DRAFT Biological Status Review Report Striped Newt

March 11, 2021

FLORIDA FISH AND WILDLIFE CONSERVATION COMMISSION 620 South Meridian Street Tallahassee, Florida 32399-1600 DRAFT Biological Status Review Report for the Striped Newt (Notophthalmus perstriatus) March 11, 2021

EXECUTIVE SUMMARY

The Florida Fish and Wildlife Conservation Commission (FWC) was petitioned by Anna Farmer on March 12, 2019 to list the striped newt (Notophthalmus perstriatus) as Threatened. Furthermore, the petitioner requested that if the striped newt failed to meet the listing criteria for Threatened, that western populations (referred to as the panhandle population in this report) be evaluated separately because recent genetic evidence suggests they are a separate evolutionarily significant unit from eastern populations. The members of the Biological Review Group (BRG) met on June 26, 2020. Group members were Kevin Enge (FWC lead), Aubrey Greene (FWC), Steve Johnson (University of Florida), Ryan Means (Coastal Plains Institute), and Dirk Stevenson (Altamaha Environmental Consulting) (Appendix 1). In accordance with rule 68A- 27.0012, Florida Administrative Code (F.A.C.), the BRG was charged with evaluating the biological status of the striped newt using criteria included in definitions in 68A-27.001, F.A.C., and following protocols in the Guidelines for Application of the IUCN Red List Criteria at Regional Levels (Version 3.0) and Guidelines for Using the IUCN Red List Categories and Criteria (Version 8.1). Information on the listing process and criteria for listing as defined in rule can be found at https://myfwc.com/wildlifehabitats/wildlife/listing-process/.

The BRG concluded from its biological assessment that statewide the striped newt met 2 criteria for designation as a Threatened species. Prior to completing the statewide assessment, the BRG opted to conduct a separate biological assessment of the panhandle population, in case subsequent input and review of its findings conclude that the striped newt as a species does not meet any of the criteria for listing as Threatened. The BRG determined that the panhandle population of the striped newt met 4 criteria for designation as a Threatened population. FWC staff recommends the statewide population of the striped newt be listed as a Threatened species.

Independent scientific review of the initial draft of the biological status review (BSR) report was sought and received from 3 scientists. All reviewers agreed that the striped newt meets criteria for listing as Threatened at a statewide level. Information and comments provided by peer review have been incorporated into this document. For a summary of peer reviewer comments, see Appendix 2.

FWC staff gratefully acknowledges the assistance of the external BRG members and BSR report peer reviewers, C. Kenneth Dodd Jr., Sarah May, and Lora Smith. The BRG in turn thanks Robin Boughton, Craig Faulhaber, Brad Gruver, Kelly O’Connor, Bradley O’Hanlon, Natalie Montero, Dan Sullivan, and Claire Sunquist for their guidance and assistance.

