flatwoods salamander breeding sites. Fire ants also have the potential of being a localized threat, particularly in disturbed areas. We consider these threats to be potential threats of moderate magnitude because 88 percent of frosted flatwoods salamander populations occur primarily on public lands where they are relatively protected from habitat destruction. D. The Inadequacy of Existing Regulatory Mechanisms Other than the National Forest Management Act and the Sikes Act, there are no existing regulatory mechanisms for the protection of the upland habitats where frosted flatwoods salamanders spend most of their lives. Refer to Factor D under “Summary of Factors Affecting the Species (Reticulated Flatwoods Salamander)” for information on the threat of the Inadequacy of Existing Regulatory Mechanisms that also applies to frosted flatwoods salamander. Longleaf pine habitat management plans that provide conservation benefits to frosted flatwoods salamanders have been written for most of these sites. They include management plans for State- and Federally-owned lands and integrated natural resource management plans (INRMPs) for Department of Defense lands. Most of the plans contain specific goals and objectives regarding habitat management, including prescribed burning, that would benefit frosted flatwoods salamanders if implemented. Multiple-use is the guiding principle on most of these public lands, however, and protection of the frosted flatwoods salamander may be just one of many management goals including timber production and military and recreational use. At the State and local levels, regulatory mechanisms are limited. The flatwoods salamander is listed as a threatened species in the State of Georgia (Jensen 1999, pp. 92-93). This designation protects the species by preventing its sale, purchase, or possession in Georgia and by prohibiting actions that cause direct mortality of the species or the destruction of its habitat on lands owned by the State of Georgia (Ozier 2008). However, there are no known frosted flatwoods salamander populations on lands owned by the State of Georgia. In 2001, the Florida Fish and Wildlife Conservation Commission (FFWCC) listed the flatwoods salamander (which includes the frosted flatwoods salamander) as a species of special concern (FFWCC 2007, p. 2) and prohibited direct take except through permit. As part of the listing process, a Statewide management plan was developed for the salamander in Florida (FFWCC 2001, p. 1-60); however, Florida regulations offer no protection against the most significant threat to the frosted flatwoods salamander—loss of habitat. In South Carolina, the flatwoods salamander is listed as endangered (South Carolina Department of Natural Resources 2008a). Prohibitions extend only to the direct take of the flatwoods salamander (South Carolina Department of Natural Resources 2008b). These regulations offer no protection against the most significant threat to the flatwoods salamander, which is loss of its habitat. In summary, although existing regulatory mechanisms provide little direct protection of frosted flatwoods salamanders (beyond the protections afforded by the Act), they do provide a degree of protection for the remaining occupied habitat, primarily on public lands. The record of management on public lands since the original listing of the flatwoods salamander in 1999 indicates that public agencies are actively pursuing longleaf pine ecosystem management programs that benefit the frosted flatwoods salamander. Frosted flatwoods salamander breeding sites on the three private land sites may, in some cases, come under the jurisdiction of the Corps (Refer to Factor D under “Summary of Factors Affecting the Species (Reticulated Flatwoods Salamander)” for discussion of section 404 of the Clean Water Act and regulatory wetlands), but most likely they are provided little regulatory protection. We have determined that the threat of inadequate existing regulatory mechanisms is primarily an ongoing threat of moderate magnitude. E. Other Natural or Manmade Factors Affecting Its Continued Existence Metapopulations are important to the long-term survival of temporary pond breeding amphibians. Refer to Factor E under “Summary of Factors Affecting the Species (Reticulated Flatwoods Salamander)” for additional information on metapopulations. Of the 25 known frosted flatwoods salamanders populations, 16 (64 percent) are supported by more than one breeding pond and may be considered metapopulations. However, 36 percent (9 out of 25) of the known frosted flatwoods salamander populations that have only a single breeding pond, any one of the many threats that may render a breeding pond unsuitable could cause the extirpation of the affected population. Invasive plant species, such as cogongrass ( Imperata cylindrica ), threaten to further degrade existing flatwoods habitat. Refer to Factor E under “Summary of Factors Affecting the Species (Reticulated Flatwoods Salamander)” for additional information on invasive species and the threat they represent, which also applies to the frosted flatwoods salamander. Frosted flatwoods salamander habitat management plans will need to address threats posed by cogongrass and other invasive plant species and include strategies to control them. Pesticides (including herbicides) may pose a threat