Species Status Assessment for the Striped Newt (Notophthalmus perstriatus) Version 1.0

Striped newt eft. Photo credit Ryan Means (used with permission).

May 2018 U.S. Fish and Wildlife Service Region 4 Jacksonville, Florida

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Acknowledgements This document was prepared by the U.S. Fish and Wildlife Service’s North Florida Field Office with assistance from the Georgia Field Office, and the striped newt Species Status Assessment Team (Sabrina West (USFWS-Region 8), Kaye London (USFWS-Region 4) Christopher Coppola (USFWS-Region 4), and Lourdes Mena (USFWS-Region 4)). Additionally, valuable peer reviews of a draft of this document were provided by Lora Smith (Jones Ecological Research Center) , Dirk Stevenson (Altamaha Consulting), Dr. Eric Hoffman (University of Central Florida), Dr. Susan Walls (USGS), and other partners, including members of the Striped Newt Working Group. We appreciate their comments, which resulted in a more robust status assessment and final report.

EXECUTIVE SUMMARY

This Species Status Assessment (SSA) is an in-depth review of the striped newt's (Notophthalmus perstriatus) biology and threats, an evaluation of its biological status, and an assessment of the resources and conditions needed to maintain species viability. We begin the SSA with an understanding of the species’ unique life history, and from that we evaluate the biological requirements of individuals, populations, and species using the principles of population resiliency, species redundancy, and species representation. All three concepts (or analogous ones) apply at both the population and species levels, and are explained that way below for simplicity and clarity as we introduce them.

The striped newt is a small salamander that uses ephemeral wetlands and the upland habitat (scrub, mesic flatwoods, and sandhills) that surrounds those wetlands. The striped newt currently extends from southern Georgia to north-central Florida. Striped newts have been divided in two regions which have been identified as having ecological differentiation due to precipitation and temperature differences. The species has been a candidate for listing under the Endangered Species Act of 1973, as amended (Act), since the 12-month finding published in 2011 (USFWS 2011).

The striped newt is one of three distinct Notophthalmus species found in North America. Striped newts are a distinct species that is more similar to and phylogenetically more closely related to the blackspotted newt (N. meridionalis) than it is to the eastern red spotted newt (N. viridescens). Striped newts have a lifespan of 12–15 years (Dodd Jr. 1993; de Magalhaes and Costa, 2009) and use aquatic and terrestrial habitats during their complex life cycle. Although they have been known to live up to 17–18 years in captivity (Wallace et al. 2009, pp. 138–139; Mark Beshel, 2018, pers. comm.) we based our analysis on the known life span from studies in the field (Dodd Jr. 1993). During the fall/winter and spring seasons and after rain events, striped newt adults migrate to isolated ephemeral wetlands that lack large predatory fish. Striped newts mate and the female deposits eggs into aquatic vegetation, and when the larvae reach maturity, they metamorphose into a gilled aquatic adult (paedomorph) that will remain in the aquatic phase or metamorphose into an immature terrestrial adult (eft). These terrestrial adults will ultimately move to the nearby uplands.

Striped newts are currently represented throughout their historical range. They were identified in

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79 breeding ponds between 1980 and 1999 and are currently identified as extant in 116 breeding ponds (Table 2). Striped newts have been identified as possibly extirpated from 57 breeding ponds and extirpated from 17 (Farmer et al. 2017). In order to maintain viability, the species should be represented by populations distributed across its range and in a variety of habitats so that there are always some populations experiencing environmental conditions that support some level of reproductive success. Currently, there are 10 known extant populations of striped newts in the Western Region and 106 in the Eastern Region of the species.

Since the early 2000s, the distribution and abundance of striped newts have not significantly changed; the spatial extent of their distribution is 95% of its historical extent. In Florida, an extensive survey effort that targeted potential suitable habitat for the species identified a large number of previously unknown breeding ponds. In Georgia, the species is still known from 5 locations with a total of 11 breeding ponds believed to be extant. Although we do not know the status of each individual population that is using each breeding pond, analyzing the current grid of occurrences suggest that populations are persisting, especially where metapopulations (multiple breeding ponds within 1 kilometer (km) of each other) occur. The extent to which stressors are impacting the species abundance is unknown, but it is likely that back to back long- term droughts would negatively impact their abundance.

Long-term drought, development, and land use changes have been historically identified as the primary causes of population loss for the striped newt. During long-term dry conditions, striped newts lose breeding habitat and their populations decline; however, studies have shown that they are able to bounce back after prolonged droughts (Dodd and Johnson 2007). Local extirpations are one of the possible outcomes of a long-term drought, as well as migration to the uplands until better breeding habitat conditions occur or they can migrate to other temporary wetlands nearby to breed (Dodd Jr. 1993).

Habitat loss due to land use changes and development is another stressor that continues to impact the species across its range. Although approximately 85% of striped newt populations are protected in conservation lands, some of the remaining populations are in private lands without any conservation. Fire suppression of the occupied habitat is also identified as a stressor of concern for the species. Lack of fire can result in the succession of natural pine forests and scrub into forests dominated by hardwoods and closed canopies with less vegetative ground cover. There is a high degree of uncertainty on how fire suppression is impacting striped newt.

We developed three future risk scenarios: 1) Conditions stay the same as currently with improvements in translocation efforts across the range of the species, low levels of climate change, and improved management on conserved lands; 2) impacts from stressors continue current trends and conservation actions continue only at Apalachicola National Forest (ANF); and 3) the effects from land use change, climate change, and fire suppression increase while repatriation efforts decrease. We focused our analysis on a 40–50 year timeframe.

Our analyses indicate that the resiliency, representation, and redundancy of the striped newt have not changed in the last 30 years. Historically, striped newts were identified in fifty-six breeding ponds (populations) in Florida and five locations in Georgia (USFWS 2011). Abundance of newts in each population is unknown and presence of newts in a breeding pond is what is used to

3 determine if a population is extant. Current populations are spread across the species' range and varying environmental conditions. Due to the large number of populations within the two ecoregions, the risk of losing large numbers of populations and thus species representation in one of the regions due to catastrophic events is unlikely.

We have determined that the habitat and population loss attributable to development and sea level rise is not likely to significantly reduce existing populations of striped newt. The vast majority of known populations of striped newt occupy conservation lands that are managed at varying degrees with prescribed fire as a management tool. Habitat loss and sea level rise will very likely impact less than 25% of the currently known suitable striped newt habitat and less than 10 of the known populations. We have moderate to low confidence in the known current condition of populations across the range, but predictions under the different scenarios show that all but five of the properties with extant populations will likely be extirpated in scenario 3; two in the Western Region and three in the Eastern Region. The species has populations distributed across all of its historical range with a lower representation in the Western Region of the species but with more breeding ponds being identified and repatriation efforts underway at ANF, Florida. Drought is expected to play a significant role as a stressor for this species but we are unclear how the species will adapt to the prolonged droughts or how resilient it is. Habitat development and sea level rise projections for the next 43 years will inundate one of the populations at Guana River WMA and may impacts a few others with salt intrusion. We have a low confidence in the likelihood of how many populations of striped newt are predicted to be extirpated in the future, but the current development and sea level rise projections only identify two possible populations that could be extirpated.

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Table of Contents CHAPTER 1 - INTRODUCTION ...... 8 CHAPTER 2 – INDIVIDUAL NEEDS: LIFE HISTORY AND BIOLOGY ...... 10 2.1 Description ...... 10 2.2 Taxonomy and Nomenclature ...... 11 2.3 Life History ...... 12 2.4 Habitat ...... 14 2.5 Diet ...... 16 2.6 Genetic distribution ...... 16 2.7 Ecological Needs ...... 19 2.7 Historical Range and Distribution ...... 21 2.8 Current Range and Distribution ...... 22 2.9 Land Ownership ...... 32 CHAPTER 3 -- CURRENT SPECIES CONDITION ...... 34 3.1 Land use changes ...... 35 3.2 Fire suppression ...... 36 3.3 Climate Change and Drought ...... 38 3.3.1 Climate Change ...... 38 3.3.2 Sea Level Rise...... 39 3.3.3 Drought ...... 44 3.3.4 Climate Change and Drought summary ...... 45 3.4 Recreation ...... 46 3.4.1 Off-road Vehicles Impacts ...... 46 3.5 Other Stressors ...... 47 3.5.1 Predation ...... 48 3.6 Combined Stressors ...... 48 3.7 Existing Regulatory mechanisms ...... 48 3.8 Current Conservation Measures ...... 51 3.9 Summary of Species Overall Current Condition: Population Resilience, Representation, and Redundancy ...... 52 CHAPTER 4 – FUTURE CONDITION SCENARIOS ...... 55 4.1 Introduction ...... 55 4.2 Development – future impacts ...... 56 4.3 Drought – Future impacts ...... 57 5

4.4 Scenario 1 ...... 60 4.4.1 Resiliency ...... 63 4.4.2 Representation...... 63 4.4.3 Redundancy...... 63 4.5 Scenario 2 ...... 63 4.5.1 Resiliency ...... 64 4.5.2 Representation...... 65 4.5.3 Redundancy...... 65 4.6 Scenario 3 ...... 65 4.6.1 Resiliency ...... 66 4.6.2 Representation...... 67 4.6.3 Redundancy...... 67 CHAPTER 5: SYNTHESIS ...... 68 LITERATURE CITED ...... 70

List of Tables and Figures Figure 1. Species Status Assessment Framework stages...... 10 Figure 2. Life-history schematic of the striped newt ...... 13 Figure 3. Ecological and genetic separation between eastern and western striped newt ...... 17 populations ...... 17 Figure 4. Genetic differentiation showing isolation by distance...... 18 Figure 5. Plot of genetic diversity (allelic richness) across geographic regions ...... 18 Figure 6. (a) Historical range of the striped newt. (b). Historical and known extant striped newt breeding ponds ...... 22 Figure 7. Current status of all known Notophthalmus perstriatus breeding ponds...... 24 Figure 8. Suitable habitat for the striped newt...... 26 Figure 9. Striped newt metapopulations across the species’ range ...... 27 Figure 10. Known striped newt breeding ponds throughout the last four decades...... 32 Figure 11. Land ownership and striped newt breeding pond locations ...... 33 Figure 12. Conceptual model for stressors impacting the striped newt ...... 34 Figure 13. Prescribed burning trends in Southern US. Kozbiar et al. 2015 ...... 38 Figure 14. Relative risk that physical changes will occur as sea level rises ...... 40 Figure 15. Mean sea level trends ...... 42 Figure 16. Sea level rise impacts to Florida coastal populations of striped newts...... 43 Figure 17. Sea level rise in Eastern Region of striped newt habitat in Georgia...... 43 Figure 18. Potential metapopulations identified for striped newt...... 54 Figure 19. Climate projections in the southeastern U.S.A...... 58 Figure 20. Summary of forecasted population condition expected under the three future condition scenarios examined...... 68

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Table 1. Life history and species needs of the striped newt ...... 20 Table 2. Striped Newt (Notophthalmus perstriatus) breeding ponds for each time period ...... 29 Table 3: Rainfall simulations using the CMIP5 models for regions within Florida and Georgia 39 Table 4. Potential striped newt suitable habitat loss due to sea level rise...... 42 Table 5. Model calculations of suitable habitat changes with various development scenarios ... 57 Table 6. Future scenarios for the striped newt...... 59 Table 7. Predicted responses of striped newt populations under three scenarios ...... 61

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CHAPTER 1 – INTRODUCTION

The striped newt (Notophthalmus perstriatus) is a small salamander that reaches a total length of up to 10 centimeters (cm) (4 inches (in)) and has a continuous red stripe along the side of its body (Conant and Collins 1991, p. 258). It is currently found in 13 counties in Florida and 6 counties in Georgia. The species was petitioned for federal listing under the Endangered Species Act of 1973, as amended (ESA), on July 10, 2008 by the Coastal Plains Institute and Land Conservancy (Means 2008). It has been a candidate for listing under the Act since the 12-month finding was published in 2011 (USFWS 2011).

The Species Status Assessment (SSA) framework (USFWS 2016) is intended to be an in-depth review of the species’ biology and threats, an evaluation of its biological status, and an assessment of the resources and conditions needed to maintain long-term viability. The intent is for the SSA Report to be easily updated as new information becomes available for a listed species, and to support all functions of the Endangered Species Program from Candidate Assessment, to Listing, to Consultations to Recovery. As such, the SSA Report will be a living document that may be used to inform Act decision making in many categories including listing, recovery, Section 7, Section 10, and reclassification decisions (the former four decision types are only relevant should the species warrant listing under the Act).

This document draws scientific information from resources such as primary peer-reviewed literature, reports submitted to the U.S. Fish and Wildlife Service (Service) and other public agencies, species occurrence information in GIS databases, and expert experience and observations. It is preceded by, and draws upon analyses presented in other Service documents, including the 12-month finding (USFWS 2011) and the Species Assessment and Listing Priority Assignment Form associated with the most recent Candidate Notice of Review (USFWS 2016).

The striped newt SSA is intended to provide the best available commercial and scientific information in determining whether to list the species as threatened or endangered and, if so, to determine whether it is prudent and determinable to designate critical habitat in certain areas. Importantly, the SSA Report is not a decisional document by the Service; rather, it provides a review of available information strictly related to the current biological status of the species and factors that may affect its future biological status. The listing decision will be made by the Service after reviewing this document and all relevant laws, regulations, and policies, and the results of a proposed decision will be announced in the Federal Register with appropriate opportunities for public input.

For the purpose of this assessment, we define viability as the ability of the species to sustain resilient populations in their ecosystem for at least 43 years. We chose 43 years because it is within the range of the available development and climate change model forecasts. Using the

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SSA framework (Figure 1), we consider what the species needs to maintain viability by characterizing the status of the species in terms of its redundancy, representation, and resiliency (USFWS 2016; Wolf et al. 2015).

 Resiliency describes the ability of a species to withstand stochastic disturbance. Resiliency is positively related to population size and growth rate and may be influenced by connectivity among populations. Generally speaking, populations need abundant individuals within habitat patches of adequate area and quality to maintain survival and reproduction in spite of a random disturbance.

 Representation is assessed at the species’ level. Representation describes the ability of a species to adapt to changing environmental conditions over time. For example, a species that has populations that exhibit geographic, genetic, or life history variation have greater ability to adapt to changing conditions. It is characterized by the breadth of genetic and environmental diversity within and among populations. Measures may include the number of varied niches occupied, the gene diversity, heterozygosity, or alleles per locus. Our analysis explores the relationship between the species life history and the influence of genetic and ecological diversity and the species' ability to adapt to changing environmental conditions over time. The analysis identifies areas representing important geographic, genetic, or life history variation (i.e., the species’ ecological settings).

 Redundancy describes the ability of a species to withstand catastrophic events (random events that have devastating consequences); it is about spreading risk among multiple populations to minimize the potential extinction of the species from catastrophic events. Redundancy is characterized by having multiple, resilient populations distributed within the species’ ecological settings and across the species’ range. It can be measured by population number, resiliency, spatial extent, and degree of connectivity. Our analysis explores the influence of the number, distribution, and connectivity of populations on the species’ ability to withstand catastrophic events (e.g., rescue effect).

To evaluate the current and future viability of the striped newt, we assessed a range of conditions to characterize the species’ resiliency, representation, and redundancy (together, the 3Rs). This SSA Report provides a thorough account of known biology and natural history and assesses the risk of threats and the limiting factors affecting the future viability of the species.

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Figure 1. Species Status Assessment Framework stages.

This SSA Report includes: (1) a description of striped newt resource needs at both individual and population levels (Chapter 2); (2) a characterization of the historical and current distribution of populations across the species’ range (Chapter 3); (3) an assessment of the stressors and conditions that contributed to the current and future status of the species and the degree to which various factors influenced viability (Chapter 4); and (4) a synopsis of the needs and stressors characterized in earlier chapters as a means of examining the future biological status of the species (Chapter 5). This document is a compilation of the best available scientific information (and associated uncertainties regarding that information) used to assess the viability of the striped newt.

CHAPTER 2 – INDIVIDUAL NEEDS: LIFE HISTORY AND BIOLOGY

In this section, we provide basic biological information about the striped newt, including its physical environment, taxonomic history, species description, known behavior, and other life history traits. We then outline the resource needs of individuals and populations. Here we report those aspects of the life histories that are important to our analyses.