Striped Newt Biological Status Review Report 2 BIOLOGICAL INFORMATION Taxonomic Classification – The striped newt (Notophthalmus perstriatus) was formally described by Bishop (1941) as Triturus perstriatus from specimens collected in Alachua and Leon counties, Florida, and Charlton County, Georgia. Carr (1940) referred to the striped newt as Triturus viridescens symmetrica (Harlan). Based on mitochondrial DNA (mtDNA) analyses of concatenated ND4 and 16S genes, and a separate analysis of the Cytochrome-B gene, the striped newt is sister to Florida specimens of the eastern newt (N. viridescens) (Matthew Fedler, FWC, pers. commun. 2020). This contradicts allozyme-based findings of Reilly (1990), who claimed that the striped newt is phylogenetically more closely related to the black-spotted newt (N. meridionalis) from southern Texas and Mexico than to the sympatric eastern newt. Several recent studies of salamander phylogeny found that results from allozyme analysis were inconsistent with those from modern DNA sequencing and analysis (Smith et al. 2018, Joyce et al. 2019). A study of mtDNA variation found that striped newt populations occur in 2 disjunct geographic regions (separated by approximately 125 km) that comprise separate evolutionarily significant units (May et al. 2011). Although these 2 regions do not share haplotypes, they have not genetically diverged sufficiently to represent separate species (Johnson 2002a, May et al. 2011). Since then, the species has been found in Taylor County, Florida, narrowing the purported distributional gap (Mays and Enge 2014), and the low-lying region along the Aucilla River apparently separates the eastern and western lineages (i.e., clades) in Florida. The Aucilla River also separates genetic lineages of the gopher frog (Rana capito) (Enge et al. 2017), which uses similar upland and wetland habitats. Studies of microsatellites and single nucleotide polymorphisms (next generation sequencing or ddRADSeq) supported genetic differentiation between eastern and western sites and found an overall pattern of isolation-by-distance (Farmer et al. 2018, Hoffman et al. 2019). Genetic clusters were present in the eastern clade in both states and in the western clade in Florida, but no gene flow existed between western clade sites in Georgia (Hoffman et al. 2019). Hereafter, the western clade will be referred to as the panhandle population and the eastern clade as the peninsular population in Florida. Life History and Habitat Requirements – Information on the species has been summarized by Dodd et al. (2005) and Enge (2019). Terrestrial adults typically migrate to breeding ponds during rains in fall and winter (some newts may migrate in spring or summer), and courtship, breeding, and oviposition occur in the water (Johnson 2002b). During the breeding season, male newts deposit spermatophores on the substrate that are subsequently picked up by female newts in their cloaca (Petranka 1998), A female may take several months to lay all her eggs (Johnson 2005), which are attached singly or in small clumps to aquatic vegetation or other objects (Carr and Goin 1955). When a larva is ca. 6 months old and measures 1.8‒3 cm snout-vent length (SVL), it either changes into an eft or remains in the pond until maturing ca. 6 months later into a paedomorph (Johnson 2002b). Efts are terrestrial juveniles, and paedomorphs are gilled adults. Sexual maturity is reached at ca. 2.5 cm SVL (Johnson 2002b). At a breeding pond in northern peninsular Florida, larvae required a minimum hydroperiod of 139 days before metamorphosing into efts (Dodd 1993). In another ephemeral wetland at the same site, Johnson (2002b) found that a striped newt paedomorph apparently only breeds once before transforming into a terrestrial adult (Johnson 2002b). This differs from other neotenic salamanders, including the mole salamander (Ambystoma talpoideum), that typically continue breeding as paedomorphs as long as water remains in the wetland. Based on the