to amphibians, such as the frosted flatwoods salamander. Refer to Factor E under “Summary of Factors Affecting the Species (Reticulated Flatwoods Salamander)” for additional information on pesticides and the threat they represent, which also applies to the frosted flatwoods salamander. However, herbicides may be a necessary tool to reduce or eliminate woody vegetation or invasive plants when the use of prescribed fire is not possible or effective (Jensen 2007, Wigley 2008). Nevertheless, pesticides should never be used in flatwoods salamander habitat unless no other habitat management tool is available; herbicide label directions should be followed closely and aerial spraying should not be used as an application technique. Under these conditions, we consider this threat to be of moderate magnitude. Studies of other ambystomatid species have demonstrated a decline in larval survival in the presence of predatory fish, as mentioned above under Factor C. One of the potential reasons for this decline may be the negative effect resulting from these fish competing with salamander larvae for invertebrate prey. The invertebrates found by Whiles et al. (2004, p. 212) in a study of larval frosted and reticulated flatwoods salamander gut contents are typical of freshwater habitats in the Southeast that do not contain predatory fish on a regular basis. The presence of predatory fish has a marked effect on invertebrate communities and alters prey availability for larval salamanders with the potential for negative effects on larval fitness and survival (Semlitsch 1987, p. 481). Wherever connections have been created between permanent water and frosted flatwoods salamander ponds, such as through installation of firebreaks or ditches, this threat from predatory fish exists. Studies of frosted flatwoods salamander populations, since the original species listing of flatwoods salamander as threatened (64 FR 15691; April 1, 1999), have been limited due to drought. Data on the numbers of adults within existing populations do not exist. However, given the low number of individuals encountered even when breeding is verified, populations are likely to be very small at any given breeding site. Small populations are at increased threat of extirpation from natural processes (genetic isolation, inbreeding depression, and drought), as well as the manmade threats described above. In summary, a variety of natural or manmade factors historically or currently threaten, or have the potential to threaten, the frosted flatwoods salamander. The loss of metapopulation structure in the distribution of frosted flatwoods salamander populations was a range-wide threat that caused historic losses of this species. It continues to be a current threat for 64 percent of the remaining frosted flatwoods salamander populations. Fire suppression and inadequate habitat management continue to cause the degradation of occupied sites, primarily on private land. Invasive plant species probably did not have much of a historic impact on salamander populations, but they are a range-wide current threat, and they are likely to become more widespread and difficult to control. Range-wide, low population densities have been a historic threat and continue to be a threat for most frosted flatwoods salamander populations, particularly due to past and current drought conditions, habitat loss, population fragmentation, and periodic reproductive failures that occur naturally in pond-breeding amphibians. The impact that competing predators may have on the salamanders' prey base, and the threat of pesticide and herbicide use, are less clear as historic threats but remain potential localized threats for the species. Therefore, while we have determined that other natural and manmade factors, such as invasive species, pesticides, and competition for the species' prey base, may threaten the frosted flatwoods salamander, the severity and magnitude of these threats are not currently known. Acting in combination with threats listed above under Factors A through D, the threats under Factor E could increase the severity of the other threats. Determination We have carefully assessed the best scientific and commercial information available regarding the past, present, and future threats to the frosted flatwoods salamander. In summary, the most significant historical threat to the frosted flatwoods salamander, as listed in Factor A (above), is loss of the majority of its habitat. A variety of localized threats (described under Factors A, C, D, and E) have the potential to impact the remaining frosted flatwoods salamander populations and habitat. These include—alterations in the hydrology of existing wetland breeding sites (including “ditching” which results in the introduction of predatory fish); incompatible forest management; ORV use; fire suppression; drought; and disease. The severity and magnitude of the threats under Factor E are not currently known. Nevertheless, we have determined that threats under this factor will exacerbate the effects of threats due to habitat loss and drought. As described in Factor E above, small populations are at increased threat of extirpation from natural processes (genetic isolation, inbreeding depression, and drought), as well as the manmade threats listed above. Finally, there are potential localized threats from