2.1 Description

The striped newt (Notophthalmus perstriatus) is a small salamander that reaches a total length of 5–10 cm (2–4 in) and a mass of 0.8 grams (g) (0.03 ounces (oz)) (Conant and Collins 1991, p. 258). There are more than three distinct life stages of the striped newt, each with varying

10 appearances (Figure 2). The larval stage is the sexually immature aquatic stage newt. Larvae have bushy external gills, a membranous tail fin, and a conspicuous lateral line, which is visible as a series of dorsolateral dashes on each side of the animal. An eft is a sexually immature terrestrial stage newt that lack gills and does not have a tail fin. A paedomorph is a sexually mature, aquatic stage newt. Finally, an adult is a sexually mature and dimorphic terrestrial or aquatic stage newt which lack gills. On adults and most efts, a continuous red stripe runs the length of the side of its trunk and extends onto the head and tail where it may become fragmented. The stripe is dark-bordered, but not so boldly and evenly as in the broken striped newt (N. viridescens dorsalis) (Conant and Collins 1991, p. 258). There may be a row of red spots along the side of the body and a faint light stripe down the center of its back. The ground color of the sides and back is olive-green to dark brown. The belly is yellow, usually sparsely marked with black specks. The skin of terrestrial newts tends to be rougher and less slimy than other salamanders. The costal grooves (grooves along the side body of salamanders used in species identification) are indistinct.

2.2 Taxonomy and Nomenclature

The currently accepted classification for the striped newt (Notophthalmus perstriatus) is (Integrated Taxonomic Information System 2016): Phylum: Chordata Class: Amphibia Order: Caudata Family: Salamandridae Genus: Notophthalmus Species: perstriatus

The striped newt was formally described by Bishop (1941, pp. 3–6) from specimens collected in Alachua and Leon counties, Florida, and Charlton Co., Georgia. Originally called Triturus perstriatus by Bishop (1941), Neill (1952, 1954) and Schmidt (1953) called the species Diemictylus perstriatus until the name became stabilized as Notophthalmus perstriatus by Smith (1953). There are three species of Notophthalmus found in North America. These include the eastern newt (N. viridescens), the blackspotted newt (N. meridionalis), and the striped newt (N. perstriatus). The three species are found in different areas throughout the United States and Mexico (Reilly 1990, p. 51). In a study of Notophthalmus spp., N. perstriatus and N. meridionalis were found to be distinct species that are more similar and phylogenetically more closely related to each other than either is to N. viridescens (Reilly 1990, p. 53),. In addition, the phylogenetic relationship (evolutionary history of an organism) of the family Salamandridae shows that the clade (group of species that includes all descendants of a common ancestor) containing newts is separate from the clade containing “true” salamanders (Zhang et al. 2008, pp. 586 and 592). The branching order of the clades for newts are: primitive newts (Echinotriton, Pleurodeles, and Tylototriton), New World newts (Notophthalmus and Taricha), Corisca-

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Sardinia newts (Euproctus), modern European newts (Calotriton, Lissotriton, Mesotriton, Neurergus, Ommatotriton, and Triturus), and modern Asian newts (Cynops, Pachytriton, and Paramesotriton). New World newts, which include Notophthalmus, originally evolved from salamandrids migrating from Europe to North America via the North Atlantic land bridge during the Mid-Late Eocene (Zhang et al. 2008, p. 595). The striped newt is considered a valid species (e.g., Zhang et al. 2008).

2.3 Life History

Striped newts have a lifespan of 12–15 years (Dodd Jr. 1993; de Magalhaes and Costa 2009) and may use aquatic and terrestrial habitats during their complex life cycle (Figure 2). Adults may inhabit isolated, temporary ponds or adjacent upland areas. Reproduction occurs in isolated, temporary ponds that lack predatory fish (introduced game fish, sunfish (Centrarchidae spp.), largemouth bass (Micropterus salmoides), and pickerel (Esox spp.)). Large predatory fish are a concern for striped newts but they successfully breed in ponds with smaller or gape-limited predatory fish (i.e., eastern mosquitofish ((Gambusia holbrooki)) (Enge et al. 2014a, pp. 60–61). Terrestrial adults migrate to these breeding ponds from adjacent uplands for courtship, copulation, and egg-laying. Females lay eggs one at a time and attach them to aquatic vegetation or other objects in the water. It may take one female several months to lay all of her eggs (Johnson 2005, p. 94). Eggs hatch and develop into externally-gilled larvae in the temporary pond environment.

Once larvae reach a size suitable for metamorphosis (18–30 millimeters (mm) (0.71–1.18 in) snout-vent length), they may either undergo metamorphosis and exit the pond as immature, terrestrial efts, or remain in the pond and eventually mature into gilled, aquatic adults (paedomorphs) (Dodd 1993, p. 609; Petranka 1998, pp. 449–450; Johnson 2005, p. 94; Dodd et al. 2005). The immature, terrestrial efts migrate into the uplands where they mature into terrestrial adults (Figure 2 c and e). Efts will remain in the uplands until conditions are appropriate (adequate rainfall) to return to the ponds to reproduce. During dry weather, they hide beneath logs and stones or in the moist humus of the forest floor (Bishop 1943, p.70). Terrestrial efts move to and from ponds during fall and winter rains (Christman and Means 1992, p. 63). Johnson (2005, p. 94) found that 25% of larvae became paedomorphs (mature while retaining larval characteristics) at his study pond. Paedomorphs will postpone metamorphosis until after they have matured and reproduced (Figure 2 d). At about a year old, they will reproduce, metamorphose, and migrate into the uplands adjacent to the pond (Johnson 2005, pp. 94–95). Once there are proper conditions (e.g., adequate rainfall) at the ponds, the terrestrial adults will move back to the ponds to court and reproduce. After returning to the ponds, the newts are referred to as aquatic adults (Figure 2 f). It is also believed that, both in the winter and during colder periods of spring and fall, entire population of a pond will disappear, presumably burrowing in the mud of the bottom (Bishop 1943, p. 70). In addition, when temporary ponds and pools dry up in late fall, many individuals are forced to bury themselves and go into

12 hibernation (Bishop 1943, p. 70).

Most of the surveys for striped newts are done during the breeding season when the newts return to the ponds and when aquatic adults and larvae can be sampled by dip netting. This sampling method is more opportunistic and takes a lot less resources than installing, maintaining, and checking drift fences and pit traps. Although it is a lot easier to dipnet for striped newts, it is very opportunistic and depends on visiting potential breeding ponds at the right time of the year when the breeding ponds have water. Detection of amphibians in their aquatic phase is easier in ponds with high density, longer length of the breeding season, and where the species stays longer in their aquatic stage (Gunzburger 2007). We did not find any published literature that evaluated the detection success of dipnetting for striped newts. Anecdotal accounts are that detectability may depend on abundance and surveying at the right time (Mays, 2017, pers. comm.). Some ponds can be visited multiple times with varying degrees of success.

Figure 2. Life-history schematic of the striped newt. Johnson 2001

The complex life-history of striped newts, their 12–15 year lifespan, and ability to live as adults in aquatic and terrestrial habitats allow individuals to survive during periods of unfavorable breeding conditions. For ideal reproduction and development, there must be enough water in ephemeral ponds for the larvae to have enough time to reach the minimum size needed for metamorphosis (Johnson 2002, p. 398). However, permanent ponds could support predatory fish that feed on aquatic-breeding amphibians (Johnson 2005, p. 94; Moler and Franz 1987, p. 235). Variable hydroperiods in breeding ponds over a long time period could result in varying reproductive success. Persistent drought conditions have shown local declines in the number of 13 striped newts detected in breeding ponds (Dodd 1993, p. 610) and heavy rainfall has contributed to the reproductive success at local ponds (Johnson 2002, p. 399). At one breeding pond, a minimum hydroperiod of 139 days (Dodd 1993, pp. 609–610) was needed for larvae to reach complete metamorphosis. Larvae undergo metamorphosis into efts after a period of 6 months, and a study showed that in order for reproduction to occur, a breeding pond must hold water long enough for larvae to complete development (Johnson 2005, p. 94). For a paedomorph to successfully reproduce, ponds must hold water for an additional 6 months to allow sufficient time for its larvae to emerge and complete metamorphosis.

Suitability of upland habitat around breeding ponds influences the pattern of immigration and emigration of newts and directional movements (Dodd 1996, p. 46; Dodd and Cade 1998, p. 337; Johnson 2003, p. 16). Newts migrate into terrestrial habitats from their breeding ponds in a direction that favors high pine sandhill habitats (Dodd and Cade 1998, p. 337). One of the studies found that 82.9% of 12 wetland breeding amphibians (including striped newts) were captured 600 meters (m) (1,969 feet (ft)) from the nearest wetland, and only a fourth of the amphibians were captured less than 400 m (1,300 ft) from the nearest wetland (Dodd 1996, p.46). Another study showed that 16% of striped newts migrated more than 500 m (1,600 ft) from breeding ponds (Johnson 2003, p. 18) and the third study showed that striped newts travelled up to 709 m (2,330 ft) from breeding ponds (Dodd and Cade 1998, p. 337). Given the above, newts typically move less than 700 m (2,296 ft) from the nearest wetland, but have an estimated dispersal distance of 1 km (3,280 ft) (Johnson 2005, p.97). We are considering this dispersal distance as the home range and distance between breeding ponds to connect populations.

For the purposes of this SSA report, we are using certain standardized terms to refer to habitat features. For instance, we define all individuals that occupy a breeding pond and the habitat surrounding the pond for 1 km as a population. As such, we use the terms "breeding pond" and "population" interchangeably throughout this SSA Report. We further define a metapopulation as local populations close enough to one another that dispersing individuals could be exchanged (gene flow) at least once per generation (Enge et al. 2014a, p. 28). Survey sites are composed of single pond populations, multiple populations, single or multiple metapopulations, and a mixture of single populations and metapopulations. Strongholds are sites that have had a mixture of populations since 2000 and/or have a robust metapopulation.

2.4 Habitat

Striped newts have a complex life cycle that requires both upland and wetland habitat (Johnson 2005, Figure 2). For reproduction, they depend on isolated ephemeral ponds with cyclic hydroperiods and that hold water long enough for larvae to develop. The cyclical nature of ephemeral ponds prevents predatory fish from inhabiting striped newt breeding ponds (Dodd and Charest 1988, pp. 87, 94; LaClaire and Franz 1990, p. 12; Moler and Franz 1987, p. 237). In addition, ponds throughout the regions may be able to hold water at various times throughout a

14 year, allowing for striped newt reproduction to occur at different times of the year. Although permanent ponds often contain large, predatory fish making them unsuitable for striped newt reproduction, striped newts will breed in semi-permanent and/or permanent ponds that lack predatory fish.

The frequency and duration of water in ephemeral ponds creates different zones of vegetation within ponds. One plant species, maidencane (Panicum hemitomon), has been found at ephemeral ponds where striped newts have been found, and seems to be a good indicator of the extent of previous flooding in ponds (LaClaire 1995, p. 88; LaClaire and Franz 1990, p. 10). Persistence of maidencane helps to reduce the rate of oxidation of organic matter, reduce soil moisture loss, and inhibit growth and establishment of upland plant species (LaClaire 1995, p. 94). The center of flooded ponds may contain aquatic floating plants (i.e., duckweed (Lemnoideae spp.); Nymphoides), and may be surrounded by vegetation with submerged roots growing along the wet edges (i.e., smartweed (Polygonum spp.). Surrounding wet areas contain tall and short emergents, such as sedges, grasses, and rushes, followed by other grasses such as bluestem grass (Andropogon virginicus) found in the drier margins of ponds. Water-tolerant shrubs or trees, such as sandweed (Hypericum fasciculatum), are found in some transitional zones between ponds and uplands (LaClaire 1995, p. 74; LaClaire and Franz 1990, p. 10).

Ephemeral ponds used by striped newts are surrounded by upland habitats of high pine, scrubby flatwoods, and/or scrub (Christman and Means, 1992, p. 62). Longleaf pine-turkey oak stands with intact ground cover containing wiregrass (Aristida stricta) are the preferred upland habitat for striped newts, followed by scrubby flatwoods and scrub (K. Enge, FWC, personal communication, 2010). Some breeding ponds are surrounded by mesic flatwoods or dry prairie, but more xeric upland habitat is found nearby. In addition, striped newts can be found in cypress- dominated dome swamps and borrow pits (Enge et al. 2014a, pp. 21, 43, 57). The habitat types that support striped newts are fire-dependent.

Striped newts use upland habitats that surround breeding ponds to complete their life cycle. Efts move from the breeding ponds to uplands where they mature into terrestrial adults. The uplands also provide habitat for the striped newt to forage and burrow during the non-breeding season (Dodd and Charest 1988, p. 95). Striped newts also use uplands to disperse to alternative ponds if the original breeding pond is destroyed or the hydroperiod is altered (Means 2006, p. 197). This shows the interdependence between upland and aquatic habitats in the persistence of populations (Semlitsch and Bodie 2003, p. 1219). Semi-aquatic species (such as the striped newt) depend on both aquatic and upland habitats for various parts of their life cycle in order to maintain viable populations (Dodd and Cade 1998, pp. 336–337; Johnson 2001, p. 47; Semlitsch 1998, p. 1116; Semlitsch and Bodie 2003, p. 1219).

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2.5 Diet

Life-history stages of the striped newt are complex and include the use of both aquatic and terrestrial habitats throughout their life cycle. Striped newts are opportunistic feeders that prey on frog eggs, worms, snails, fairy shrimp, spiders, and insects (adult and larvae) that are of appropriate size (Dodd et al. 2005, p. 889; Christman and Franz 1973, pp. 134–135; Christman and Means 1992, pp. 62–63). Christman and Franz (1973, p. 135) found that newts were attracted to frog eggs by smell. Feeding behavior of newts has only been documented with aquatic adults; little is known of the feeding habits in the terrestrial stage (Dodd et al. 2005, p. 889).

2.6 Genetic distribution

Recent studies looked at whether populations of striped newt that occur in two regions separated by 125 km (78 mi) exhibit genetic differences (May et al. 2011, p. 1442; Hoffman 2012, p. 5). The regions are described as having ecological differentiation due to precipitation and temperature differences that would show these two regions are separate conservation units (May et al. 2011, p. 1442; Hoffman 2012, p. 5). One region consists of populations located in peninsular Florida and southeastern Georgia, and the other region consists of populations located in northwestern Florida and southwestern Georgia (Dodd and LaClaire 1995, p. 42; Franz and Smith 1999, p. 13). Studies have shown that there is gene flow between localities within each region, but no genes were shared between regions (May et al. 2011, p. 1446). In addition, it has been shown that genetic exchange between populations is minimal or nonexistent due to upland habitat fragmentation that has limited long-distance dispersals and restricted gene flow (Johnson, 2001, pp. 107,113–115). However, a separate study, May et al. (2011, p. 1448) concluded that the genetic divergence was not great enough to support a taxonomic separation of the Eastern and Western Regions into two distinct species. May et al. (2011, p. 1447) ran niche-based distribution models that showed there were significant climatic and environmental differences between the two regions when considering temperature and precipitation (Figure 3). The Western Region is characterized by lower mean temperatures and more extreme winter cold, coupled with higher variation in temperature and precipitation. The genetic differences between the regions have not been associated with any differences in physiology or ecology between the regions, so currently we cannot determine, with any measure of confidence, whether any of the genetic differences affect key life history stages.

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Figure 3. Ecological and genetic separation between eastern and western striped newt populations as presented in May et al. 2011.

Since these genetic studies, a population has been found at Big Bend WMA in Taylor County, Florida, in the purported distributional gap (Mays and Enge 2014, p. 275). This population is closest to the Western Region, but its genetic affiliation is closest to the eastern striped newts (Hoffman 2017, pers. comm.). Further work by Hoffman (unpublished data) shows that there is significant genetic structure and a significant pattern of isolation by distance (Figure 4). In addition, Hoffman describes that striped newts from western breeding ponds and eastern isolated ponds exhibit reduced genetic diversity compared to newts from the eastern stronghold breeding ponds (Figure 5). Sites (breeding pond sites) showed that gene flow between ponds was occurring only at a local scale for nearby ponds, and that all sites were genetically identifiable. The genetic information on the striped newt does show variation across its range, but no genetic information has shown a cause-effect relationship or correlational association with the newt’s physiology or key life history attributes.

From this point on we will be referring to the Eastern Region as the populations located in eastern Florida and eastern Georgia, as defined by the ecological model shown above (Mays et al. 2011). We will use the term Western Region as the term for the populations found in western Florida and western Georgia, also as defined in the model shown above from Mays et al (2011). The current species stronghold is defined as striped newt populations found in the Eastern Region in five public lands in Florida: Camp Blanding, Jennings State Forest, Ocala National Forest, Ordway-Swisher Biological Station, and Triple N Ranch Wildlife Management Area (WMA). For example, in Ocala National Forest, striped newts have been found in 38 breeding ponds since 2006 (Enge et al. 2014a, p. 83).