Striped Newt Biological Status Review Report 3 presence of populations of paedomorphic striped newts in permanent wetlands situated in presumably unsuitable upland habitats, such as dense sand pine (Pinus clausa) scrub, paedomorphs may exhibit reproductive plasticity. Non-gilled aquatic adults have never been found in these scrub wetlands (Kevin Enge, FWC, pers. obs.), suggesting the absence of terrestrial life stages. Additional evidence supporting the possibility of multiple reproductive episodes by paedomorphs in at least some populations is the wide variation in size of paedomorphs observed in some ponds, indicating that some paedomorphs are likely older than 1.5 years. Also, striped newts maintained in captivity in totally aquatic environments may breed more than once while retaining their gills (Mark Mandica, Amphibian Foundation, pers. commun. 2020). Little is known regarding feeding habits of terrestrial adults and efts. Aquatic adults opportunistically feed upon frog eggs, fairy shrimp, spiders, and larval and adult insects (Christman and Franz 1973, Dodd et al. 2005). Striped newts are primarily found in xeric upland habitats, most often sandhill but also in scrub, scrubby flatwoods, and adjacent mesic flatwoods. In Osceola County, Florida, newts apparently use mesic flatwoods and dry prairies besides patches of scrub and scrubby flatwoods (Enge et al. 2020). Primary upland habitats around 111 striped newt ponds in Florida were sandhill (59.5%), scrub (16.2%), scrubby flatwoods (7.2%), mesic flatwoods (7.2%), upland pine forest (4.5%), dry prairie (1.8%), xeric hammock (1.8%), and disturbed (1.8%) (Kevin Enge, unpubl. data). Habitat types are based on the Florida Cooperative Land Cover Map (version 3.0). Striped newts are rare or absent in unburned sandhills that have been invaded by hardwoods (Greenberg et al. 2003). Striped newts migrating from a pond favored the direction leading to sandhill habitat and were found up to 709 m from the nearest wetland (Dodd and Cade 1998). Johnson (2003) found that 16% of striped newts in his study migrated more than 500 m from ponds. Little is known regarding the terrestrial existence of efts and adults, although newts (primarily efts) are occasionally found under logs (Enge 2019). Striped newts typically breed in ephemeral or semipermanent sinkhole ponds, marshes, dome swamps, and borrow pits that lack large, predatory fish species (Christman and Means 1992, Dodd et al. 2005, Enge et al. 2014a) and have diverse herbaceous vegetation, including maidencane (Panicum hemitomon), witch grasses (Dichanthelium spp), spikerushes (Eleocharis spp.), lesser creeping rush (Juncus repens), and dotted smartweed (Persicaria punctata). Wetlands used by striped newts in Florida (N = 111) were depression marsh (58.6%), basin marsh (13.5%), dome swamp (13.5%), sinkhole pond (11.7%), borrow pit (1.8%), and sandhill lake (0.9%) (Kevin Enge, unpubl. data). These natural wetland habitat types are defined by the Florida Natural Areas Inventory (2010), except for sinkhole pond, which refers to a deep depression marsh that rarely, if ever, dries. In scrub habitat that is presumably unsuitable for survival of efts and terrestrial adults, breeding ponds are typically permanent and continuously inhabited by paedomorphs and/or larvae. Alternatively, scrub ponds are depressions in large wet prairie systems, predominately vegetated by blue maidencane (Amphicarpum muehlenbergianum), that are typically dry and may provide suitable upland habitat (Enge et al. 2014a). Nonpredatory fishes were present in 23.6% of 106 striped newt ponds surveyed in Florida (Kevin Enge, unpubl data). Striped newts and other Notophthalmus spp. have lifespans of ca. 12‒15 years (Forester and Lykens 1991), and a striped newt lived >17 years in captivity (Wallace et al. 2009). Some adults at the Jacksonville Zoo and Gardens are estimated to be 13‒19 years old (Means et al. 2017). This long lifespan enables populations to survive unfavorable stochastic environmental events,

Striped Newt Biological Status Review Report 4 such as prolonged droughts, that affect reproduction (Dodd 1993, Dodd et al. 2005). Striped newts recolonized a pond that had been dry for 10 years (Dodd and Johnson 2007). Population Status and Trend – The U.S. Fish and Wildlife Service (USFWS) was petitioned by Means et al. (2008) to list the striped newt as Threatened under guidelines of the Endangered Species Act because of an enigmatic population decline in the Munson Sandhills of the Apalachicola National Forest (ANF), coupled with apparent declines in much of the rest of its global range, long-term droughts, groundwater withdrawal, degradation of wetlands by off- road vehicles, and upland and wetland habitat degradation from fire suppression. The 12-month finding determined that the striped newt warranted listing because of the threats of habitat loss, disease, inadequate regulatory mechanisms, and drought (USFWS 2011). It concluded that regulatory mechanisms were inadequate to prevent further loss of breeding ponds (Factor D) throughout the striped newt’s range, and existing regulations also did not protect striped newts on private lands. Long-term regional droughts (Factor E) had a negative impact on the long-term persistence of striped newts. The USFWS found that listing the striped newt was warranted in 2011, but this action was precluded by higher priority listings. The federal listing status of the striped newt remained as “warranted but precluded” for 7 years until the USFWS (2018) determined that it did not currently warrant listing. Reasons stated for this status change were 1) threats to the species are of lower magnitude than previously thought; 2) new populations have been discovered, resulting in increased resiliency, redundancy, and representation; 3) past conservation efforts, including captive rearing and release of striped newts, have helped reestablish populations in ANF; and 4) 85% of populations currently occur on conserved lands (USFWS 2018). During a 1990‒1995 survey of 297 ponds in Florida, Franz and Smith (1999) found striped newts in only 38 ponds in the 5 known strongholds plus Faver-Dykes State Park and Rock Springs Run State Reserve. They documented striped newts in 4 ponds at 30 historical sites and at 34 new ponds. Franz and Smith (1999) believed that the paucity of recent records strongly suggested a serious decline throughout its Florida range and recommended both state and federal listing. In 2000–2003, Johnson and Owen (2005) visited 51 historical sites (64 ponds) in 11 counties in peninsular Florida and ranked the habitat quality of the wetland and surrounding uplands in terms of their potential to support newts. They ranked 26 sites in Clay, Marion, and Putnam counties as excellent, but they ranked 22 sites in 9 counties as having moderate to no potential to support newts. A survey of 227 ephemeral ponds on commercial forest lands in northern Florida in 1996−1998 found striped newts in only 4 ponds in Clay and Putnam counties (Wigley et al. 1999), suggesting that some aspect of commercial forestry is incompatible with this species. Striped newts have not been found on private property in the eastern portion of the Munson Sandhills since the 1980s when it was converted to a dense sand pine plantation (Means and Means 1998, Means and Means 2005). Surveys of 265 ephemeral ponds in the Munson Sandhills of ANF in 1995‒2007 located 18 breeding ponds (Means and Means 1998), but no larvae and only 3 adults were found in 2 ponds during surveys in 2001‒2008 (Means et al. 2008, U.S. Fish and Wildlife Service 2011). Extensive resurvey efforts in the Munson Sandhills in 2011 and 2012 (Means et al. 2012) failed to detect any striped newts. A population that was present in the late 1970s at St. Marks National Wildlife Refuge in Wakulla County is apparently extinct, despite the presence of suitable habitat (Dodd et al. 2007). By 2012, in response to rapid enigmatic extirpation of the striped newt in ANF, as well as to an apparent global decline, a partnership of researchers, zoos, and agencies teamed up to create a