fire ants, pesticides, and invasive plants for which the extent of impact is yet undeterminable, but that we believe are legitimate threats due to both their impact on surrogate species and their prevalence in the types of habitats used by this species. Only 25 frosted flatwoods salamander populations are known. Ten (40 percent) of these populations are supported by only one breeding site. A population with only one breeding site has a tenuous future just given randomly varying environmental factors without considering the additional threats of habitat destruction and degradation that further threaten these populations. As noted previously, habitat with the range of the frosted flatwoods salamander is currently experiencing drought conditions. Palis et al. (2006, pp. 5-6) studied a frosted flatwoods population in Florida during a drought from 1999-2002. This study documented 3 consecutive years of reproductive failure and a steady decline in adult immigration to the site for breeding as the drought progressed. Catastrophic reproductive failure occurs even in healthy populations of pond-breeding amphibians. When it does occur, the modeling efforts of Taylor et al. (2005, p. 796) showed that each year of reproductive failure raises the threshold of survival required to achieve persistence and imposes the possibility of extirpation even under otherwise favorable environmental conditions. Taylor et al. (2005, p. 799) reminds us that particularly with small populations or low population growth rates (as exists with the frosted flatwoods salamander) effects of reproductive failure are made worse by demographic stochasticity. Even in populations with multiple breeding ponds, amphibian populations may be unable to recolonize areas after local extirpations due to their physiological constraints, relatively low mobility, and site fidelity (Blaustein et al. 1994, pp. 60, 67-68). For frosted flatwoods salamander, 40 percent of populations have only one breeding pond. If the habitat at that site is destroyed, recolonization would be impossible and the population supported by that breeding pond would be extirpated. Habitat loss on private lands is an imminent threat that is compounded by a variety of other factors. Fire suppression on private lands occupied by the frosted flatwoods salamander represents one of the biggest threats to the species' habitat and the continued existence of the species on these sites. However, 60 percent of frosted flatwoods salamander populations have an improved chance of surviving demographic and environmental stochasticity given that the distribution of breeding sites occurs within an adult salamander's dispersal distance. We believe that, when combining the effects of historical, current, and projected habitat loss and degradation, historical and ongoing drought, and the exacerbating effects of disease, predation, small population size, and isolation, the frosted flatwoods salamander continues to be likely to become an endangered species throughout all of its range within the foreseeable future. We believe these threats, particularly the threats to populations resulting from habitat degradation and fragmentation, small population size, and drought, are current and are projected to continue into the future. We have determined that these threats are operating on the species and its habitat with a moderate degree of magnitude throughout most of its range and with a moderate degree of severity, as discussed above. Based on the best available scientific and commercial information, we have determined that the preferred action is for the frosted flatwoods salamander to retain its status as a threatened species under the Act. Without the protection of the Act, significant management of threats would likely occur on public lands; however, there is still substantial risk of loss of ponds to drought and disease and, on private lands, a variety of potential threats (for example, introduction of fish, predation, pesticides), and imminent threats (for example, fire suppression, invasive species, and development). As discussed previously, declines resulting from drought can occur within only a few years. In the case of the frosted flatwoods salamander, 40 percent of populations have only one breeding pond. If the habitat at that site is destroyed, recolonization would be impossible and the population supported by that breeding pond would be extirpated. This could occur within a few years given recurring drought conditions and existing threats. While not in immediate danger of extinction, the frosted flatwoods salamander is likely to become an endangered species in the foreseeable future throughout all or a significant portion of its range if the present trends that negatively affect the species, and its limited and restricted habitat, continue. Furthermore, because these threats to the species are of comparable magnitude and severity across all of the species' range, we have determined that an analysis of whether a specific portion of the range might require a different listing status is not warranted at this time. Available Conservation Measures Conservation measures provided to species listed as endangered or threatened under the Act include recognition of the status, increased priority for research and conservation funding, recovery actions, requirements for