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Figure 4. Genetic differentiation showing isolation by distance. Hoffman et al. 2017, unpublished data.

Figure 5. Plot of genetic diversity (allelic richness) across geographic regions. Hoffman et al. 2017, unpublished data.

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2.7 Ecological Needs In this section, we describe the ecological needs of the striped newt at the different hierarchical levels of individual, population, and species (Table 1). 1. Individual Resource Needs (Figure 2): a. Egg: Ephemeral wetlands/ponds that retain water for at least 3–6 months, with emergent vegetation. b. Larvae: small invertebrates (zooplankton) for feeding, hydroperiod of at least 6 months for larvae to reach metamorphosis. c. Efts (immature terrestrial stage): uplands, small invertebrates for feeding. d. Paedomorphs (sexually mature aquatic stage): ponds or wetlands that retain water for at least 6 months, aquatic invertebrates, frog eggs for feeding, mates. e. Adults (sexually mature terrestrial): upland habitat, terrestrial invertebrates for feeding, mates. f. Adult (aquatic stage newt): ponds or wetlands that retain water for at least 6 months, mates, aquatic invertebrates, frog eggs for feeding.

2. Population Needs 2.1. Resource Needs and/or Circumstances: Striped newts must reproduce (fertilization and development) in an aquatic environment. They are known only to reproduce in ponds that are devoid of large, predatory fish species. They inhabit and reproduce in ephemeral wetlands that are isolated and dry out frequently because the isolation and dry downs prevent the establishment of predatory fish populations or eliminate predatory fish populations that became established during floods. Multiple suitable breeding ponds of varying sizes and hydroperiods should be found within a 1 km (population footprint/distribution) of each other and within 1 km of suitable upland habitat, so that the likelihood of all of the breeding ponds drying out during random, adverse events such as extended drought periods is low. For terrestrial and aquatic habitat to support survival, growth, and reproduction, frequent fire regimes are needed. Wetlands need to burn regularly when they are dry to maintain breeding habitat.

2.2. Population-level resiliency: Populations (defined as individual breeding ponds) must have enough individuals (population growth) so that after a random event (extended drought) occurs that limits reproduction, the newt populations are able to survive in the uplands and find mates in the breeding ponds after the adverse environmental conditions subside. Metapopulations (a cluster of breeding ponds within 1 km of each other) provide a higher level of resiliency to those populations. There needs to be enough individuals that make it to efts and terrestrial adults that can survive extended adverse environmental conditions (predatory fish invasion or drought) to maintain a population in less adverse years. That is, populations must always be large enough to persist through periods of stochastic events (i.e., severe storm events, prolonged droughts, etc.). However, in a

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metapopulation, populations may occupy, colonize, or be locally extirpated from breeding ponds at any point in time (Dodd et al. 2007, p.150). This interplay of scenarios in a metapopulation creates a higher level of resiliency for those populations.

3. Species Needs 3.1. Resource Needs and/or Circumstances: Connectivity among populations of striped newts is not documented well, but a suitable upland corridor (high pine, scrub, dry prairie, and pine flatwoods) that connects various ephemeral wetlands and allows for terrestrial adults to move between breeding ponds is likely needed. Apparent impediments to dispersal include unmanaged natural forests (fire suppression), developed areas, intensive agriculture, and industrial pine plantations.

3.2. Species-level redundancy: Currently, striped newts are identified as extant in 114 breeding ponds, have been previously collected from but are possibly extirpated in 57 ponds, and are reasonably certain to be extirpated from 17 breeding ponds (Figure 7; Farmer et al. 2017). These identified breeding ponds are part of 44 potential metapopulations (Enge et al. 2014a, p. 51) in Florida and at least 2 in Georgia, as well as and multiple populations represented by known single breeding ponds.

3.3. Species-level representation: Populations must be distributed in a variety of habitats throughout the range so that there are always some populations experiencing conditions that support some level of reproductive success. Especially in dry years, populations in peninsular Florida (Eastern Region) have less precipitation variations and breeding ponds hold water for enough time to allow reproduction to occur. Populations should be represented in the Eastern and Western regions of the species as presented in Section 2.6. There are currently 5 extant populations in the Western region and over 20 populations in the Eastern region of the species.

Table 1. Life history and species needs of the striped newt Life Stage Resources and/or circumstances needed for Resource Information INDIVIDUALS to complete each life stage Function Source (BFSD*)

Larvae Feed and grow in ephemeral wetlands/ponds. FS Johnson (immature aquatic 2001; stage newt) Johnson 2002

Efts (immature Uplands, longleaf pine forest, sandhill, and FSD Johnson terrestrial stage scrub habitat. 2001;

20 newt) Johnson 2002

Terrestrial Uplands, longleaf pine forest, sandhill, and FSD Johnson Adults (non- scrub habitat. 2001; gilled, sexually Johnson 2002 mature)

Paedomorphs Isolated ponds and/or ephemeral wetlands that BFS Johnson (sexually mature dry relatively often so fish won’t establish. 2001; aquatic stage newt) Johnson 2002

Eggs Aquatic vegetation in ephemeral S Johnson wetlands/ponds. 2001; Johnson 2002

Aquatic Isolated ponds and/or ephemeral wetlands that BFS Johnson Adults (sexually dry relatively often so fish won’t establish in 2001; mature aquatic ponds that dry frequently. Aquatic vegetation Johnson 2002 newt with tail fin) in wetlands/ponds to lay eggs.

* B=breeding; F=feeding; S=sheltering; D=dispersal

2.7 Historical Range and Distribution

The historical range (pre-1980) of the striped newt was likely similar to the current range (Dodd et al. 2005, p. 887). However, loss of native longleaf habitat, fire suppression, and the natural patchy distribution of upland habitats used by striped newts have resulted in fragmentation of existing populations (Johnson and Owen 2005, p. 2; Figure 6b). The range extends from the Atlantic Coastal Plain of southeastern Georgia to central Florida and west to Wakulla and Leon County in the Florida panhandle (Dodd et al. 2005, p. 887), upward to Taylor County in western Georgia (Jensen and Klaus 2004, p. 403). Prior to 2014, there was thought to be a 125-km (78- mi) separation between the western and eastern portions of the striped newt’s range (Dodd et al. 2005, p. 887; Dodd and LaClaire 1995, p. 42; Franz and Smith 1999, p. 12; Johnson 2001, pp. 115–116).

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a b

Big Bend WMA, Taylor County

Figure 6. (a) Historical range of the striped newt. (b). Historical and known extant striped newt breeding ponds (taken from Farmer et al. draft)

Figures 6a and 6b provide an illustration of the historical range of the striped newt and a comparison of the historical and extant breeding ponds within that range. Figure 6b was developed from various sources of data (Enge 2011, pp. 22–24; Enge et al. 2014a, pp. 130–133; Farmer et al. 2017, entire; J. Jensen, Georgia Department of Natural Resources (GADNR), personal communication, 2013). Historical records (pre-1980) in Florida describe 44 locations that include nine breeding ponds, in 14 counties from the panhandle east of the Apalachicola River to central peninsular Florida (Farmer et al. 2017). In Georgia, historical records document the striped newt from 23 breeding ponds in 11 counties extending from southeastern and south- central Georgia west to Lowndes and Wilcox counties (Farmer et al. 2017). A distributional gap between panhandle and peninsular populations in Florida seems to cross diagonally to Georgia, but it is apparent from the data but this may be an artifact of sampling opportunities or sampling success. Striped newts are sampled by dip netting during the breeding season. Successful surveys depend on specific timing and water availability in the breeding ponds/wetlands. Dip-netting is the best sampling technique for striped newts at the moment, but overall detectability of newts in a breeding pond may be low unless larvae are present in high densities.

2.8 Current Range and Distribution

This section describes the known current distribution of breeding ponds as reported in the literature, known areas where metapopulations exist, and the breeding ponds where striped newts are thought to be extirpated. Known extant, possibly extirpated, and extirpated populations have been recently identified in the literature and through additional surveys. Breeding ponds were considered extant when a pond and surrounding uplands remained suitable and if the most recent surveys (since 2000) found striped newts, and if a historical breeding pond was within 1 km of

22 an extant population (Farmer et al. 2017, p. 586). Breeding ponds are considered possibly extirpated if they had no recent observations (since 2000), or when multiple recent surveys had failed to detect the species (Farmer et al. 2017, p. 587). Striped newts were presumed extirpated from a breeding pond where the pond or the majority of immediately adjacent uplands were no longer suitable (Farmer et al. 2017, p. 587). The recent records (2000–2017) show that Florida has 105 extant populations and Georgia has 11 extant populations. We will be describing data that has shown populations in breeding ponds since 1980 but will be focusing on the last 30 years and describing the current condition of those populations.

Populations of striped newts in Florida have changed throughout the last decades but are still present in most of the historical areas. Records from Florida identify striped newts as extant in 77 breeding ponds in eight counties from 1980–1999 (Farmer et al 2017, p. 598). There are12 extant and 3 possibly extinct metapopulations identified for the striped newt (40 breeding ponds, including 2 unknown breeding ponds in 2 possibly extinct populations) in Ocala National Forest, 10 probably extant metapopulations (16 breeding ponds) in Camp Blanding Military Reservation, 5 extant metapopulations (18 breeding ponds) in Jennings State Forest, 5 possibly extinct metapopulations (18 breeding ponds) in Apalachicola National Forest (ANF), and 4 probably extant metapopulations (18 breeding ponds) in Ordway- Swisher Biological Station (Enge et al. 2014a, p. 51). Current surveys (2000–2017) have detected striped newts in 105 breeding ponds in 12 counties (Table 2; Figure 7; Farmer et al. 2017, p. 598). Most of the observations are from the “stronghold areas” (areas with multiple metapopulations) of the striped newt in peninsular Florida (Eastern Region). These stronghold areas are the same five public lands where metapopulations were identified in 2014: Camp Blanding, Jennings State Forest, Ocala National Forest, Ordway-Swisher Biological Station, and Triple N Ranch Wildlife Management Area (WMA). In 2016, surveys at ANF, where striped newts were thought to be extirpated and where translocation efforts have been focused, found a few “wild” adults in one of the historical breeding ponds; these metapopulations may not be extirpated. Records from 2000– 2016 were from 106 breeding ponds in thirteen counties (Farmer et al. 2017, Hill 2018, pers. comm.); all of these records are from breeding ponds that have been surveyed multiple years.

Multiple researchers have offered estimates of extant breeding ponds. These estimates differ due to the time periods that the researchers looked at. Current evaluation of those records indicates that striped newts have been extirpated from 17 breeding ponds, possibly extirpated from 40 breeding ponds, and extant at 105 breeding ponds (Figure 7) in Florida (Farmer et al. 2017, p. 588; Hill 2018, pers. comm.). This analysis found that twenty-five ponds in the peninsula Florida were not occupied during recent surveys but were placed in the extant category because of their proximity (< 1 km) to ponds with recent records.

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Figure 7. Current status of all known Notophthalmus perstriatus breeding ponds in Georgia and Florida, USA, classified as: extant, possibly extirpated, or extirpated. Based on recent surveys and information from Bishop (1941), Williamson and Moulis (1994), Dodd and LaClaire (1995), Franz and Smith (1999), and Krysko et al. (2011). Taken from Farmer et al. (2017); new records of two breeding ponds discovered in 2017 are not shown in this graph, total extant breeding ponds is 116.

Four ponds identified as extirpated by Farmer et al. (2017) included two ponds on the Lochloosa Wildlife Conservation Area in Alachua County whose uplands were converted to silviculture and a pond on Guana Tolomato Matanzas National Estuarine Research Reserve with upland habitat affected by fire suppression. Another breeding pond on private land in Leon County, Florida was deepened and converted to a horse pond and no longer holds striped newts. Ponds identified as possibly extirpated at ANF (with the exception of the one pond with recent records) and the Watermelon Pond tract of Goethe State Forest appear to still have suitable habitat but have not had striped newt observations during multiple recent surveys (Farmer et al. 2017).

Since the species has been a candidate (USFWS 2011, entire), striped newts have been detected in five breeding ponds in the Florida panhandle (Western Region), including a single pond on Apalachicola National Forest (ANF) and four ponds in a newly discovered population on a private plantation in Jefferson County (Hill and Sash 2015, p. 557; Hill 2018, pers. comm.). On February of 2014, researchers found three adult striped newts (2 males and 1 female) in one pond, and two adult males in a small, algae-lined hole of a dry depression marsh in the Spring Creek Unit of Big Bend WMA located in Taylor County, Florida (Mays and Enge 2014, p. 275).

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These discoveries fill a previously thought gap in the range, as the newly-discovered pond is located 43 miles east of the previously-known closest pond in Wakulla County, Florida, and 52 miles west of the previously-known closest pond in Gilchrist County, Florida. In addition, surveys have found a new metapopulation of striped newts at Triple N Ranch WMA, Osceola County (Eastern Region), which extended the known range 57 km SSE from the southernmost record on the University of Central Florida, Orange County (Enge et al. 2014a, p. 2, 10 and 40– 42). In 2015, striped newts were discovered for the first time in a pond on Merritt Island National Wildlife Refuge, Volusia County (Enge et al. 2015, p. 557).

Conservation efforts initiated by the Coastal Plains Institute in 2013 began a reintroduction program in the Munson Sandhills. The program has had success releasing captive bred striped newts at breeding ponds at ANF in Florida. The program has observed juvenile efts exiting the translocation ponds in 2013–2015, released larvae returning to one pond as mature adults in 2015–2016, and the sampled a first-generation larvae in 2016 (Ryan Means et al. unpublished. reports). First observations since 2007 of native striped newts at ANF occurred in 2016. This is the first evidence of breeding by native animals there since 1998 (Farmer et al. 2017; Ryan Means et al. unpublished reports).

The Florida Fish and Wildlife Conservation Commission (FWC) developed a striped newt habitat suitability model (Endries et al. 2009, pp. 45–46). The model identified three landcover classes: (1) Breeding (bay, cypress swamp, freshwater marsh, wet prairie), (2) primary upland (sandhill, xeric oak scrub, sand pine scrub), and (3) secondary upland (hardwood hammocks and forests, pinelands, and shrub and brushland). Breeding habitat was limited to patches that were less than 9 hectare (ha) (22 acres (ac)) in size and which were contiguous with upland habitats. The primary upland habitats included in the model were those areas contiguous and within 1,000 m (3,300 ft) of breeding habitat. Secondary upland habitat was included for areas that were contiguous and within 500 m (1,600 ft) of primary uplands and 1,000 m (3,300 ft) of breeding habitat. Their GIS analysis found a total of 244,576 ha (604,360 ac) of potential habitat (Endries et al. 2009, p. 45) (Figure 8). Of the potential habitat, 122,724 ha (303,257 ac) occurred on 124 sites within public lands, but only 64 of these sites had greater than 40 ha (100 ac) of potential habitat. The remaining habitat was found on privately owned lands in patches that were greater than 79 ha (195 ac) (Endries et al. 2009, pp. 45–46). Of the potential habitat found on public lands, 55% occurred on Ocala National Forest (ONF), 8% on Camp Blanding Joint Training Center, 6% on Withlacoochee State Forest, 5.3% on Apalachicola National Forest (ANF), and 2.9% on Jennings State Forest (K. Enge, FWC, personal communication, 2010). However, no records of striped newt occurrences have been found at Withlacoochee State Forest, even though this appears to be suitable habitat. Ocala National Forest has 67,514 ha (166,831 ac) of potential habitat and 38 occupied ponds, making it the largest “stronghold” for metapopulations for striped newts in Florida (Enge et al. 2014a, pp. 40 and 51). Striped newts are also found in ponds throughout peninsular Florida at Ordway-Swisher Biological Station, Camp Blanding Joint

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Training Center, Jennings State Forest, Goethe State Forest, Rock Springs Run State Reserve, Faver-Dykes State Park, Seminole State Forest, Big Bend WMA, Guana River WMA, Pumpkin Hill Creek Preserve State Park, and Triple N Ranch.

Figure 8. Suitable habitat for the striped newt. Endries et al. 2009

Analysis of striped newt breeding pond density shows metapopulations in areas of the identified species strongholds in Florida and Georgia (Florida: Camp Blanding, Jennings State Forest, Ocala National Forest, Ordway-Swisher Biological Station, and Triple N Ranch WMA; Georgia: Big Pond, Sandhills WMA) (Figure 9). This analysis also shows a cluster of breeding ponds at ANF in Florida where historically the striped newt was found in large numbers (Johnson 2005, p. 95; Johnson and Owen 2005, p. 7; Enge et. al., FWC, 2014a; USFWS 2011) and where re- introduction efforts have been ongoing.

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Figure 9. Striped newt metapopulations across the species’ range. Known multiple breeding ponds within 1000 m of each other.