Striped Newt Biological Status Review Report 5 proactive conservation strategy for the striped newt within the former ANF stronghold. This multiphase project would proceed with repatriations of captive-raised newts back into select historical ANF breeding ponds. At the inception of the striped newt repatriation project, western clade striped newts could reliably be found in only 1 pond in Georgia (Means et al. 2012). This site became the source for the establishment of assurance colonies within several zoos in the Southeast. F1 offspring produced by parents from the assurance colonies were routinely released into recipient ANF ponds from 2013 through 2020. In 2015, 2 western clade striped newt ponds were discovered in Jefferson County, Florida (Hill and Sash 2015). In 2016, a relict population was rediscovered in a single ANF breeding pond, where striped newts had last been detected in 2006 (Means et al. 2016, Farmer et al. 2017). Because of these discoveries of additional striped newt breeding ponds in the Florida panhandle, 3 genetic stocks are now available to source individuals for repatriation efforts. Repatriation results in ANF have shown some glimpses of success, including observations of reproduction, recruitment, and a few returning adults. However, more years of intensive repatriation and monitoring efforts will be needed to establish robust, self-sustaining populations (Ryan Means, Coastal Plains Institute, pers. commun. 2020). Farmer et al. (2017) compiled 44 locational records (9 breeding ponds) in 14 Florida counties prior to 1980. They documented 83 locations (77 breeding ponds) in 8 counties in 1980‒1999 and 106 locations (all breeding ponds) in 13 counties in 2000‒2016. Dipnet surveys in 2010‒2014 found striped newts in 22 historical and 24 new ponds in 8 counties but failed to find newts in 18 known ponds at ANF, 12 ponds at Ocala National Forest (ONF), 9 ponds at Ordway-Swisher Biological Station, 8 ponds at Camp Blanding Military Reservation, and 5 ponds at Jennings State Forest (Enge et al. 2014a). These results suggest that striped newt populations may be declining in population strongholds, but low detection probabilities and low water levels some years may have accounted for the lack of success. Although striped newts have been recorded since 2005 from most known ponds in ONF, 3 subpopulations (not included in Table 2) may now be extinct (Enge et al. 2014a). Dodd (1993) documented a population decline at a pond on Ordway Swisher Biological Station due to persistent drought, but the pond was recolonized 10 years later (Dodd and Johnson 2007). Coastal Plains Institute staff documented population extirpations at 17 ponds in ANF between 1998 and 2012 (Means et al. 2008, Means et al. 2012). This decline is believed to be the result of multiple factors working in unison, including upland and wetland habitat degradation from fire suppression or alteration, long-term drought, or potentially an unidentified pathogen. Direct evidence of fire ant predation on emergent striped newt metamorphic individuals also has increased during striped newt repatriation project operations (Means et al. 2015; Ryan Means, pers. commun. 2020). The number of striped newt populations has declined as the human population has increased and converted suitable habitat to urban, agricultural, and other land uses. Anthropogenic habitat conversion has destroyed approximately half of Florida’s historical wetland coverage (Dahl 2005) and modified over 98% of historical longleaf pine (Pinus palustris) forests, threatening populations of wetland-dependent fauna (Meshaka and Babbitt 2005). Means and Means (1998) claimed that urban sprawl and land conversion since 1824 had extirpated the species from the Tallahassee Red Hills region, but a subpopulation has since been discovered in northern Jefferson County (Hill and Sash 2015). The first record of the striped newt in Florida was in 1922 from Lake Jackson north of Tallahassee, Leon County, and the first breeding pond in the Tallahassee Red Hills was discovered in 1969 (Enge et al. 2014a). This pond was dredged, deepened, and stocked with game fish in the 1980s, extirpating the striped newt population