Federal protection, and prohibitions against certain activities. Recognition through listing results in public awareness and conservation actions by Federal, State, and local agencies; private organizations; and individuals. The Act provides for possible land acquisition and cooperation with the States, and for conservation actions to be carried out for listed species. Section 7(a) of the Act, as amended, requires Federal agencies to evaluate their actions with respect to any species that is listed as endangered or threatened and with respect to its critical habitat, if any is being designated. Regulations implementing this interagency cooperation provision of the Act are codified at 50 CFR part 402. Section 7(a)(2) of the Act requires Federal agencies to ensure that activities they authorize, fund, or carry out are not likely to jeopardize the continued existence of a listed species or result in destruction or adverse modification of critical habitat. If a Federal action may affect a listed species or its critical habitat, the responsible Federal agency must consult with us under the provisions of section 7(a)(2) of the Act. Federal agency actions within the species habitat that may require consultation as described in the preceding paragraph include management and any other landscape altering activities on Federal lands administered by the Department of Defense, Fish and Wildlife Service, and U.S. Forest Service; issuance of section 404 Clean Water Act permits by the Corps; construction and management of gas pipeline and power line rights-of-way by the Federal Energy Regulatory Commission; and construction and maintenance of roads or highways by the Federal Highway Administration. The Act and its implementing regulations set forth a series of general prohibitions and exceptions that apply to all endangered wildlife. The prohibitions, codified at 50 CFR 17.21 for endangered wildlife, in part, make it illegal for any person subject to the jurisdiction of the United States to take (includes harass, harm, pursue, hunt, shoot, wound, kill, trap, capture, or collect; or to attempt any of these), import, export, ship in interstate commerce in the course of commercial activity, or sell or offer for sale in interstate or foreign commerce any listed species. It is also illegal to possess, sell, deliver, carry, transport, or ship any such wildlife that has been taken illegally. Certain exceptions apply to agents of the Service and State conservation agencies. We may issue permits to carry out otherwise prohibited activities involving threatened or endangered wildlife species under certain circumstances. Regulations governing permits are codified at 50 CFR 17.22 for endangered species. You may obtain permits for scientific purposes, to enhance the propagation or survival of the species, and for incidental take in connection with otherwise lawful activities. Critical Habitat Previous Federal Actions For information about previous Federal actions regarding designation of critical habitat for flatwoods salamanders, see our proposed rule (73 FR 47258) published on August 13, 2008. This notice included revisions to the proposed designation of critical habitat published in the Federal Register on February 7, 2007 (72 FR 5856), and announced the availability of our draft economic analysis of the proposed critical habitat designation (DEA). On September 18, 2008, we published a notice in the Federal Register (73 FR 54125) providing supplemental information on the status of the frosted flatwoods salamander. On October 8, 2008, we published a notice in the Federal Register which extended the public comment period on the proposed rule and provided the time, date, and location of our public hearing (73 FR 58922). We held a public hearing on October 22, 2008. The extended public comment period ended on November 3, 2008. Summary of Comments and Recommendations As stated above, since the proposed rule addressed both listing and critical habitat, comments received combined these two issues. Therefore, we are presenting the combined comments and responses for these issues, below. In the 2007 proposed rule, we requested written comments from the public on reasons why we should or should not designate critical habitat for the flatwoods salamander (72 FR 5856). We contacted appropriate Federal, State, and local agencies; scientific organizations; and other interested parties, and invited them to comment on the proposed rule. We also issued press releases and published legal notices in the Jasper County Sun, Pensacola News Journal, The DeFuniak Springs Herald-Breeze, Savannah Morning News, Tallahassee Democrat, The Albany Herald, Miller County Liberal, The Berkeley Independent, The Florida Times-Union, The News Herald , and The Post and Courier newspapers. During the open public comment period, we received a request to hold a public hearing, however a public hearing was not held at that time. Due to new information that became available on threats to the flatwoods salamander and the reclassification in the taxonomy of the species, we asked for an extension of our court-ordered deadline on the designation of critical habitat for the flatwoods salamander to include the new information. Subsequently, a new proposed rule was written and published in the Federal Register on August 13, 2008 (73 FR 4725). For