As mentioned above striped newts have been found within the Munson Sandhills at ANF (Western Region). This site represents a small physiographic region within the Gulf Coastal Plains in Florida (Means and Means 1998a, p. 3). Striped newts have only been located in the western portion of the Munson Sandhills within the ANF. No newts have been found in the eastern portion of the sandhills since the 1980s, when the area was converted to a dense sand pine (Pinus clausa) plantation (Means and Means 1998a, p. 6). Habitat characteristics of occurrences in Apalachicola National Forest and St. Marks NWR versus those in northern and southern Ocala National Forest, where populations are extant, were compared and showed that the populations are separated by similar latitudinal distances but that the Western Region has

27 more annual and winter/fall precipitation, more disturbed areas, and closer proximity to major roads than populations from the Ocala National Forest (Enge 2014, p. 98).

Striped newt distribution continues north of the Munson Sandhills to the Tallahassee Red Hills, Tifton Uplands, and Dougherty Plain in southwestern Georgia (Western Region). Until recently, the Tallahassee Red Hills were no longer thought to support striped newts. Prior to 2015, striped newts were documented once in a breeding pond found in the Red Hills, but this site was dredged, deepened, and stocked with game fish in the 1980s and no longer supports newts (Means and Means 1998b, pp. 6, 15). However, in May 2015, March 2016, and July 2017, Pierson Hill and Kim Sash found striped newt larvae from a total of four ponds at the Dixie Plantation in northern Jefferson County (Hill and Sash 2015, p. 557; Hill 2018, pers. comm.). This represented a new county record and in 2015 the only known extant breeding pond in the Red Hills region. Three of the four breeding ponds where newts are found at Dixie Plantation have high diversity and/or abundant submergent wetland vegetation and emergent grasses. The upland habitat is mature mixed pine woodland with fire-maintained old field vegetation (Hill 2018, pers. comm.). One of the populations is in a large semi-permanent depression marsh with several non-predatory fish where paedomorphic striped newts are abundant and the population is estimated to be in the thousands (Hill 2018, pers. comm.). Three other breeding ponds at the Dixie Plantation had much lower numbers of newts sampled.

In addition, wild or native striped newt adults (1 male, 2 females) were captured in a single pond in the Munson Sandhill region in 2016 (Means 2016, pers. comm.). This was the first time since the beginning of the breeding program where wild striped newts have been found at ANF (Means 2017, pers. comm.). Additional striped newts have been released at ANF by the Coastal Plains Institute and partners in multiple years. As part of the captive breeding efforts, the program collected striped newt larvae in 2017 from the Dixie Plantation, Jefferson County, FL and from Big Pond, Sandhills WMA, GA. These larvae will be taken to various Zoo programs to breed additional newts for release at ANF. We conclude that there are 105 extant populations, 4 populations presumed extirpated, and 38 populations potentially extirpated in Florida. Many of the extant populations are part of metapopulations in the stronghold areas.

In Georgia, the striped newt is currently known to occur in five and likely six separate locations: Fort Stewart (Eastern Region), Alapaha River WMA (Eastern Region), Private Tract (Eastern Region), Joseph W. Jones Ecological Research Center (Ichauway Reserve) (Western Region, potentially extirpated), Sandhills Wildlife Management Area (Western Region), and Ohoopee Dunes Wildlife Management Area (Eastern Region) (Jensen 2010, pers. comm.; Smith 2010 and 2017, pers. comm.; Stevenson 2000, p. 4; Stevenson and Cash 2008, p. 252; Stevenson et al. 2009a, pp. 2–3; Farmer et al. 2017). Most of these locations are within the Dougherty Plain (Baker County), Tifton Uplands (Irwin, Lanier, and Lowndes Counties), and the Barrier Island Sequence (Bryan, Camden, Charlton, Evans, and Long Counties) (Dodd and LaClaire 1995, pp.

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40–42). From 1993 to 1994, Dodd and LaClaire (1995, p. 40) found striped newts in one pond each at five sites in Irwin, Baker, and Charlton Counties, and a series of ponds at Ft. Stewart in Bryan and Evans Counties. Striped newts were first found on Trail Ridge in 1924 near Okefenokee National Wildlife Refuge (ONWR), but this area has been highly modified since the 1940s (Dodd 1995, p. 44; Dodd and LaClaire 1995, pp. 39–40), and newts are no longer found in this area, except for possibly in the ONWR. In 2008, a new striped newt site was found in Georgia in Camden County, which is the first record for this county since 1953 (Stevenson et al. 2009b, p. 248). In 2013, newts were detected in the Sandhills WMA, Lentile Tract (Irwin County), and Ohoopee Dunes WMA (Emanuel County) ponds (J. Jensen, GADNR, personal communication, 2014).

Compiled breeding pond records of striped newts from 49 unique locations in 16 counties in Georgia, (Figure 7; Table 2) may indicate that fragmentation has impacted some of these breeding ponds (Farmer et. al., 2017). Before 1980, records document the striped newt from 23 breeding ponds in 11 counties extending from southeastern and south-central Georgia west to Lowndes and Wilcox counties. Eight counties have records from 1980–1999 from 21 breeding ponds, and records for this period documented the species in southwestern Georgia at Ichauway Reserve (Baker County) and as far north as Taylor County in the Fall Line sandhills region (Jensen and Klaus 2004; Kenneth Dodd, Jr., unpubl. report). Recent records (2000–2016) in six counties document the species in nine breeding ponds. During the last five years, records of striped newts have been collected in conservation lands (public and privately owned lands that are identified as having natural resource value that are being managed at least partially for conservation purposes) and one private lands location. eDNA sampling has identified striped newts at Fort Stewart but their locations are not certain (Roy King, 2017, pers. comm.) and the size of the populations is unknown. Survey efforts in the last three years have not been able to identify striped newts at Ichauway Reserve. This site has good habitat conditions and the reasons for the apparent species decline in this area is unknown (Smith 2017, pers. comm.). Extirpation and low density numbers are two theories at the moment.

Table 2. Striped Newt (Notophthalmus perstriatus) breeding ponds for each time period and number of ponds with extant populations by state and property. Data from Farmer et al. 2017 and new information on recent surveys in 2017. Property (County) Pre- 1980– 2000– 2012– Extant 1980 1999 2016 2016 Florida Apalachicola National Forest (Leon) 1 18 5 1 1 Big Bend WMA (Taylor) 0 0 2 2 2 Camp Blanding Military Reservation 1 11 10 7 8 (Clay)

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Faver-Dykes SP (St. Johns) 0 1 3 1 3 Goethe SF (Alachua, Levy) 0 0 5 0 1 Guana River WMA (St. Johns) 0 0 3 3 3 Guana Tolomato Matanzas NERR (St. 0 1 1 0 0 Johns) Jennings SF (Clay) 0 11 13 13 17 Lochloosa WCA (Alachua) 1 2 0 0 0 Merritt Island NWR (Volusia) 0 0 1 1 1 Ocala National Forest (Lake, Marion, 3 15 40 12 39 Putnam) Ordway-Swisher Biological Station 0 12 7 5 13 (Putnam) Private land (Gilchrist) 1 0 0 0 0 Private land (Jefferson) 0 0 2 2 4 Private land (Leon) 1 0 0 0 0 Private land (Putnam) 0 3 0 0 0 Private land (St. Johns) 0 1 0 0 0 Pumpkin Hill Creek SP (Duval) 0 0 1 0 1 Rock Springs Run SP (Lake, Orange) 0 1 3 0 3 Seminole SF (Lake) 0 0 1 0 1 Triple N Ranch WMA (Osceola) 0 0 9 9 9 University of Central Florida (Orange) 1 1 0 0 0 Florida Total 9 77 106 56 106 Georgia Alapaha River WMA (Irwin) 0 1 1 1 1 Sandhills WMA (Taylor) 0 0 1 1 1 Fort Stewart Military Installation 2 12 2 0 6 (Bryan, Liberty, Long) Ichauway Reserve (Baker) 0 4 2 0 0 Ohoopee Dunes WMA (Emanuel) 0 1 1 1 1 Okeefenokee NWR (Charlton) 2 1 0 0 0 Private land (Bryan) 1 0 0 0 0 Private land (Camden) 1 0 2 0 2 Private land (Charlton) 8 0 0 0 0 Private land (Jenkins) 0 2 0 0 0 Private land (Screven) 1 0 0 0 0 Private land (Wilcox) 1 0 0 0 0 Unknown locations (Charlton, Early, 7 0 0 0 NA Emanuel, Lanier, Lowndes) Georgia Total 23 21 9 3 11

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Of the 49 known Georgia breeding ponds, Farmer et al. (2017) determined that striped newts were extirpated from 13 ponds, possibly extirpated from 17 ponds, and extant at 11 ponds (Figure 7). Striped newts were extirpated from eight historical sites from the Trail Ridge on the eastern side of Okefenokee Swamp in Charlton County (Dodd and LaClaire 1995; C. Kenneth Dodd, Jr., unpubl. report). In this area, commercial forestry practices (fire suppression, ditching, clear cutting, and bedding) have significantly diminished habitat quality on lands outside of Okefenokee National Wildlife Refuge (Eastern Region). Striped newts were extirpated from Chesser’s Island inside the refuge because of hydrological alterations to ponds on the island from ditching. Striped newts may still be extant in one breeding pond on the refuge where the habitat is intact and managed with prescribed fire, although multiple recent surveys have failed to detect the species (Jensen 2017, pers. comm.; Farmer et. al., 2017). Single breeding ponds in Bryan, Wilcox, and Camden counties are no longer suitable for striped newts due to long-term wetland fire suppression, wetland destruction, and urban development, respectively. Similarly, two ponds in Evans and Long Counties on Fort Stewart (Eastern Region) are no longer suitable for the species because of fire suppression in both, and ditching of one of the ponds.

Striped newts are currently extant in multiple breeding ponds on Fort Stewart and on a private land in Camden County where the species was discovered in 2008 (Table 2). Recent (2000−2016) records exist for two Fort Stewart ponds (Eastern Region), although newts may still occur at four other ponds with suitable habitat that are near ponds with recent records. Additional extant newt localities in Georgia, based on a single pond each, occur on the Ohoopee Dunes WMA (Emanuel County), Sandhills WMA (Taylor County), and Alapaha River WMA (Irwin County). Multiple ponds on Fort Stewart and Okefenokee National Wildlife Refuge are suggested to be possibly extirpated because of a lack of recent records despite multiple recent surveys. All ponds on Ichauway Reserve in Baker County, GA are thought to be possibly extirpated because the species has not been detected there since 2006 despite good habitat and multiple survey efforts. A breeding pond in the Sandhills WMA, Taylor County, GA (Western Region) has been reliable for collection of striped newts for the breeding program in Florida. Twenty-five striped newt larvae were collected on August 17, 2017 (Means 2017, pers. comm.).

Our analysis of survey efforts in Florida and Georgia, as well as data presented by Farmer et al. 2017 shows that 105 breeding ponds are extant in Florida and 11 in Georgia (Figure 10). There are 57 other populations that have been identified as “potentially extirpated” but these may or may not be currently occupied by striped newts. Prior to 1980, there were nine known striped newt breeding ponds in Florida and 23 in Georgia. Known breeding ponds increased in Florida during 2000–2017 as did survey efforts during that timeframe (Enge et al. 2014a and Farmer et al. 2017); overall, we consider this increase in known ponds to be due to an increase in survey effort, and not an actual population expansion. In Georgia, known number of extant breeding ponds decreased but survey efforts were not increased as was the case in Florida where habitat

31 was analyzed and potential breeding ponds were targeted for sampling (Farmer et al. 2017). Although striped newts were previously found in 5 ponds at the Ichauway Reserve (Baker County), none have been collected since 2006; the lack of sightings is thought to have been caused by severe droughts in the last three decades (Farmer et al. 2017) and potential low population densities (Smith 2017, pers. comm.). Efforts from 2012–2017 show an apparent decline in presence of striped newts in surveyed breeding ponds but experts consider breeding ponds located near occupied breeding ponds (< 1 km) to be extant (Johnson 2005; Farmer et al. 2017). The decline in extant populations in Georgia from 21 to 11 known ponds is believed to be due to drought impacts as mentioned above, habitat alteration, and fragmentation (Farmer et al. 2017; Jensen 2017, pers. comm.).

Striped Newt Breeding Ponds (Populations) 120

100

80

60 Florida Georgia

40 BreedingPonds

20

0 Pre-1980 1980-19992000-20162012-2016 Extant

Figure 10. Known striped newt breeding ponds throughout the last four decades. Solid lines show detection in striped newt breeding ponds from 1980 to 2016. The far right of the graph shows the number of breeding ponds where striped newts are believed to be extant.

2.9 Land Ownership

In Florida, GIS analysis of striped newt suitable habitat estimated that half of the 244,576 ha (604,360 ac) of remaining suitable habitat is currently on public lands (Endries et al. 2009). Of the 105 breeding ponds identified as extant, 103 are located in public conservation lands (Table 2, Figure 11). In Georgia, during the last five years, striped newts have been detected in only five properties that have one to multiple breeding ponds. Four of the properties are located on public

32 conservation lands and one is in private ownership without a conservation easement.

Figure 11. Land ownership and striped newt breeding pond locations

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CHAPTER 3 -- CURRENT SPECIES CONDITION

The habitat and demographic characteristics that are most influential to population resiliency were identified earlier under the Ecological Needs section. This included the species needs throughout its life cycle and the population needs. The stressors and their cause and effect upon these factors and the subspecies as a whole were identified through published literature and unpublished reports and are listed below (Figure 12). Of the listed stressors, two have been previously identified by experts as main drivers of changes to striped newt populations: land use changes (human development or habitat conversion or invasion of predatory fish), and climate change (prolonged drought and shift in precipitation events).

Potential stressors impacting the species include:  Land use change (habitat loss due to development or land conversion);  Climate changes: drought, shifts in seasonal timing and amounts of precipitation and rainfall, and sea level rise;  Fire Suppression  Recreation (off-roading)  Predation

Figure 12. Conceptual model for stressors impacting the striped newt. Identified stressors (left boxes), species variables boxes on the right of the model

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3.1 Land use changes

Habitat loss and alteration from land use changes has resulted in temporary or permanent loss of habitat for the striped newt. Examples of land use change impacting the species include silviculture (conversion of natural pine forests to intensely managed pine plantations), urban development, solar farms, and other agricultural practices. Current striped newt habitat distribution relative to historical (pre-1980) suitable habitat has decreased approximately 17% in Florida and in Georgia where striped newt populations have been fragmented (Endries et al. 2009; Farmer et al. 2017, p. 592). Using the available habitat suitability model for Florida presented by Endries et al. (2009), we calculated a loss of suitable species habitat of 101,715 acres (41,163 ha) from 2003 to 2014. This represents 16.8% of the identified suitable habitat in Florida. However, it is important to note that suitable habitat is not analogous to occupied habitat (there is more suitable habitat for striped newts than occupied habitat). At the time of the 2011 12-month finding, habitat loss due to silvicultural practices was identified as one of the primary threats impacting the species (USFWS 2011, p. 32922).

Land conversion from agricultural practices such as silviculture may be one of the largest land use changes in the species range (Enge 2014a, Farmer et al. 2017, page 590). In Georgia, land use changes due to agriculture dominate the uplands of the Coastal Plains. Striped newts are extirpated from eight historical sites from the Trail Ridge on the eastern side of Okefenokee Swamp in Charlton County (Dodd and LaClaire 1995; C. Kenneth Dodd, Jr., unpubl. report; Farmer et al. 2017). The extirpation is thought to be a result of commercial forestry practices (i.e., fire suppression, ditching, and bedding) which have significantly diminished habitat quality on lands outside of the Okefenokee National Wildlife Refuge. Forestry practices have also been the cause of multiple extirpations as mentioned above. In addition to direct habitat loss due to agricultural practices, associated changes in soil and hydrology due to agricultural practices continue to impact remaining isolated wetlands in Georgia (GADNR 2015; Stuber et al. 2016).

Habitat loss from the conversion of natural pine forests to intensely managed silviculture has greatly disrupted the dispersal of striped newts between breeding ponds and upland habitat. Means and Means (1998a, p. 6) found that striped newt habitat at the Munson Sandhills varied due to differences in silvicultural practice between the eastern and western portions of the Sandhills. In the western portion of the Sandhills found within ANF, native groundcover remains in the second-growth longleaf pine forests, where striped newts spend most of their adult life. However, the eastern portion of the Munson Sandhills has been clear-cut, roller-chopped, and planted in sand pine (Pinus clausa), which is now a closed canopy with little native ground cover. Surveys of ponds located in the eastern Munson Sandhills found no striped newts after the site was converted to sand pine plantations (Means and Means 1998a, p. 4; Means and Means 2005, pp. 58–59; Means 2008, p. 30).