Striped Newt Biological Status Review Report 6 (Means and Means 1998). Numerous historical striped newt records once occurred around Gainesville, Alachua County, but the only recent records in the county are from the southwestern corner in the Watermelon Pond area, where the species was last documented in 2010 (Farmer et al. 2017). Urban sprawl prevents movement of newts between breeding ponds, decreasing genetic variability of newts and making them more vulnerable to extinction. A striped newt habitat model for Florida identified 244,576 ha of potential habitat, but only 38% was located on managed lands (Endries et al. 2009). A subsequent habitat model using more accurate landcover imagery and more data points identified 158,706 ha of potential habitat, 88.4% of which was considered good (Enge et al. 2014a). Conservation lands contained 50.2% of the total habitat and 49.4% of the good habitat, with 104 conservation lands containing at least 50 ha of potential habitat (Enge et al. 2014a). Geographic Range and Distribution – The range of the striped newt extends from southern Georgia to central Florida from the Atlantic Coast westward to Leon and Wakulla counties in Florida and Taylor County in southwestern Georgia (Farmer et al. 2017). In peninsular Florida, the range extends east to Nassau County and south along the Atlantic coast to Merritt Island National Wildlife Refuge in Volusia County (Enge et al. 2015). Mid-peninsular populations extend south to Half Moon Wildlife Management Area (WMA) in Sumter County (Johnson and Dwyer 2000) and Triple N Ranch WMA in Osceola County (Enge et al. 2014b, 2020). The striped newt occurs on the northern Brooksville Ridge but is apparently absent from large tracts of sandhill habitat farther south in Citrus and Hernando Counties, such as tracts of Withlacoochee State Forest. However, a 1930s record from Floral City, Citrus County (A. Carr, University of Florida, field notes), suggests that the species might have historically occurred farther south on the Brooksville Ridge. Recent discoveries of striped newts in the Spring Creek Unit of Big Bend WMA, Taylor County (Mays and Enge 2014), and Livingston Place (formerly named Dixie Plantation) in Jefferson County (Hill and Sash 2015) have partially filled in the purported distributional gap between panhandle and peninsular populations. The species’ historical range was probably similar to its current range (Fig. 1), but many populations have likely been lost because of extensive habitat modification by humans (Dodd et al. 2005, Farmer et al. 2017).

Striped Newt Biological Status Review Report 7

Fig. 1. Locality records for the striped newt (from Enge 2019). Blue symbols are records pre- 1980 and orange symbols are records 1980‒2018. Circles represent voucher specimens, triangles represent credible but unvouchered records, and squares represent county record.