the 2008 proposed rule, we requested written comments from the public on known or suspected threats to the frosted flatwoods salamander and the reticulated flatwoods salamander and any information on the need to change the status of either species; reasons why we should or should not designate critical habitat for the two species; and on the DEA (73 FR 47258). We contacted appropriate Federal, State, and local agencies; scientific organizations; and other interested parties, and invited them to comment on the proposed rule. We also issued press releases and published legal notices in the Jasper County Sun, Northwest Florida Daily News, Pensacola News Journal, Savannah Morning News, Tallahassee Democrat, The Albany Herald, The Berkeley Independent, The Florida Times-Union, The News Herald , and The Post and Courier newspapers. Based on a request received during the public comment period, we held a public hearing and information meeting on October 22, 2008, at Pensacola Junior College in Pensacola, Florida. During the comment period for the first proposed rule that opened on February 7, 2007, and closed on April 9, 2007, we received 23 comments directly addressing the original proposed critical habitat designation: five from peer reviewers; three from Federal agencies; three from State agencies; and 12 from organizations or individuals. During the comment period for the second proposed rule that opened on August 13, 2008, and closed on November 3, 2008, we received 79 comments directly addressing the reclassification in the listing of the flatwoods salamander into two species; the proposed designation of the reticulated flatwoods salamander as endangered; the maintenance of the listing of the frosted flatwoods salamander as threatened; the revised proposed critical habitat designation for the two species; and the DEA. Of these latter comments, 44 comments were received either in written form or through the portal at: http://www.regulations.gov ; three of these were from Federal agencies, none were from State agencies, one was from a local government, and 40 were from organizations or individuals. Thirty-five of the 79 comments were made during the public hearing held on October 22, 2008; one of these was from a Federal agency, one was from a State agency, one was from a state senator, four were from local governments, and 28 were from organizations or individuals. The following summary statistics are provided on the 23 comments received on the 2007 proposed rule. In total, 7 commenters supported the designation of critical habitat for the flatwoods salamander, 3 opposed the designation, and 13 were neutral regarding the designation. These following summary statistics are provided on the 79 comments received on the 2008 proposed rule. Nine commenters sent comments during the 2008 open comment period and also commented at the public hearing. An individual, group, or agency responding multiple times was only counted once as none of these commenters' opinions of the proposed rule differed between responses. In total, 33 commenters supported the proposed rule, 34 opposed the proposed rule, and 3 were neutral regarding the proposed rule. Comments received were grouped into 7 general issues specifically relating to the subjects in the 2008 proposed rule and the DEA, and are addressed in the following summary. We have incorporated comments into this final rule as appropriate. Peer Review In accordance with our policy published on July 1, 1994 (59 FR 34270), and current Department of the Interior guidance, we solicited expert opinions for both the 2007 and 2008 proposed rules from five knowledgeable individuals with scientific expertise that included familiarity with the species, the geographic region in which the species occurs, and conservation biology principles. We received responses from all five of the peer reviewers on the 2007 proposed rule and from four of the five peer reviewers on the 2008 proposed rule. We reviewed all comments received from the peer reviewers for substantive issues and new information regarding flatwoods salamander critical habitat. We combined peer reviewer comments from both years. The peer reviewers generally concurred with our methods and conclusions and provided additional information, clarifications, and suggestions to improve the final critical habitat rule. Peer reviewer comments are addressed in the following summary and incorporated into the final rule as appropriate. Peer Review Comments (1) Comment: Three of the peer reviewers emphasized the importance of the Eglin Air Force Base-Hurlburt Field metapopulation to the survival of the reticulated flatwoods salamander and questioned whether adequate habitat management, especially fire management, could be conducted if the highway proposed for the area was approved ( see also Comment 15). They discussed the inclusion or exclusion of military lands which have approved Integrated Natural Resource Management Plans (INRMPs), including these two sites as well as Navy Outlying Landing Field (NOLF) Holley (see also Comment 16). These peer reviewers were concerned about the finite period of the plans and the potential for decreased conservation efforts if INRMPs are revised when renewed. One peer reviewer recommended that NOLF Holley be included in critical habitat because the Navy's natural resources