Land use change from urban development is also impacting striped newt habitat. In 10 coastal 35

Georgia counties, the human population is expected to increase 51% by 2030 (Center for Quality Growth and Regional Development 2006, p. 4), but no estimate of impact on native habitats due to potential increased urban development was provided. Striped newts have been found within five of these coastal counties in Georgia, including Bryan, Camden, Long, Liberty, and Screven Counties (Franz and Smith 1999, p. 13, Stevenson 2000, pp. 6–7). In two of the five counties identified above, striped newts are believed to be extirpated.

Land use change is a current stressor that continues to cause ongoing habitat loss; we expect this stressor will continue to result in loss of unprotected suitable habitat into the future. Land use from commercial forestry practices is decreasing population resiliency and therefore overall species redundancy in Georgia. It is identified as a primary cause of population loss. Although a significant amount of the remaining and known striped newt habitat is in conservation lands (Figure 6), approximately 14–20% of the identified suitable habitat may be lost due to destruction by development. However, habitat loss due to land use changes alone does not appear to be a significant factor in the reduction of currently known occupied striped newt habitat. We expect that habitat loss due to road construction and other practices in conservation lands will be less than 5% in the next 43 years. Additionally, over 85% of the known extant populations are located in lands protected in conservation (Farmer et al. 2017, Appendix 2; Figure 6). Most of these conservation lands are larger than the dispersal range of the species and thus would provide upland habitat for the terrestrial phase of the striped newt and connectivity within metapopulations.

3.2 Fire suppression

Striped newt habitat is fire-dependent. Naturally ignited fires and prescribed burning maintain an open canopy and reduce forest floor litter. An open canopy provides sunlight necessary for the ground cover growth needed by newts for foraging and sheltering. Fire is also an important factor for wetland vegetation (LaClaire and Franz 1990, p. 10; Means 2008, p. 4). Historically, fire would be naturally ignited in the uplands during the late spring and early summer and would sweep through the dry pond basins, reducing organic matter and killing encroaching upland plant species (Means 2008, p. 4; Myer 1990, p. 189). Fire suppression in uplands that buffer breeding ponds allows fire-intolerant hardwoods to shade out herbaceous understory needed by striped newts for foraging and sheltering. As a result, fire shadows may form along the upslope wetland and upland boundary. In addition, the surrounding wetlands are often burned in the dormant season when wetlands are holding water. Wetlands need to burn when they are dry to reduce the build-up of organic matter and decrease hardwood encroachment (Farmer et al. 2017). Most of Florida’s woodlands were fire suppressed between 1920 and 1950 due to the U.S. Forest Service, timber farmers and other land owners that saw wild fires as a threat to communities and to forests (MacAllister and Harper, 1998, p.36). Silvicultural practices, development, roads, and urbanization have also contributed to the exclusion of fire in uplands as well as in striped newt breeding ponds.

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Fire suppression at many sites with newt breeding ponds has been concurrent with the conversion of uplands to pine plantations (Johnson 2005, p. 97). Lack of fire can result in the succession of natural pine forests converting to fire-intolerant species and forests dominated by hardwoods (Means 2008, pp. 27–28). The succession of natural pine forest to more shade- tolerant species, such as oaks and hickories, can result in the loss of ground cover such as wire grass that is needed by striped newts for shelter and foraging (Means 2001, p. 31). However, this shift in habitat to fire-intolerant species can be managed and ameliorated through controlled burns. Frequencies of prescribed burns in these uplands need to take place in a 1- to 3-year cycle to provide suitable habitat for striped newts (Johnson and Gjerstad 2006, pp. 287–292) and 10– 20 years for scrub habitat (FWC 2012, p. 303). This would also reduce the naturally woody components around the ephemeral ponds, and stimulate flowering of grasses used by the newts along the pond margins (Means 2006, p. 196).

Examples of areas where lack of fire has been a concern for the species can be found throughout the species' range. At St. Marks National Wildlife Refuge (SMNWR), surveys conducted in 2002–2005 and again in 2009 were not able to locate any newts at 34 ponds (K. Enge, FWC, personal communication, 2010; Dodd et al. 2007, p. 29). While SMNWR still provides a rich amphibian fauna, the last known observation of striped newts was in 1978 (Dodd et al. 2007, p. 29; Farmer et al. 2017, p. 590). Some areas of the species habitat at SMNWR appeared to be too degraded in 2010 to be suitable for striped newts; this was due to the lack of fire (Enge 2010, pers. comm.).

In Georgia, lack of prescribed fire has also impacted some of the historical sites. Farmer et al. (2017) considered striped newts to be extirpated from Chesser’s Island inside the Okefenokee National Wildlife Refuge because of hydrological alterations to ponds on the island from ditching (Farmer et al. 2017, p. 589); however, in areas where the habitat is intact and managed with prescribed fire, the species is believed to be extant. Additional breeding ponds in Fort Stewart are also considered unsuitable for striped newts due to long-term fire suppression and ditching of one of the ponds. Although two of the breeding ponds in Fort Stewart have been extirpated, striped newts are found in several other breeding ponds and eDNA sampling has identified the striped newt in several breeding ponds (Roy, 2017, pers. comm.).

In recent years, prescribed fire has been constantly increasing in managed lands; however, it is needed in many more. In Florida, some public land managers do not currently have the resources to implement effective habitat management programs (Howell et al. 2003, p.10). Although Florida and Georgia’s prescribed fire programs burn more than 404,686 ha (1,000,000 ac) a year (Melvin 2012, pp. 7–12), reports document that less than 25% of public land managers had been ranked as having an excellent prescribed burn program (Florida Department of Environmental Protection 2007, p. 1). In addition, while the amount of prescribed burning being conducted in the southeastern United States, including Georgia and Florida, is still increasing, the rate of that 37 increase has slowed over time, especially in private lands (Figure 13) (Kozbiar et al. 2015). It is likely that fire suppression may decrease habitat quality at local and regional level but not to a level of decreasing overall species viability.

Figure 13. Prescribed burning trends in Southern US. Kozbiar et al. 2015

3.3 Climate Change and Drought

3.3.1 Climate Change

The terms “climate” and “climate change” are defined by the Intergovernmental Panel on Climate Change (IPCC). The term “climate” refers to the mean and variability of different types of weather conditions over time, with 30 years being a typical period for such measurements (IPCC 2013, p. 1450). The term “climate change” thus refers to a change in the mean or variability of one or more measures of climate (for example, temperature or precipitation) that persists for an extended period, whether the change is due to natural variability or human activity (IPCC 2013, p. 1450).

The following scientific and ecological information on climate change includes summarized work by the National Climate Team and staff of the United States Fish and Wildlife Service (USFWS) from the 2014 publication entitled "Climate Change Impacts in the United States: The Third National Climate Assessment" (NCA) (Carter et al. 2014, entire). This team also summarized the 2013 publication from the International Panel of Climate Change (IPCC) entitled "Highlights of the IPCC 5th Assessment Report: The Physical Science Basis of Climate Change (WGI); Summary for Policymakers" (IPCC, 2013, entire). This information is being further condensed with a primary focus on Florida (USFWS 2017, entire).

The National Climate Assessment reports that average precipitation in northeast and south Florida has increased by +5 to +10% since 1900, and central and panhandle Florida regions have

38 decreased by -5 to -10% (Walsh et al. 2014, pp 32–35). Heavy downpours have increased over the last 30 to 50 years. There is currently a 27% increase in the frequency of heavy downpours in Florida since the 1970s (USFWS 2017, p. 4).

Climate simulations using the CMIP5 models indicate changes of precipitation seasonally for the State of Florida (Table 3). Projections for Georgia, North Florida and the Panhandle regions are for wetter fall/winter and drier summers (Ingram et al 2013, pp. 27–29). Changes in precipitation patterns may limit the striped newt's ability to breed and may provide fewer opportunities to breed during dry years. Shifts in the breeding season for the striped newt to take advantage of the wetter season is likely as FWC staff has witnessed breeding in the fall when the ponds are dry in the spring (FWC, 2018, comments). In Central Florida, projections are for wetter falls and winter seasons. The impact of drier rainy seasons is the potential drying of breeding ponds before striped newts are able to complete their aquatic larval stage and metamorphose into terrestrial efts. Prolonged periods of drought could also cause hydrological changes in the breeding ponds and loss in vegetative cover in the upland habitat. Heavy downpours may be beneficial to the species if the events are prolonged enough to maintain breeding ponds throughout the aquatic breeding cycle. However, heavy downpours could also have a negative impact if ponds overflow and flood waters carry large predatory fish to otherwise isolated ponds that support striped newts as was the case in the Panacea Unit at SMNWR (Enge et al. 2014a, pp. 92-93). During wet years, flooding can occur, as ponds overflow and large predatory fish may colonize the breeding ponds impacting localized populations. In addition, sheet flow through the uplands is a natural phenomenon during hurricanes and tropical storms and it is more common in poorly-drained habitats, such as pine flatwoods and dry prairie, but occasionally occurs in xeric habitats (Enge et al, 2014a, p.92). These hydrological alterations create conditions that facilitate fish colonization by connecting isolated wetlands with ditches and more permanent bodies of water. Also, creation of nearby roadside ditches, canals, borrow pits, or fish ponds serve as sources of fish introductions (Enge et al. 2014a, p.93).

Table 3: Rainfall simulations using the CMIP5 models for regions within Florida and Georgia, (USFWS 2017, p. 5; Carter et al. 2014). Regions Winter Spring Summer Fall Panhandle 0 to -10% 0 to +10% 0 to -10% +10 to +20% North Florida 0 to -10% 0 to +10% -10 to -20% +10 to +20% Central Florida 0 to +10% 0 to -10% -10 to -20% +10 to +20% Southern 0 to -10% 0 to +10% -10 to -20% +10 to +20% Georgia

3.3.2 Sea Level Rise

Potential changes in sea level rise were analyzed to determine future impacts to coastal

39 populations of striped newts. Currently, data show that over the next century changes in sea level are expected to follow the global trend reasonably closely, but with variations on shorter time scales (Mitchum et al. 2017). Global sea level rose about 20 cm (8 in) in the last century and is projected to rise another 0.3–1.2 m (1–4 ft) in this century (Carter et al. 2014, pp. 400–402) (Figure 14). Further analysis shows that unless greenhouse gas emissions are reduced, sea level in Florida will most likely increase by 1–2 m (3.2–6.6 ft) over the next 50 to 100 years (Mitchum et al. 2017).

Figure 14. Relative risk that physical changes will occur as sea level rises (Hammar-Klose and Thieler 2001).

In order to understand sea level trends, we examined monthly mean sea level data from 1925– 1983 in Daytona, Florida; data from 1935–2016 in Savannah, Georgia (Eastern Region); and data from 1967–2016 in the Florida panhandle (NOAA 2017). Relative Sea Level Rise in Feet, on top of MHHW (Mean Higher High Water) was calculated using NOAA's 2015 mea sea level (MSL) trends and the USACE Sea Level Change Curve Calculator. Sea level rise projections were mapped at decadal intervals (2040–2100 for MHHW and MSL). We calculated the potential loss of striped newt suitable habitat using a GIS analysis of the state landcover data and elevation layers in conjunction with projected sea level rise. This projection resulted in a loss of 3,526 ha (8,714 ac) of striped newt suitable habitat by the year 2060 if sea level rise follows the more extreme projection, C5 (Table 4) (NOAA, 2017).

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The mean sea level trend in Daytona, Florida (Eastern Region) is 2.32 mm/year with a 95% confidence interval of +/- 0.63 mm/yr, which is equivalent to a change of 104 mm (0.34 ft) in 45 years (Figure 15a). In Savanah, Georgia the mean sea level trend is 3.23 mm/year with a 95% confidence interval of +/- 0.28 mm/yr, which is equivalent to a change of 145 mm (0.477 ft) in 45 years (Figure 15b). For the Florida Panhandle (Western Region), the mean sea level trend is 2.14 millimeters/year with a 95% confidence interval of +/- 0.66 mm/yr, which is equivalent to a change of 97 mm (0.32 ft) in 45 years (Figure 15c).

a

b

41

C

Figure 15. (a) Mean sea level trend in Daytona, Florida (Eastern region). (b) Mean sea level trend in Savanah, Georgia (Eastern Region). (c) Mean sea level trend in Apalachicola, Florida (Panhandle Florida) (Western Region).

Table 4. Potential striped newt suitable habitat loss due to sea level rise.

Striped Newt Inundated Inudated Inundated Inudated Inundated Inudated suitable Habitat (ac) 2040 (ac) Percent 2060 (ac) Percent 2100 (ac) Percent 502,645 C1 1,623 0.3% C1 1,871 0.4% C1 2,470 0.5% 502,645 C2 1,984 0.4% C2 2,800 0.6% C2 5,466 1.1% 502,645 C3 3,083 0.6% C3 5,561 1.1% C3 12,658 2.5% 502,645 C4 3,625 0.7% C4 6,793 1.4% C4 16,476 3.3% 502,645 C5 4,581 0.9% C5 8,714 1.7% C5 23,260 4.6%

We evaluated all of the extant coastal striped newt populations for potential impacts to sea level rise, based on SLR projections of 0.1 m (0.34 ft) in the next 40–50 years (NOAA 2017; Carter et al. 2014). Our analysis found three striped newt populations in the Guana River WMA would be impacted by saltwater intrusion into breeding ponds and by sea level rise (Figure 16) per the SLR projections for 2100, but only one population would be impacted by current SLR projections. Other populations analyzed were the Big Bend WMA and the Guana Tolomato Matanzas National Estuarine Research Reserve. These populations did not appear to be impacted by SLR in the time projections we analyzed.

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Figure 16. Sea level rise impacts to Florida coastal populations of striped newt in the Eastern and Western Region.

Figure 17. Sea level rise in Eastern Region of striped newt habitat in Georgia.

We would like to note again that the habitat we evaluated is suitable habitat and not just the known, currently occupied habitat. Within the 3,526 ha (8,714 ac) of striped newt suitable habitat that is projected to be lost by the year 2060, only one of the Eastern Region populations in the Guana River WMA is likely to be inundated. We evaluated all of the coastal populations within the species range and none of the other populations would be lost due to SLR. The population in

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Camden, GA and Merritt Island NWR which were the closest would not be inundated. Therefore, we determined that SLR would have a low impact on the redundancy and resiliency of the species due to the loss of one population in the next 40–50 years.

3.3.3 Drought

Dry consecutive days in most of Florida are expected to vary widely in the future, with possible changes from either a 10% decrease to a 20% increase (USFWS 2017). An analysis done of long-term climate data revealed marked differences between the Florida panhandle and peninsula (Enge et al. 2014a, p. 3–5 and 29–30). Since 1900, the Florida Climate Center shows, that every decade at least one severe and widespread drought somewhere within Florida has occurred, with the most severe droughts occurring in 1906, 1927, 1945, 1950, 1955, 1961, 1968, 1980, 1984, 1998, and 2006 (Collins et al. 2017, p. 584–586). Drought indices showed a significant trend of increasing drought over time in the peninsula (excluding the northern peninsula) but not in the panhandle in Florida. Drought periods in Georgia are also expected to follow the peninsular Florida trends (Carter et al. 2014, pp. 405–406).

Droughts can limit or completely eliminate suitable conditions for striped newt breeding (Dodd et al. 2007, p.151). When striped newts' breeding ponds are dry, they may attempt to disperse to other available wetlands within their dispersal distance (1 km) (Enge 2014; Dodd et al. 2007). In metapopulations, local extirpations or poor recruitment from individual ponds are offset by colonization of individuals from nearby ponds and availability of other breeding ponds (Marsh and Trenham 2001; Dodd et al. 2007; Enge 2014). Single, isolated populations are more vulnerable because they do not have a source population from which they can be recolonized or other breeding ponds that may still provide breeding habitat for a source population (Dodd et al. 2007). In the Western Region, three of the populations are single breeding pond populations. The populations at Dixie Plantation and ANF have multiple breeding ponds available. The Georgia population that has been used as a source for the ANF translocation newts is found in a single breeding pond.

Even though the striped newt is a species adapted to drought periods, the effects of a long-term drought have contributed to the decline of striped newts from breeding ponds at not only the ANF populations in Florida, but at breeding sites throughout Florida and Georgia. Droughts normally occur in cycles and amphibian populations fluctuate with drought conditions (Dodd 1992, pp. 138–139). However, droughts lasting several years (more than 4) were found to have affected reproductive success, resulting in population level declines (Dodd 1992, p. 139; Dodd and Johnson 2007, p. 150; Petranka 1998, p. 450). Surveys conducted at the Camp Blanding Joint Training Center in 2000 to 2001 during a drought did not find any striped newts due to dry breeding ponds. In previous years, surveys found 7 to 10 sites with newts (Gregory et al. 2006, p. 487) and they were found in 8 breeding ponds in 2011 (Enge 2014) which may show that even

44 though there were dry conditions, there isn’t evidence of a population decline at Camp Blanding. Striped newts are adapted to changes in hydrological conditions, and surveys depend on good wet years and timing for detection of the species.