Quantitative Analyses – A population viability analysis (PVA) apparently does not exist for the striped newt. An attempt at conducting a PVA was unsuccessful, so a potential habitat model was created instead for Florida (Endries et al. 2009). Breeding wetland habitat was limited to patches <9 ha in size, and primary upland habitats were included within 1,000 m of breeding habitat, which restricted upland habitat patches to 79 ha. The GIS analysis found 244,576 ha of potential habitat; 122,724 ha occur on 124 sites on public lands, but only 64 of these sites have >40 ha of potential habitat (Endries et al. 2009). Of the potential habitat on public lands, 55% occurs on ONF, 8% on Camp Blanding Military Reservation, 6% on Withlacoochee State Forest, 5.3% on ANF, and 2.9% on Jennings State Forest (USFWS 2011). Enge et al. (2014a) developed a Maxent potential habitat model that identified 158,706 ha of potential habitat, 88.4% of which was considered good quality. The variables land cover and distance to sandhill had the strongest positive influence on the model, whereas elevation and canopy cover had the least influence. Conservation lands, including state, federal, or local lands and conservation easements that retain lands in their natural condition, contain 50.2% of the total habitat and 49.4% of the good habitat, with 104 conservation lands containing at least 50 ha of potential habitat. Ocala National Forest has 3.2 times more potential habitat (22,644 ha) than the conservation land in second place, Withlacoochee State Forest, where striped newts have not been found. The amount of potential habitat in the other 4 strongholds for the species is 4,127 ha at Camp Blanding Military Reservation, 3,332 ha at ANF, 1,128 ha at Jennings State Forest, and 819 ha at Ordway-Swisher Biological Station (Enge et al. 2014a).

Striped Newt Biological Status Review Report 8

BIOLOGICAL STATUS ASSESSMENT Threats –The greatest threat to striped newts is loss and alteration of xeric upland habitats resulting from commercial and residential development, silviculture, agriculture, and mining (USFWS 2011, Farmer et al. 2017). Intact xerophytic upland ecosystems inhabited by striped newts have suffered severe losses in Florida, particularly longleaf pine-dominated sandhill (Means and Grow 1985, Myers 1990, Kautz 1998, FWC 2006). Infrequent and/or dormant season fire in sandhill and upland pine forests habitats leads to degradation of the upland and wetland habitats critical for striped newts. In an altered fire regime in these xeric upland habitats, woody-stemmed, broad-leafed vegetation eventually replaces the herbaceous ground cover that is believed to provide important microhabitat for striped newts. Similarly, altered fire regimes in ephemeral ponds also favor woody-stemmed vegetation and reduction of semiaquatic herbaceous vegetation believed to provide crucial breeding habitat and cover. Panhandle populations of the striped newt occur in sandhill and upland pine forest, but peninsular populations can occur in mesic or scrubby flatwoods, dry prairie, oak or sand pine scrub, and xeric hammock (Enge et al. 2014a, Enge et al. 2020). Suitable fire frequencies in these other habitats vary, and some striped newt populations persist in uplands and wetlands that rarely burn. Cogongrass (Imperata cylindrica), a perennial grass native to Southeast Asia, is one of the leading threats to the ecological integrity of native herbaceous flora, including that in the longleaf pine ecosystem in the southern United States (Jose et al. 2002). Cogongrass can outcompete wiregrass, altering the soil chemistry, nutrient cycling, and hydrology of an infested site (Jose et al. 2002). Cogongrass fires are typically 15‒20% hotter and more intense than natural fires in southern pinelands (MacDonald 2007), killing most aboveground vegetation except trees (Lippincott 2000) and possibly amphibians, including ones sheltering underground (Enge et al. 2014a). Urbanization associated with human population growth will result in habitat loss in the future. According to Carr and Zwick (2016), Florida is projected to add nearly 15 million residents from 2010 to 2070 and developed land will increase by approximately 10‒15%, depending upon the scenario used. However, all known extant striped newt populations except for 1 population in Duval County occur on conservation lands, which will not lose habitat from population growth. However, urban development around conservation lands will likely result in habitat degradation because of the inability to properly manage habitat by prescribed burning. It is often difficult, if not impossible, to provide correct ecological burns to some conservation lands because of the proximity of human settlements and structures to wildlands and the fear that either smoke or fire may negatively impact humans. This is likely to worsen in the coming decades as Florida’s population grows. Continued improper ecological use of fire is expected to negatively impact remaining striped newt populations. Groundcover alteration from altered fire regime, development, road construction, and vehicular mortality also interrupt potential gene flow among populations. Degradation and destruction of striped newt breeding ponds can result in population extinctions, particularly in areas with few suitable wetlands and fragmented uplands. Government regulatory agencies provide varying degrees of protection to wetlands, but they do not protect the small, ephemeral wetlands that striped newts often use for breeding sites (USFWS 2011). Exclusion and suppression of fire from wetlands often leads to degradation of breeding