manager and forester had informed him that the Navy no longer had use for the field and that Santa Rosa County was interested in acquiring it. They concurred with the 2008 proposed rule (73 FR 47258) that included these military lands in proposed critical habitat. Our Response: Eglin Air Force Base (Eglin) has assured the Service that they “will not allow negative impacts to the salamander habitat” on the base (DoAF 2008a, p. 1). The Commander of Eglin stated that, “Eglin will ensure that the proposed Bypass road, and any actions associated with it, will not prevent implementation of the conservation measures identified in the INRMP for the flatwoods salamander” (DoAF 2008b, p. 1). The Service has reassessed the Eglin INRMP and determined that, with the Air Force's recent assurances, it will provide a conservation benefit for the reticulated flatwoods salamander. The Department of the Navy has assured the Service that the Navy has no intention of transferring ownership of NOLF Holley and the INRMP will continue to be implemented for this site as written (Department of the Navy 2008, p. 2). We conduct annual reviews of the INRMPs for all the military bases with known flatwoods salamander populations and reassess their conservation benefits and implementation. All the involved military bases have assured the Service of their future compliance with their INRMPs (see citations above). As a result of this analysis, Eglin, Hurlburt Field, and NOLF Holley have been removed from the final critical habitat designation for the reticulated flatwoods salamander. (2) Comment: One peer reviewer stated that the locality record used as the basis for proposed critical habitat unit RFS-5 is based on a poor quality photograph of a single larva collected in 1998 and that the larva in the photograph is likely a mole salamander ( Ambystoma talpoideum ). The site of the locality record and at least 100 other wetlands in the area have been surveyed since 2002 during suitable immigration and emigration periods. Many mole salamanders have been captured, but no reticulated flatwoods salamanders. It is the opinion of the peer reviewer that the original identification of the collected larva as a reticulated flatwoods salamander was in error. Our Response: We agree that there is no verifiable evidence that flatwoods salamanders occupy habitat represented by Unit RFS-5, as originally proposed. Therefore, this unit has been removed and the final critical habitat designation for the reticulated flatwoods salamander has been revised based on this comment. (3) Comment: One peer reviewer stated that habitat within proposed critical habitat unit FFS-2, located on Tate's Hell State Forest, is no longer suitable for the frosted flatwoods salamander. Since an adult flatwoods salamander was discovered there in 1998, hydrological restoration of the likely breeding site has been conducted and altered the site to a more permanently flooded wetland. Surveyors sampled the site in 2002, 2003, and 2004 but were unsuccessful in documenting any flatwoods salamander larvae within the wetland. The peer reviewer believes the wetland restoration project and the historically poor upland management of the area have resulted in the loss of flatwoods salamander habitat at this site on Tate's Hell State Forest. Our Response: Based on the peer reviewer's comment and the site visit information, we believe Unit FFS-10, as originally proposed, no longer contains the PCEs essential to the conservation of the flatwoods salamander. Therefore, this unit has been removed from the final critical habitat designation. (4) Comment: One peer reviewer stated that maps in the proposed rule are not sufficient for delineating actual boundaries of proposed critical habitat. The peer reviewer suggested using road or topographic maps and aerial photography. Our Response: The printing standards of the Federal Register are not compatible with using road or topographic maps and aerial photography. We constructed the critical habitat units using a Geographic Information System (GIS). The resulting critical habitat GIS shapefiles are available by request from the Mississippi Field Office (see FOR FURTHER INFORMATION CONTACT ). The shapefiles can be laid over other layers (aerial photography, roads) to get more precise locality information. (5) Comment: One peer reviewer stated that reticulated flatwoods salamander units in southwest Georgia (unit names in 2008 proposed rule (73 FR 47258; August 13, 2008) are RFS-10, subunits A and B, respectively) may have agricultural land that does not contain the primary constituent elements and should be removed. Our Response: The peer reviewer did not have access to our GIS data when we received this comment. When constructing the units in question, we used aerial photography to verify the presence of the primary constituent elements on the areas and that all agricultural land was excluded from RFS-10, subunits A and B. Public Comments General Biological Comments (6) Comment: One commenter cited studies which described flatwoods salamander breeding sites as roadside ditches and borrow pits, as well as natural habitats. This commenter believed that this shows the adaptability of the species and the likelihood that suitable breeding habitat could be created for the salamanders. Several commenters during the public