Striped newts will respond to drought conditions in several ways: (1) temporary extirpation; (2) migration to adjacent areas with better habitat conditions; and (3) survival in upland habitat, with recolonization once water has returned (Dodd 1993, p. 612). As observed in a population on the Ordway-Swisher Biological Station, after a prolonged drought in 1988–1990, striped newts returned to Breezeway Pond which was monitored for a ten year period and where captured striped newts fluctuated between 34 to 970 with a total of 2,489 striped newts during that period (Dodd et al. 2007, p. 607). In 1988, larval striped newts were collected from the pond and from other ponds at the Biological Station (Dodd et al. 2007, p. 150–151). From 1996–1998, there were 8,127 striped newts counted entering and exiting one-shot pond at Ordway-Swisher Biological Station (Dodd et al. 2007, p.151). Two additional ponds at the Station were also recolonized after the drought period (Fox Pond with 32 individuals sampled and Blue Pond) and currently 13 breeding ponds are occupied (Dodd et al. 2007, p. 151; Farmer et al, 2017).

Another example is at Apalachicola National Forest where even with the return of water, striped newts had not been identified (Means 2013, pers. comm.), but there may be initial signs that a natural recolonization of the breeding ponds is possible. Striped newts used to be abundant in 19 ponds on the forest, but repeated surveys of all of the historical and nearby suitable ponds had not found striped newts until for the first since 2007, “native” striped newt adults (1 male, 2 females) were captured in a single pond in ANF in 2016 (Means 2016, pers. comm.). This may be evidence that with the return of water and habitat improvements at ANF these changes are benefiting the existing population of striped newts, and the original population of newts may be slowly recolonizing the historical breeding ponds and becoming more abundant where detection of the species is more probable.

Extended droughts are likely to have an impact on the resiliency, representation, and redundancy of the striped newt. The species has shown resiliency in the last decades where metapopulations exist and there has been evidence of “recolonization” or the return of striped newt to breed in breeding ponds/wetlands where they had been thought to be extirpated. Drought conditions are expected to be worst in the Eastern Region where the species is more secure at this moment and exists in metapopulations; therefore, though the drought conditions will be more severe in this area, we do not expect a substantial impact on the species.

3.3.4 Climate Change and Drought summary

We evaluated the available historical weather data and the species biology to determine the likelihood of effects that climate has had and will continue to have on striped newts. Sea level

45 rise will only inundate one population of striped newts and is likely to impact two others by salt intrusion. In the southeastern United States, droughts typically display a relatively shorter duration and are mostly ameliorated by the periodic occurrence of tropical cyclones (Ingram et al. 2013, p.15). Severe and prolonged droughts may be a stressor for the striped newt when drought conditions are extended; however, areas where metapopulations exist have maintained striped newt populations throughout drought periods. The likelihood that severe drought and climate related shifts in precipitation will continue to affect the species across its current range is likely, but the potential magnitude is unknown give uncertainties in temperature and precipitation projections.

3.4 Recreation

3.4.1 Off-road Vehicles Impacts

Off-road vehicle (ORV) use impacts upland and wetland habitats and has the potential to cause direct mortality of individual amphibians on both public and private lands (Enge 2014, p. 95). Off-road vehicles have destroyed the littoral zone of some striped newt breeding ponds in Apalachicola National Forest (USFWS 2011; Bruce Means, unpubl. report). At their study ponds in the Munson Sandhills at ANF, Means et al. (1994, pp. 6–7; 2008, pp. 11, 16) found that ORV use had degraded the littoral zone of the breeding ponds into barren sandy beaches unsuitable for striped newts. The littoral zone provides shallow, warm water where small aquatic invertebrates are concentrated, providing food for newts. Off-road vehicle use also destroys the grasses and grass-like vegetation around the ponds needed by newts for protection from predators such as wading birds (Means et al. 2008, p. 11). In 1994, 27 of 100 ponds at ANF were found to be damaged by ORV use, including 3 of 18 striped newt ponds (Means et al. 1994, pp. 6–7). By 2006, ORV impacts were documented at nearly every pond at ANF (Means et al. 2008, p. 16). However, by 2010, the ANF closed the forest to ORV use to protect the striped newt ponds (C. Petrick, USFS, personal communication, 2010) and although this stressor has not been completely removed, the activities have been significantly reduced.

Breeding ponds at Ocala National Forest (ONF) are also being impacted by off-roading. In 2010, striped newts were discovered in a pond at ONF, but subsequent surveys in multiple years have been negative. This pond has sustained ORV damage along its margins. Overall, recreational use of ORVs on ONF, ANF, and other conservation lands has seriously impacted the vegetative cover in some pond basins making it unsuitable or unattractive to striped newts (Enge 2011, pp. 9–17; Means et al. 2008, pp. 11). Many areas and trails at ONF have been closed to prevent off- road vehicle impacts to striped newt breeding ponds. An off-road vehicle and sport truck threat to wetlands on public and private lands continues despite regulations and enforcement efforts (Carrie Sekerak, 2018, pers. comm.). Greater efforts to keep vehicles out of wetlands are tasking budgets, manpower, and law enforcement capacities (Carrie Sekerak, 2018, pers. comm.).

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Impacts of ORVs are localized impacts that affect populations found only in the immediate area. Although ORV use continues to be an issue at the ONF, it has been resolved at ANF by prohibition of off-roading, and since it has not been identified in other areas as a significant stressor, we believe this stressor is unlikely to have a significant impact on striped newt populations and or the species resiliency.

3.5 Other Stressors

Other potential stressors include disease, predation, and overutilization. Disease can be difficult to detect in pond-breeding amphibians. In addition, because documenting mortality in striped newts is very unlikely, it increases the difficulty of detecting diseases that may impact the species. Some researchers believe that disease pathogens represent one of the potential causes of decline of the striped newt (Blaustein and Johnson 2003, pp. 87–92; Enge 2014, pp. 101–102; Farmer et al. 2017, p. 590). The presence of chytrid fungal infections could particularly threaten populations of striped newts, as they may not have the resiliency to recover after a population crash caused by this disease (Ouellet et al. 2005, p. 1437). However, this disease has never been identified in the striped newt. A group of viruses belonging to the genus Ranavirus has been shown to affect some local populations and cause localized die-offs of amphibians (Gray et al. 2009a, p. 244) but none have been detected in striped newts.

A disease caused by a fungus-like protist (Amphibiocystidium viridescens) has been reported in eastern newt populations (Raffel et al. 2008, p. 204). Specifically, evidence of mortality and morbidity due to infection with this disease, and the potential importance of secondary infections as a source of mortality, were reported (Raffel et al. 2008, p. 204). Also, Cook (2008a) found a striped newt in captivity to be infected with a protistan parasite that has caused disease in other species of amphibians, but there is no evidence to indicate that this protist is widespread in striped newts in the wild. This parasite, currently identified as Dermomycoides spp. (Cook 2008a, p. 2), caused disease resulting in a complete loss of recruitment of the Mississippi gopher frog population in Harrison County, Mississippi, in 2003.

Although disease is identified as a potential explanation for the extirpation of striped newts in areas of Florida and Georgia, especially in the Munson Hills in the Florida Panhandle, we are not aware of any disease-associated striped newt die-offs in wild populations (Means et al. 2008; Enge 2014, p. 190–191; Farmer et al. 2017, p. 590). Some efforts are underway for a variety of known and novel pathogens in striped newts from breeding ponds in Georgia and Florida to test for the presence of the chytrid fungi Batrachochytrium dendrobatidis and B. salamandrivorans, which have caused amphibian die-offs in other parts of the world (Martel et al. 2014; Horner et al. 2017, pp. 5–8) but currently diseases are not a significant factor affecting the species.

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3.5.1 Predation

Evidence of newt breeding has only been documented in ephemeral ponds that lack large predatory fish, including sinkhole ponds (lime sinks), depression or basin marshes, wet prairies, dome swamps, and borrow pits that are typically vegetated with emergent grasses, sedges, and forbs (Enge 2011, p.2). The cyclical nature of ephemeral ponds prevents predatory fish from inhabiting breeding ponds (Dodd and Charest 1988, pp. 87, 94; LaClaire and Franz 1990, p. 12; Moler and Franz 1987, p. 237) and the isolated breeding ponds that are being used by striped newt and that hold water all year round usually do not have a source of fish nearby that can reach the ponds if they are flooded. Although wetland changes and the introduction of predatory fish have impacted striped newts at their breeding ponds, we have not found evidence that predation is a significant stressor to the species and a combination of stressors is more likely to cause extirpations than predation alone. Predation becomes a stressor when large predatory fish are introduced or colonize striped newt breeding ponds. During wet years, flooding can occur, as ponds overflow and large predatory fish may colonize the breeding ponds impacting localized populations.

3.6 Combined Stressors Impacts from a single stressor can be exacerbated when those impacts are additive and/or cumulative and where they can have a significant effect on the species or species' populations. Although we don’t have evidence of the cumulative impact of any stressors on the striped newt and its habitat, habitat alteration and drought is a good example of a potential combination of stressors that may have an impact on the species. Droughts are a naturally occurring event in the ecology of the striped newts, but prolonged droughts can worsen threats to already small populations and exacerbate the degradation and fragmentation of striped newt habitat that may be already taking place. These situations can lead to localized extirpation of striped newts.

Another combination of stressors that are of concern if they are acting in an additive fashion is fire suppression and drought. In the event of a prolonged fire suppression of striped newt habitat, hardwood vegetation can take over ephemeral wetlands and dry them out or make them unsuitable for striped newts. Although the potential for cumulative impacts of stressors can be significantly detrimental to the striped newt, we have seen evidence of recolonization of breeding ponds after prolonged droughts and fire suppression, indicating that the species may be able to respond to some level of alterations in their habitat.

3.7 Existing Regulatory mechanisms

While many States in the southeastern United States do not regulate those activities affecting wetlands that are exempt from section 404 of the Federal Clean Water Act (CWA) (33 U.S.C.1251 et seq.), Florida is the only state known to regulate isolated wetlands. In Georgia, 48 there are no state laws that protect isolated wetlands. Lack of protection for upland habitat under wetland statutes can result in loss of recruitment of efts and paedomorphs into the terrestrial adult population, which would reduce the potential for the population to persist during extended periods of drought (Semlitsch 1998, p. 1116).

Federal statutes and regulations such as the CWA regulate the dredge and fill activities that adversely affect wetlands. Section 404 of CWA regulates the discharge of dredge or fill materials into wetlands but not the isolated ephemeral wetlands which the striped newt mostly uses. In addition, the Department of Interior through the National Wildlife Refuge System Administration Act of 1966 (NWRAA; 16 U.S.C. 668dd–668ee) administers National Wildlife Refuges for the conservation, management, and restoration of fish, wildlife, and plant resources and their habitats for the benefit of the American people. The NWRAA requires that the refuge system ensures the biological integrity, diversity, and environmental health of refuges be maintained, and requires development and implementation of a comprehensive conservation plan (CCP) for each refuge. The CCP must identify and describe the wildlife and related habitats in the refuge and actions needed to correct significant problems that may adversely affect wildlife populations and habitat (16 U.S.C. 668dd(e)). Also, striped newt habitat within national wildlife refuges is protected from loss due to urban development. Striped newts have historically been observed at St. Marks National Wildlife Refuge (SMNWR) in Florida and Okefenokee National Wildlife Refuge (ONWR) in Georgia, although in fewer numbers than previously known and/or haven’t been found since 2010 (ONWR 2010, pers. comm.; Jensen 2018, pers. comm.).

On military installations, the Department of Defense (DOD) must conserve and maintain native ecosystems, viable wildlife populations, Federal and State listed species, and habitats as vital elements of its natural resource management programs, to the extent these requirements are consistent with the military mission (DOD Instruction 4715.3). Amendments to the Sikes Act (16 U.S.C. 670 et seq.) require each military department to prepare and implement an integrated natural resources management plan (INRMP) for each installation under its jurisdiction.

Recently, the Camp Blanding Joint Training Center signed a Candidate Conservation Agreement with Assurances with the USFWS and the Florida Fish and Wildlife Conservation Commission to manage enrolled lands in the agreement to benefit multiple species, including the striped newt. Currently there are eight extant breeding ponds at Camp Blanding. The installation will maintain a 2–5 year burn schedule on approximately 10,000 acres of flatwoods and 5,000 to 16,000 acres of sandhill habitat. All ephemeral wetlands will be allowed to burn at the same time as their associated uplands during the growing season. In addition, the installation will not allow vehicle operation within known wetland areas and will maintain natural hydrology (USFWS et al. 2017, pp. 13–22).

The Forest and Rangeland Renewable Resources Planning Act (16 U.S.C. 36), of 1974, as

49 amended by the National Forest Management Act of 1976 (16 U.S.C. 1600 et seq.), requires that each national forest be managed under a forest plan which must be revised every 10 years. Regulations governing preparation of forest plans are found in 36 CFR 219. The purpose of a forest plan is to provide an integrated framework for analyzing and approving future, site- specific projects and programs, including conservation of listed species. Identification and implementation of land management and conservation measures to benefit striped newts vary between forests. For example, on the National Forests in Florida, striped newts are not designated as a species for which special management prescriptions are implemented. There are no specific land management objectives for striped newts on the National Forests in Florida. The Land and Resource Management Plan for the National Forests in Florida (U.S. Forest Service 1999, entire) provides for the restoration of longleaf pine forest through various management areas located at Apalachicola National Forest (ANF) and Ocala National Forest (ONF). Metapopulations of striped newts are found at both of these forests. However, a decline of striped newt populations at ANF has occurred over the past 10 years (Means et al. 2008, p. 6) to a point where extirpation from this area is determined likely by CPI researchers (Means et al. 2012, p.14).

Florida and Georgia have different statutes and regulations that may offer protections to the striped newt. The striped newt is not currently a State-listed species in Florida. However, ephemeral ponds in Florida have some protection under Florida State regulations. The five Water Management Districts (WMDs) and the Florida Department of Environmental Protection regulate wetland protection. The WMDs include isolated wetlands in the Environmental Resource Permit process, which requires a permit for any activities that would impact a wetland (SJRWMD 2010, p. 1). Under the WMDs permitting process, mitigation for impacts to wetlands below a minimum permitting threshold size of 0.2 ha (0.5 ac) is not addressed unless the wetland supports an endangered or threatened species, is connected by standing or flowing surface water at seasonal high water level to one or more wetlands that total more than 0.2 ha (0.5 ac), or is of more than minimal value to fish and wildlife (SJRWMD 2010, p. 1). Although the known striped newt breeding ponds ranged in size from 0.02 to 12.22 ha in Florida (median size of 0.48 ha) they may not meet all of the requirements for protection (Enge et al. 2014a, p. 57). In Georgia, a State statute requires that any rule and regulation promulgated for protected species (including the striped newt) shall not affect rights on private property or in public or private streams, nor shall such rules and regulations impede construction of any type (Ga. Code Ann. section 27-3- 132(b)). Georgia’s Endangered Wildlife Act of 1973 establishes statutory protection for protected species (Ga. Code Ann. section 27-3-130–133). Georgia Board of Natural Resources Rule (Chapter 391-4-10) mirrors the statue, but includes permitting for research under a scientific collecting permit (Ga. Code Ann. section 27-2-12). Any implementing regulations are constrained by these statutory requirements, and therefore can only prohibit collection, killing, or selling of individual newts. There are no regulatory or permitting mechanisms in place in Georgia to address habitat destruction or striped newt mortality resulting from development

50 projects on private lands. Consequently, striped newts and their habitat in private ownership in Georgia are vulnerable to ongoing and future habitat loss and mortality.

Current Federal, State, and local regulations provide some protection for striped newts and their habitat on private lands; striped newts have additional protections on conservation lands. In Georgia, striped newt habitats on private lands are not protected under State regulations, even though the striped newt is listed as threatened in that State. The status of striped newts on private lands is unknown, but the majority of the extant breeding ponds are found in conservation lands where they are offered protection from development. Regulatory mechanisms at the local, State, and Federal levels provide varying degrees of protection to wetlands, but do not protect the small, ephemeral wetlands that striped newts use for breeding sites. However, as previously stated, the majority of the current known distribution of the species is found in conservation lands. In addition, many regulations do not address management needs of the striped newt.