hearing thought that flatwoods salamander habitat could be relocated or constructed elsewhere as an alternative to protecting the existing occupied sites through critical habitat designation. Other commenters at the public hearing stated that this was not possible, as flatwoods salamanders are tied to specific soils and forest-wetland types that need to be present in a landscape context. These commenters expressed support for protecting existing sites. Our Response: Flatwoods salamanders are known to breed in wetlands that dry on a seasonal basis. The Service is aware of records of flatwoods salamander larvae occurring in ditches and borrow pits. However, whether larvae were successful in developing into adult salamanders at these sites is unknown. The ponded breeding sites must hold water long enough and have a sufficient food source to allow salamander development and metamorphosis. They must also be free of predaceous fish and toxic substances. In addition, there are a number of biotic and abiotic factors that are likely essential for flatwoods salamanders at breeding sites that are currently unknown. Experimental relocations should be an action of last resort for these species and may be explored through the recovery process, if deemed necessary. Adequacy and Extent of Critical Habitat (7) Comment: Two commenters stated that critical habitat designation on any lands approved under the Sustainable Forestry Initiative and Sustainable Forestry Certification Program (SFI) is unnecessary and redundant (not warranted). These lands are already recognized as habitat for listed species under the certification program and participants in the program are required to safeguard and protect threatened and endangered species. Participants are expected to implement scientifically based management practices and adaptive management strategies as appropriate. Provisions of this program are not legally binding; however, participants must comply to stay in the program. Therefore, lands under SFI programs should not require special management considerations. The commenters believed designation would not significantly increase or contribute to the likelihood of recovery of the species because the vast majority of lands are either in public ownership or managed according to SFI standards. Therefore, the commenters asserted that critical habitat offers little or no additional management protection and no additional conservation benefit. Our Response: The criteria for designating sites as critical habitat are whether the sites provide the features essential for the conservation of the species and whether those features may require special management consideration or protection. Under section 4(b)(2) of the Act, the Secretary may exclude an area from critical habitat if the benefits of such exclusion outweigh the benefits of specifying such areas as part of critical habitat. We also consider whether landowners having proposed critical habitat on their lands have developed any conservation plans for the area, or whether there are conservation partnerships that would be encouraged by designation of, or exclusion from, critical habitat. Included in this analysis would be whether or not conservation plans have species-specific management prescriptions, or other management approaches, that are coupled with assurances of implementation. The commenter presented a general statement about SFI programs. However, the Service did not receive any comments from specific private landowners within proposed critical habitat that identified themselves as participants in SFI programs nor did we receive any SFI conservation plans for analysis. Therefore, there is no new information indicating that removal of lands under SFI from critical habitat is warranted. (8) Comment: Several commenters stated that private land should be excluded from critical habitat designation. One commenter suggested that the Service should offer incentives, such as voluntary cooperative agreements as a conservation policy for private lands in lieu of critical habitat designation. These commenters stated that it would not be beneficial to flatwoods salamanders to designate critical habitat on private land since designation would be a disincentive for the landowners to continue conservation efforts for this species, would likely increase threats resulting in a net loss of conservation benefit, and eventually result in the extirpation of flatwoods salamanders on private lands. Our Response: Section 4 of the Act requires the Service to use the best available scientific data in designating critical habitat. Private lands are not exempted from this analysis. Flatwoods salamanders have been listed since 1999 and protection from “take” under section 9 has been in effect since that time. The Service knows of no situation where a private landowner has knowingly destroyed or mismanaged flatwoods salamander habitat as a result of this listing. Critical habitat only applies to those lands where there is a Federal nexus (a connection or link to the Federal government). In some cases, private lands may be affected if the landowner is undertaking a project that requires Federal funding or permit. However, the Service believes most application of the protection provided by critical habitat will not affect private lands. Programs are available to provide