3.8 Current Conservation Measures

As we indicated above, current Federal, State, and local regulations do not protect the vast majority of striped newts or their habitat on private lands. In Georgia, striped newt habitats on private lands are not protected under State regulations, even though the striped newt is listed as threatened in that State. The status of striped newts on private lands is unknown, but is likely threatened by ongoing land uses such as development and silviculture. Regulatory mechanisms at the local, State, and Federal levels provide varying degrees of protection to wetlands but do not protect the small, ephemeral wetlands that striped newts use for breeding sites. However, the U.S. Forest Service (USFS) has now restricted or closed ORV use in sensitive biological communities, such as wetlands (USFS 2010, p. 1), at both ANF and ONF. ORVs have historically been a recurring issue in or around ponds at ANF and ONF. Changes at ANF and ONF have made ORVs off-limits in the Munson Sandhills and the ephemeral ponds in the ONF where striped newt ponds were being affected by ORV use (Petrick 2006, pers. comm.).

Additionally, conservation efforts initiated by the Coastal Plains Institute (CPI) in 2013 in collaboration with the U.S. Forest Service, Florida Fish and Wildlife Conservation Commission and other partners resulted in a reintroduction program in the Munson Sandhills. The program has had success releasing captive bred striped newts at breeding ponds at ANF in Florida. The program has observed juvenile efts exiting the translocation ponds in 2013–2015, observed released larvae returning to one pond as mature adults in 2015–2016, and sampled a first- generation larvae in 2016 (Ryan Means et al. unpublished. reports). At the conclusion of year 5 of the study, a total of 1,062 striped newt larva and 126 striped newt adults had been released over the length of the entire project (Means et al. 2015, page i).

Striped newt repatriation efforts and precautionary habitat measures to ensure repatriation success and enhance breeding habitat on ANF (Means et al. 2011, p. 2) have already shown

51 some success. Also, various zoos have joined the effort to captive breed striped newts collected in Georgia and Florida to be released at ANF. “Native” striped newt adults (1 male, 2 females) were captured in a single pond in the Munson Sandhill region in 2016 (R. Means, Coastal Plains Institute, personal communication, 2016). This may be evidence that habitat improvements at ANF are benefiting the existing population of striped newts and the original population of newts is slowly re-appearing.

In 2017, the Camp Blanding Joint Training Center signed a Candidate Conservation Agreement with Assurances (CCAA) with the USFWS and the Florida Fish and Wildlife Conservation Commission to manage enrolled lands in the agreement to benefit multiple species, including the striped newt. The Camp Blanding CCAA will provide protection and management of key habitat for two candidate species, the gopher tortoise and striped newt, and 20 other species, 14 of which have been petitioned for listing. It is designed to deliver conservation at the habitat scale. The Florida Armory Board will implement conservation actions on 46,507 acres and manage the six habitat types. There are 11 breeding ponds identified at Camp Blanding with eight of them currently considered extant. The installation will maintain a burn schedules on approximately 10,000 acres of flatwoods and 5,000 to 16,000 acres of sandhill habitat. All ephemeral wetlands will be allowed to burn at the same time as their associated uplands during the growing season and will maintain their natural hydrology (USFWS et al. 2017, pp. 13–22).

Another ongoing conservation action that is almost completed is the Quail Country CCAA. The Quail Country CCAA’s goal is to implement conservation actions that reduce and/or eliminate threats to the multiple species covered by the CCAA and which includes the striped newt (Quail Country CCAA, 2017, in draft). The CCAA covers non-federal properties and private lands within 5 Florida Counties (Gadsden, Jefferson, Leon, Madison, Wakulla) and 24 Georgia Counties (Baker, Brooks, Calhoun, Colquitt, Crisp, Decatur, Dodge, Dooly, Dougherty, Grady, Lee, Macon, Marion, Mitchell, Pulaski, Schley, Sumter, Talbot, Taylor, Terrell, Thomas, Webster, Wilcox, Worth). The property owners that will likely be enrolling in the Quail Country CCAA are known to have a strong land management ethic. Management actions include widespread use of prescribed fire which will benefit both upland and ephemeral wetland habitat used by the striped newt. Many of these lands also practice sustainable forestry practices, have native groundcover and longleaf pine which are suitable habitat for striped newt. The CCAA is expected to be signed this year (2018).

3.9 Summary of Species Overall Current Condition: Population Resiliency, Representation, and Redundancy

We have described the best available information on the current condition of known populations. Based on the described conditions, the resiliency of the known striped newt populations can be considered to be moderate with a 60–90% probability of persistence. There are at least 28

52 metapopulations identified with a total of 114 extant breeding ponds in 20 properties across its range. The population abundance is unknown, but evidence indicates continual changes in detection throughout the species range. The most consistently surveyed areas are locations in peninsular Florida where the stronghold of the species appears to be. The Ocala National Forest, Ordway-Swisher Biological Station, Jennings State Forest, Camp Blanding Joint Training Center, and Triple N Ranch WMA are five strongholds for the striped newt in the Eastern Region of the species. In the Florida Panhandle, the Dixie Plantation (private property in Jefferson County, FL) has a recently discovered metapopulation with four breeding ponds currently identified. Abundance in one of the Dixie Plantation breeding ponds is considered high, probably numbering in the thousands (Pierson Hill, 2017, pers. comm.) Populations across the species range have persisted through time and have shown to be resilient to long-term droughts as seen in the Ordway-Swisher example described above. Suitable habitat exists across the range of the species in private and conservation lands.

Redundancy for the striped newt is considered to be moderate. We’ve identified the potential of more than 28 metapopulations (example of some give on Figure 17) and multiple single breeding pond populations that are distributed across the range of the species. Enge et al. (2014a, pp. 51- 52) had identified at least 44 of 46 striped newt metapopulations as extant throughout Florida. We were conservative in our calculations but believe that more than 40 are extant throughout the species range. Despite the species distribution across the entire range, populations in the Western Region are the ones that show the highest decline of the species (Farmer et al. 2017). Populations at ANF are not abundant, populations at Ichauway Preserve have not been found in over five years, and translocation efforts are just showing reproduction success at ANF. There are currently 5 properties in the Western Region that were identified as extant by Farmer et al. 2017. We identified four populations as extant within the Dixie Plantation (Western Region) as a current stronghold in the Florida Panhandle.

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Eastern range

Western range

Figure 18. Potential metapopulations identified for striped newt. Dashed line divides (generally) the eastern and western range of the species.

Habitat between breeding ponds in metapopulations, as per satellite view analysis, is available in most of the connected breeding ponds found in conservation lands. We expect that connectivity within the breeding ponds in a metapopulation is occurring and expected to be taking place through passive dispersal. The long-term viability of each metapopulation will depend on the number of breeding ponds (populations) and the severity of threats that each metapopulation is facing. Genetic diversity is shown to be higher within the species strongholds in the Eastern Region, likely occurring due to the proximity of the populations with each other. The greatest stressors to the species’ redundancy are the loss of habitat from current land use changes and long-term droughts. Land use changes in Georgia have caused fragmentation of habitat near striped newt populations and may be cause for striped newt from some areas.

The striped newt is currently divided into two genetic “regions”: Western Region and Eastern Region. Hoffman (2017, pers. comm.) found that genetic variability is higher at the strongholds in the Eastern Region (peninsular Florida, eastern Georgia) but that there is relatively little

54 genetic exchange except for local genetic exchange among nearby ponds, and most likely constrained within their dispersal distance of 1km. Ecological variation occurs throughout the range where ephemeral ponds are found near sandhill uplands, flatwoods, and scrub habitat. No behavioral or morphological variation is described for the species and given the available data it does not appear to be associated with genetic variation. Environmental, precipitation, and temperature differences have been identified for the Eastern and Western Regions (May et al. 2011, p. 1447) but these have also been changing in the last decades (Enge et al. 2014a, pp. 97- 101).

CHAPTER 4 – FUTURE CONDITION SCENARIOS

4.1 Introduction

To analyze species’ viability, we consider the current and future availability or condition of resources. To examine the potential future condition of the striped newt, three future scenarios were developed. The scenarios focus on a range of conditions based on climate change scenarios and projections for land development. The range of what is likely to happen in each scenario will be described based on current condition and how resiliency, representation, and redundancy would be expected to change. The levels of certainty or uncertainty are addressed in each scenario. Given that there is uncertainty as to exact future trends of many stressors, these future scenarios are meant to explore the range of uncertainty and examine the species' response across the range of likely future conditions.

In the case of minor stressors, we have no information that the magnitude of some stressors will change from their current condition. These stressors include disease and collection. For the purpose of this assessment, we generally define viability as the ability to sustain populations over time; to do this a species must have a sufficient number and distribution of healthy populations to withstand changes in its biological (e.g. predators, disease) and physical (e.g. habitat loss, climate change) environment, and environmental stochasticity (e.g. flooding, drought, and storm surge). We chose 43 years as the time frame for our analysis because it is within the range of the available development, Florida 2060 Development Projections, 2007 and updated in 2015, (Zwick and Carr 2006; Carr and Zwick 2016) and climate change model forecasts (NOAA 2017). We looked at expected changes in development and sea level rise in 2040, 2060, 2070, and 2100 but focus on changes that are expected in the next 40–50 years.

We measure population resiliency based on the following scale:

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Overall Condition Risk Persistence over 43 Probability of years Persistence High Low Very Likely 90–100% Moderate Moderate Likely 60–90% Unlikely to likely Low High 0–60% as not

The table below (Table 6) describes three plausible scenarios where potential impacts to striped newt populations by land use changes, climate impacts and other stressors were considered.

4.2 Development – future impacts

Possible future development is very likely to continue across the range of the striped newt. In addition, the loss of habitat due to conversion of natural pine forests to more intense silvicultural management regimes will continue, at lower rates, in interior portions of the range of the striped newt. Suitable habitat that is projected to be lost in all of these scenarios is privately owned and not currently under conservation. Development models in Florida show that development has already impacted approximately 41,163 ha (101,715 ac) (16.8%) of striped newt suitable habitat. Future projections calculated using striped newt suitable landcover and the Florida 2060 Development projections (Carr and Zwick 2016; Zwick and Carr 2006) show that 14,681 ha (36,279 ac) (7.2%) will be lost by the year 2040 and 21,595 ha (53,362 ac) (10.6%) will be lost by year 2060 (Table 5). The worst case scenario with full development and no conservation actions taken into consideration throughout the development phase is projected to cause a potential loss of 29,514 ha (72,932 ac) (14.5%) of striped newt suitable habitat in Florida by 2070. Best case scenario for development, with the conservation of additional striped newt habitat, is projected to cause a 25,497 ha (63,005 ac) (12.5%) loss of striped newt suitable habitat.

In Georgia, regulatory or permitting mechanisms in place do not address habitat destruction or striped newt mortality resulting from development projects on private lands. Habitat suitability models for the striped newt were not developed for Georgia and development projections calculations specific to habitat suitability of the striped were not made. However, development trends in the southern half of the state of Georgia show development to increase from 10–20% in areas where striped newts may be found.

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Table 5. Model calculations of suitable habitat changes with various development scenarios in Florida using models from Endries et al. 2009 and Carr and Zwick 2016 Habitat Striped newt Suitable Habitat/Data Hectare Acre Lost (ac) Percent

Current: 2003–2014 (FWC Model) 244,576 604,360 101,715 16.8%

2040 Developed (fl2060) 203,413 502,645 36,279 7.2%

2060 Developed (fl2060) 203,413 502,645 53,362 10.6%

2070 Developed Trend-Worse ( fl 2070) 203,413 502,645 72,932 14.5% 2070 Developed Alternative-Best ( fl 2070) 203,413 502,645 63,005 12.5%

4.3 Drought – Future impacts

The future scenarios include the projected changes in temperature and precipitation from climate change in the southeast region. The projected increases in average annual temperature by the late 21st century (compared to the late 20th century) vary from +3 to +7 °F rangewide depending on location and the emissions scenario used. Extreme heat events in FL are projected to increase relative to 1986–2005. By the late 21st century, the average temperatures on the hottest days will be 3 ºF to 8 °F hotter. Due to the already released, human-induced emissions of greenhouse gases (GHGs) present in the environment, another +0.5 °F increase in surface air temperature would be expected, even if there was a sudden end to all human-induced GHG emissions (Carter et al. 2014, p. 25). The southeast region coastal area is projected between 2041–2070 to experience approximately thirty to forty more days a year temperatures above 95 °F, compared to recent historical levels (1971–2000) (Figure 18).

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Figure 19. Climate projections in the southeastern U.S.A. (Carter et al. 2014, pp. 399)

Major wildfires occur during drought conditions. For example, in Florida, the most significant wildfires from 1981 to 2008 occurred during dry years (Collins et al 2017, pp. 586–588). The projections for wildfire susceptibility are moderate to high throughout most of peninsular Florida and striped newt habitat (Collins et al 2017, p.587). Precipitation projections are less certain, but many models project decreases in precipitation during the summer across the range of the species (Carter et al. 2014, pp. 399.). Projections of future changes in precipitation show substantial shifts in where and how precipitation will fall. Models are in agreement regarding changes in tropical storm and hurricane rainfall events. Greater rainfall rates are expected with about a 20% increase near the center of storms. Scientists continue to research the expectation of precipitation changes in other severe storms (USFWS 2017, pp 4–5). Dry consecutive days are expected to increase up to 20% in the Eastern Region of the species by 2100. While dry conditions are expected to increase in the summer across the species range, fall conditions are expected to be wetter. This is important because it is when striped newts migrate to breeding ponds to start reproduction. The species is more likely to persist in the stronghold and in the Western Region where the presence of multiple breeding ponds in the metapopulations and a less likelihood of drier summers, respectively, will provide better conditions for the striped newt.

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Table 6. Future scenarios for the striped newt. SCENARIO 1 Land use Fire Other Conservation Climate change Likelihood change Suppression Stressors actions Less than 1% Habitat Low greenhouse of ephemeral restoration emission; Low SLR wetlands across surrounding (C1, table 4), Reduced trend or Prescribed fire in the range of the breeding ponds Stronger storms remains the managed lands species have and uplands; (increased same as current increases to predatory fish. Increased magnitude) with trend; Land better manage Unlikely ORV impacts efforts of heavier downpours, conversion for fire to habitat is repatriation of Less days higher remains the dependent minimized in striped newts at than 95 degrees F, same species all ANF and Wetter rainy seasons conservation locations in (fall) lands GA SCENARIO 2 Land use Fire Other Conservation Climate change Likelihood change Suppression Stressors actions Moderate emissions, Introduction of moderate SLR (C3, predatory fish Table4), Stronger in ephemeral Current efforts Effectiveness of storms (increased wetlands across of repatriation management on magnitude) with the range is of striped Current trend existing heavier downpours, minimal. ORV newts and Very Likely continues protected lands Temperature, impacts to restoration of continues at precipitation and habitat are habitat at ANF current range drought trends minimized in are continued. continue as current conservation trends lands SCENARIO 3 Land use Fire Other Conservation Climate change Likelihood change Suppression Stressors actions High greenhouse emissions, High SLR curve (C5, Table 4); Introduction of Increase in the predatory fish Effectiveness of average temperature in ephemeral management on 3–8 degrees F by wetlands across existing 2100; Longer Increase in the range Decrease of protected lands drought periods, current increasing efforts of decreases where Likely higher temperatures population trend predation. repatriation of persistence of days that would and development ORV impacts striped newt species and increase evaporation to habitat habitat of wetlands, more increases in deteriorates frequent stronger conservation storms (increased lands magnitude) with heavier downpours

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4.4 Scenario 1

Under Scenario 1, factors that influence populations and habitat conditions of striped newts are expected to improve. Climate change effects are predicted to be minimal under the low emissions models (USFWS 2017), so effects of increased temperatures, storms, precipitation events, and droughts counteract each other. Increased precipitation and increased storm frequency will counteract the effects of increased temperature and seasonal droughts. Land use changes are also predicted to have less of an impact in this scenario and more land is purchased for conservation. Because wetter breeding seasons and increased conservation are predicted in this scenario, it is expected that the species will maintain most existing populations, increase the number of populations in some areas, and/or increase abundance in some of the populations (Table 7, Figure 19).

Western Region. There are 5 currently extant populations in the Western Region of the species. If conditions continue to improve at Apalachicola National Forest, this site has the potential for maintaining multiple metapopulations of striped newts. Currently, the resiliency of this population is low, but under this scenario we assume that the repatriation of the breeding ponds continues to successful and expands to the other 16 historical breeding ponds. In addition, “native” populations of striped newt are expected to become more abundant and re-colonize historical breeding ponds. This scenario also considers the repatriation of striped newts in other Georgia populations such as the Ichauway Reserve where suitable habitat is managed and the re- introduction of the species would have a high likelihood of success. Pond persistence and land use change pressure would not change under this scenario for two of Georgia’s isolated populations (Sandhills WMA and Alapaha River WMA) and the single breeding pond populations are predicted to persist in these isolated conditions. Habitat conditions would stay the same or improve for the Dixie Plantation population in Jefferson County, FL, and this would result in an improvement in breeding success for this population.