funds to private landowners for managing habitat for listed species, as well as permits that can be obtained to protect private landowners from the take prohibition when such taking is incidental to, and not the purpose of, the carrying out of an otherwise lawful activity. Private landowners may contact their local Service field office to obtain information about these programs and permits. (9) Comment: Several commenters expressed concern about the potential for being prosecuted for adverse modification if private properties designated as critical habitat are fire suppressed. The commenter requested a definition for fire suppression and an explanation of practices related to fire suppression that would be problematic. The commenter requested that the Service offer emergency exemptions from adverse modification for human life or property. Our Response: At this time, the Service is unaware of any Federal actions that would leave a private landowner vulnerable to prosecution for adverse modification due to fire suppression. Federal actions related to fire suppression that might lead to adverse modification would include improper implementation of management plans on Federal lands. If suppression of a wildfire is necessary as an emergency Federal action relating to human health and safety within occupied habitat of a listed species or designated as critical habitat, an after-the-fact consultation can be conducted. Under the statutory provisions of the Act, we determine destruction or adverse modification on the basis of whether, with implementation of the proposed Federal action, the affected critical habitat would remain functional to serve its intended conservation role for the species. However, when considering fire suppression as a threat, we refer to a Federal action which will lead to elimination of fire as a management tool and allow thick underbrush and mid-story to shade out the herbaceous ground cover. Fire suppression, in this sense, leads to deterioration of flatwoods salamander habitat quality and potentially adverse modification of critical habitat. (10) Comment: One commenter requested that the Service should consider a buffer width less than 1,475 feet (ft) (450 meters (m)) around known breeding ponds when defining and designating critical habitat units and stated that the designation of this distance was arbitrary because it was based on a different salamander species. The commenter suggested a 534 [sic] ft (164 m) buffer width as calculated by Semlitsch (1998, p. 1113). This commenter also references Palis et al. (2002, pp. 1-20) that is provided as support for a smaller buffer width around known flatwoods salamander breeding ponds. Another commenter disputed the scientific basis for rounding up the buffer radius to 1,500 ft (457 m) from 1,476 ft (450 m) when constructing critical habitat units. Several commenters requested that the buffer width used in calculating critical habitat units be increased to 5,576 ft (1,700 m), since this is the maximum distance flatwoods salamanders have been reported to disperse and this would create connectivity between known occurrences. Our Response: Semlitsch (1998, p. 1113) combined movement data in five States for six species of ambystomatid salamanders, which had been collected over a period of several decades. Using these data, we generated a 538-ft (164-m) buffer width from a wetland's edge into the terrestrial habitat, which would create an area that he stated would encompass 95 percent of a population of one of these species. However, Semlitsch (1998, p. 1117) pointed out that the values used in this calculation probably underestimate the actual buffer needed for some species of salamanders. In addition, he specifically mentioned the flatwoods salamander as one of the species that may require more habitat than the area created by using the 538-ft (164-m) buffer width (Semlitsch 1998, p. 1117). The Service selected a buffer width of 1,475 ft (450 m) from the Semlitsch (1998, p. 1115) paper to use for the flatwoods salamander calculation. This is the maximum value used in his calculations for the marbled salamander. We chose this species because it was recommended by researchers as a model for the flatwoods salamander due to its similar life history (Taylor et al. 2005, pp. 792) and because it uses habitat in a similar way. Although adult marbled salamanders occupy hardwood forests rather than pine forests, they breed underground and in temporary ponds. The 1,475 ft (450 m) value corresponded well with data collected over 20 years by Means et al. (1996, p. 435) which described estimated movements of flatwoods salamanders of 984 ft (300 m) to 1,640 ft (500 m) between upland habitat and breeding ponds in relatively homogeneous habitat. The Service used the value of 1,476 ft (450 m) to estimate the size of activity areas used by flatwoods salamanders in the original listing rule. This value also was used originally in draft management guidelines for flatwoods salamanders that the Service wrote in conjunction with the flatwoods salamander recovery team. During review of these draft guidelines, several members of the rec [Text truncated at 120,000 characters. The full text is on the page linked above.] --- Source: Frix Law Library, https://www.frixlaw.com/law-library/documents/fr%3AE9-2403. Public record. Not legal advice.