Eastern Region. Currently, there are 15 properties in the Eastern Region with a total of 114 extant breeding ponds for striped newts, believed to be extirpated in 14 properties but potentially extant in more than 40 additional breeding ponds. Under this scenario, the 14 extirpated sites would stay extirpated unless striped newts are released in these sites. Habitat conditions would improve and populations in the five stronghold sites (Ocala NF, Camp Blanding Training Center, Ordway-Swisher Biological Station, Jennings SF, and Triple N Ranch), which have multiple metapopulations, would disperse into additional available breeding ponds. Habitat conditions would stay the same and/or slightly improve for all sites maintaining the extant populations. We predict that stable and/or slightly improved habitat conditions would allow for the Fort Stewart Military Installation metapopulation to increase and expand to other historical breeding ponds found at the Installation. In addition, of the properties identified as extirpated we believe at least two are possibly extant (Goethe State Forest and ONWR). Four breeding ponds were extant in

60 surveys done between 2008 and 2011 in Goethe State Forest (Enge 2011, pp. 5–25).

Table 7. Predicted responses of striped newt populations under three scenarios Current Scenario 1 Scenario 2 Scenario 3 Western Region Apalachicola National Low Moderate Low Low Forest (Leon County, FL) Private land (Jefferson Moderate High Moderate Moderate County, FL) Alapaha River WMA Low- Low Low Low (Irwin County, GA) extirpated Sandhills WMA (Taylor Moderate Low Low Low County, GA) Ichauway Reserve (Baker Presumed Possibly Possibly Low County, GA) extirpated extirpated extirpated

Current Scenario 1 Scenario 2 Scenario 3 Eastern Region Big Bend WMA (Taylor Moderate Moderate Low Low County, FL) Camp Blanding Military Reservation (Clay County, High High High Moderate FL) Faver-Dykes SP (St. Johns Moderate Moderate Moderate Moderate County, FL) Goethe SF (Alachua, Levy Possibly extant Low Extirpated Extirpated Counties, FL) Guana River WMA (St. Low Low Low Low Johns County, FL) Guana Tolomato Matanzas Presumed NERR (St. Johns County, Low Extirpated Extirpated extirpated FL) Jennings SF (Clay County, Moderate- High High High FL) Low Lochloosa WCA (Alachua Presumed Extirpated Extirpated Extirpated County, FL) extirpated Merritt Island NWR Low- Low Low Low (Volusia County, FL) Extirpated Ocala National Forest (Lake, Marion, Putnam High High High High Counties, FL)

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Ordway-Swisher Moderate- Biological Station (Putnam High High High High County, FL) Private land (Gilchrist Presumed Extirpated Extirpated Extirpated County, FL) extirpated Private land (Leon County, Presumed Extirpated Extirpated Extirpated FL) extirpated Private land (Putnam Presumed Extirpated Extirpated Extirpated County, FL) extirpated Private land (St. Johns Presumed Extirpated Extirpated Extirpated County, FL) extirpated Pumpkin Hill Creek SP Low Low Low Low (Duval County, FL) Rock Springs Run SP Low- (Lake, Orange Counties, Low Low Low extirpated FL) Seminole SF (Lake County, Low Low Low Low FL) Triple N Ranch WMA High High High Moderate (Osceola County, FL) University of Central Presumed Florida (Orange County, Extirpated Extirpated Extirpated extirpated FL) Fort Stewart Military Installation (Bryan, Low Moderate Low Low Liberty, Long Counties, GA) Ohoopee Dunes Natural Area (Emanuel County, Low Low Low Low GA) Okefenokee NWR Possibly extant Low Extirpated Extirpated (Charlton County, GA) Private land (Bryan Presumed Extirpated Extirpated Extirpated County, GA) extirpated Private land (Camden Moderate Moderate Moderate Low County, GA) Private land (Charlton Presumed Extirpated Extirpated Extirpated County, GA) extirpated Private land (Jenkins Presumed Extirpated Extirpated Extirpated County, GA) extirpated Private land (Screven Presumed Extirpated Extirpated Extirpated County, GA) extirpated

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Private land (Wilcox Presumed Extirpated Extirpated Extirpated County, GA) extirpated

4.4.1 Resiliency

Scenario 1 projects conditions beneficial to striped newt populations and striped newt breeding conditions if current conservation actions improve the populations by the predictive time horizon. In this scenario, all populations that are extant are predicted to remain extant (Table 7), a few could become extant, and for some their condition would improve slightly. Specifically, stronghold populations in the Eastern Region of the striped newt are predicted to be highly resilient and to remain as the stronghold of the species and disperse into additional historical breeding ponds. The ANF and the Dixie Plantation populations in the Western Region are predicted to be moderately and highly resilient, respectively, while twelve other sites are predicted to be characterized by populations with low resiliency. No extirpations that have not already occurred are predicted under this Scenario.

4.4.2 Representation

Under Scenario 1, it is predicted that the striped newt will retain current levels of representation and potentially slightly increase its level of representation in the Western Region with repatriation efforts. As such, the species will continue to remain extant in the Eastern and Western Regions. At the population level, metapopulations in the species strongholds in Florida are predicted to have high resiliency, while the remaining isolated populations will have moderate to low resiliency (Table 7, Figure 19).

4.4.3 Redundancy

Under Scenario 1, it is predicted that the striped newt will maintain existing levels of redundancy, with varying resiliency in the Eastern and Western Regions of the species. The five stronghold locations in the Eastern Region and the Dixie Plantation population in the Western Region are predicted to have multiple, highly resilient populations. The species is represented at 23 of the 34 historical sites and in both the Eastern and Western Regions.

4.5 Scenario 2

Under Scenario 2, factors that influence populations and habitat conditions of striped newts are expected to stay the same or continue at current trends. Under the current greenhouse emissions projections, climate change is predicted to be moderate, SLR projections are moderate (C3, Table3), stronger storms (increased magnitude) with heavier downpours are expected; and wetter rainy seasons (fall) are anticipated, along with increases in temperature and prolonged droughts

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(USFWS 2017). Land use changes are expected to continue at the current projections with a potential loss of up to 14% of suitable striped newt habitat (Table 5) and with no changes or with insignificant changes to the current lands in conservation. A wetter breeding season is expected but may not be enough to counteract frequent prolonged droughts (Table 7, Figure 19). In this scenario, repatriation and restoration efforts at Apalachicola National Forest continue but no other repatriation efforts are added.

Western Region. There are five properties currently with extant populations in the Western Region of the species. Under Scenario 2, repatriation efforts at Apalachicola National Forest continue and this site has the potential for maintaining and improving populations of striped newts. Currently, the resiliency of the population at ANF is low, but under this scenario we assume that the repatriation of the breeding ponds and the recolonization of native striped newts continue successfully and striped newt numbers increase in the extant breeding ponds. Pond persistence and land use change pressure is expected to increase slightly under this scenario. Two of the Western Region populations in Georgia are isolated single breeding pond populations (Sandhills WMA and Alapaha River WMA); these populations are predicted to persist in these isolated conditions. The Dixie Plantation population in Jefferson County, FL is under conservation and active fire management; we predict that under this scenario this metapopulation persists.

Eastern Region. Under this scenario, the populations at the 15 sites with extant populations would persist. Habitat conditions would vary from the wetter rainy season to more frequent prolonged droughts, but it is uncertain how more frequent storm events with heavy downpours would impact striped newt habitat. We predict that water will be retained longer during the wetter periods and striped newt will shift their breeding season where needed to breed successfully. In the five stronghold sites (Ocala NF, Camp Blanding Training Center, Ordway- Swisher Biological Station, Jennings SF, and Triple N Ranch) under this scenario, water will be available in multiple breeding ponds in most of the breeding seasons and striped newt populations will persist. It is too uncertain to say if breeding ponds will stay wet for 6 months at a time, but those areas with metapopulations and multiple breeding ponds in each site will be where striped newts are more likely to breed successfully and persist. We predict habitat conditions will stay the same and or slightly deteriorate for all sites. These conditions will allow for all extant populations in the Eastern Region to persist.

4.5.1 Resiliency

Scenario 2 projects the continuation of current conditions or slightly deteriorated conditions for the timeframe analyzed. Striped newt breeding conditions will be optimal to good, and striped newt populations (especially areas with metapopulations) will persist at current numbers. Because changes predicted for this scenario impact only a few of the known populations and

64 breeding ponds, all populations that are extant are predicted to remain extant (Table 7, Figure 19), and recolonization is not expected for any of the extirpated populations. Specifically, stronghold populations in the Eastern Region of the striped newt are predicted to be highly resilient to drought conditions and to remain as the stronghold of the species. The ANF and the Dixie Plantation populations in the Western Region are predicted to have a low resiliency and moderate resiliency, respectively, while twelve other sites are predicted to be characterized by populations with low resiliency. No extirpations that have not already occurred are predicted under this scenario.

4.5.2 Representation

Under Scenario 2, it is predicted that the striped newt will retain current levels of representation in the western and Eastern Region. Striped newts will continue to remain extant in the Eastern and Western Regions with potential breeding decrease during prolonged droughts. At the population level, metapopulations in the species strongholds in GA and FL are predicted to have high resiliency, while the remaining isolated populations will have low to moderate resiliency (Table 7, Figure 19). Even with good habitat, populations at the Ichauway Preserve are believed to be “possibly extirpated.” Under this scenario, we predict that if striped newts are still found at Ichauway they will recolonize the breeding ponds.

4.5.3 Redundancy

Under Scenario 2, it is predicted that the striped newt will maintain existing levels of redundancy with varying resiliency in the Eastern and Western Regions of the species. The five stronghold locations in the Eastern Region and the Dixie Plantation population in the Western Region are predicted to continue to have multiple, highly resilient populations. The species will be represented at 23 of the 34 historical sites and in both the Eastern and Western Regions. None of the extant populations are predicted to be extirpated.

4.6 Scenario 3

Under Scenario 3, weather conditions and habitat conditions for striped newts are expected to deteriorate significantly. Under the increased greenhouse emissions projections, climate change is predicted to cause high SLR (C5, Table 3), stronger storms (increased magnitude) with heavier downpours, wetter rainy seasons (fall), increase in the average temperature up to 8 ºF, more frequent 95 ºF or higher days, and more frequent prolonged droughts (USFWS 2017). In this scenario, there is an increase in development and other land use changes for a potential loss of up to 20% of striped newt suitable habitat (Table 5) and no additional lands in conservation. A wetter breeding season is expected but may not be enough to counteract frequent prolonged droughts (Table 7). In this scenario, repatriation and restoration efforts at Apalachicola National

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Forest are minimal and no other repatriation efforts are pursued.

Western Region. In Scenario 3, of the five (5) properties with currently extant populations in the Western Region of the species, repatriation efforts at Apalachicola National Forest continue, but with no additional restoration efforts. We predict that SLR will not impact any of the populations in the Western Region. Also, we predict that populations of striped newt at ANF will have low resiliency. Populations at ANF are believed to currently have low resiliency, and under this scenario we predict that the populations stay at low abundances. Pond persistence and land use change pressure is expected to increase under this scenario. Two of the Western Region populations in Georgia are isolated single breeding populations (Sandhills WMA and Alapaha River WMA) but are located in conservations lands. We predict that these populations persist in these isolated conditions in low abundances due to expected wetter conditions but may be extirpated if long drought periods occur frequently. The Dixie Plantation population in Jefferson County, FL, is considered to have high resiliency; under this scenario, we predict that populations at this site will persist due to the wetter conditions predicted for the Western Region. Populations at Ichauway Preserve stay extirpated.

Eastern Region. Under this scenario, we are less certain about the future habitat conditions in the Eastern Region of the striped newt, especially for the isolated populations and those areas with high land use change pressure. With varying habitat conditions, wetter breeding seasons may or may not counteract prolonged drought periods. In events of frequent storm events that bring heavy rains, striped newt breeding ponds may stay wet long enough for newts to complete their life cycle. We are less certain of the species response under this scenario in the Eastern Region due to lower precipitation forecasts but we predict that water will be retained long enough (5–6 months) in existing breeding ponds for striped newts to complete their life cycle and metamorphose in time to begin their terrestrial phase. In the five stronghold sites (Ocala NF, Camp Blanding Training Center, Ordway-Swisher Biological Station, Jennings SF, and Triple N Ranch), we predict that under this scenario, that water will be available in multiple breeding ponds for most of the breeding seasons and striped newt populations will persist by taking advantage of the hydrological differences of the multiple ponds. Habitat conditions will deteriorate in open areas where drought and higher temperatures will evaporate water more quickly, and those populations with isolated ponds will not be able to complete the breeding cycle every year. In addition, sea level rise is predicted to impact the coastal populations and at least one (Guana River WMA) striped newt location and one of the populations will be lost to sea level rise (Fig 14).

4.6.1 Resiliency

Scenario 3 projections indicate that current conditions will deteriorate in the analyzed timeframe. It is uncertain, but we predict striped newt breeding conditions to deteriorate in some of the

66 current sites. Specifically, sea level rise will inundate a population of the Guana River WMA site and will increase salt water intrusion in others. Isolated populations that have high land use changes surrounding the populations will have bigger changes in habitat conditions. Changes under this scenario will impact most of the known populations and breeding ponds, but all of the populations found in conservation lands that are extant are predicted to remain extant (Table 7). Stronghold populations are highly resilient to drought conditions, and they will remain as moderately resilient under this scenario. The ANF and the Dixie Plantation populations in the Western Region are predicted to have a low and moderate resiliency, respectively, the metapopulation nature of these sites have built-in redundancy and the species can recolonize breeding ponds after drought periods. For ANF, extirpation is likely to occur if translocation efforts do not continue and/or abundant populations aren’t established in multiple breeding ponds. Extirpation of the Guana River WMA and the population in private property in Camden County, GA is very likely to occur.

4.6.2 Representation

Under Scenario 3, it is predicted that the striped newt will retain lower levels of representation in the Eastern and Western Regions. Striped newts will continue to remain extant in the Eastern and Western Regions with potential breeding decreases during prolonged droughts and extirpation from two known sites (Guana River WMA and Camden County, GA). At the population level, metapopulations in the species strongholds in GA and FL are predicted to have moderate resiliency, while the remaining isolated populations will have low resiliency (Table 7) and will persist. Even with good habitat, populations at the Ichauway Preserve are believed to be “possibly extirpated.” Under this scenario, as well as in Scenario 2, we predict that if striped newts are still found at Ichauway they will recolonize the breeding ponds. Because over 85% of the known extant populations are found in conservation lands, land use changes will have a lesser impact on the species and the species will still be represented throughout its entire range.

4.6.3 Redundancy

Under Scenario 3, it is predicted that the level of redundancy for striped newt populations will decrease, with varying and decreased resiliency in the Eastern and Western Region of the species. The five stronghold locations in the Eastern Region and the Dixie Plantation population in the Western Region are predicted to continue to have multiple resilient metapopulations. The species will be represented at 20 of the 34 historical sites and in both the Eastern and Western Regions, only two of the extant populations are predicted to be extirpated.

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Figure 20. Summary of forecasted population condition expected under the three future condition scenarios examined.

CHAPTER 5: SYNTHESIS

We have determined that the habitat and population loss attributable to development and sea level rise is not likely to significantly reduce existing populations of striped newt. The vast majority of known populations of striped newt occupy conservation lands that are managed at varying degrees with prescribed fire as a management tool. Habitat loss and sea level rise will very likely impact less than 25% of the currently known suitable striped newt habitat and less than 10 of the known populations. We have moderate to low confidence in the known current condition of populations across the range, but predictions under the different scenarios show that all but five of the properties with extant populations will likely be extirpated in scenario 3; two in the Western Region and three in the Eastern Region. The species has populations distributed across all of its historical range with a lower representation in the Western Region of the species but with more breeding ponds being identified and repatriation efforts underway at ANF, Florida. Drought is expected to play a significant role as a stressor for this species but we are unclear how the species will adapt to the prolonged droughts or how resilient it is. Habitat development and sea level rise projections for the next 43 years will inundate one of the

68 populations at Guana River WMA and may impacts a few others with salt intrusion. We have a low confidence in the likelihood of how many populations of striped newt are predicted to be extirpated in the future, but the current development and sea level rise projections only identify two possible populations that could be extirpated.

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