Striped Newt Biological Status Review Report 9 ponds through shrub encroachment, peat buildup, reduction in herbaceous emergent vegetation, and increased evapotranspiration shortening hydroperiods (LaClaire 2001). For population recruitment, a wetland must contain water for at least 6 months (Johnson 2005). Herbaceous vegetation, bladderwort (Utricularia spp.), or comparable submerged plants are needed for egg deposition sites and for larval and adult cover from predators. Land managers often use fire lines to exclude prescribed fire from dry wetlands to prevent problems with smoke management or muck fires, particularly if the wetlands are associated with wildland urban interface (Bishop and Haas 2005). Erosion from adjacent unpaved roads can lead to siltation and sedimentation of ponds, and runoff from paved roads can pollute ponds with petrochemicals and other toxic substances to amphibians (LaClaire 2001). Pond degradation also results from garbage dumping and off-road vehicle (ORV) use (Means and Means 1998, LaClaire 2001). Use of ORVs in pond basins can cause direct mortality and affect habitat quality by altering pond contours, herbaceous vegetation, and hydrology (LaClaire 2001). Large ORV tires may break the organic hardpan beneath a pond, causing water to drain out and shortening the hydroperiod (LaClaire and Franz 1990). “Mudding” by ORVs has been observed in striped newt ponds in ANF and ONF, despite efforts by the U.S. Forest Service to prevent this practice. Means et al. (2008) documented extreme vegetative destruction at some ANF ponds. Feral hogs (Sus scrofa) may root up wetlands while they are dry, reducing vegetation, altering microtopography, and increasing eutrophication from feces (Enge et al. 2014a), and 1 striped newt pond on Triple N Ranch WMA was impacted by grazing cattle (Enge et al. 2020). The hydrology of many of Florida’s depression marsh wetlands may already have been significantly influenced by anthropogenic-caused impacts related to groundwater withdrawals. North Florida has undergone extreme shifts in groundwater potentiometric levels (i.e., “groundwater contours”) (Grubs and Crandall 2007). The hydrologic impact has been documented for the first time across regional hydrologic divides between the Suwannee River and St. Johns River water management districts, and there are numerous examples across the entire state where groundwater withdrawals are significantly shifting the historical directional flow of localized groundwater supplies (FWC 2011). Some ephemeral wetlands are independent of ground water or surface aquifer water, but other wetlands are being impacted by hydrological alterations related to groundwater withdrawal (Guzy et al. 2006). Groundwater withdrawal can shorten hydroperiods or even eliminate ephemeral wetlands. In addition to drought, groundwater consumption by an expanding human population in Tallahassee might exacerbate drying of ephemeral wetlands in the Munson Sandhills to the south (Ryan Means, pers. commun. 2020). Hydrology of all known ephemeral ponds within the known habitat of the striped newt in ANF is driven primarily by fluctuations of the underlying aquifer system (Kish and Means 2012, Means and Means 2019). Ephemeral wetlands in the Munson Sandhills are associated with a 5–10 m layer of clean sands containing thin lenses of clay that rest upon limestone units comprising the regional aquifer system (Kish and Means 2012, Means and Means 2019). Approximately 85% of ponds are within 2 m of the groundwater surface, and ponds fill when the groundwater surface is very near to the base elevation of the ponds and high seasonal rainfall provides additional water to the surface sand aquifer, producing a local “cap” resting on top of the main groundwater surface (Kish and Means 2012). Water availability in ephemeral wetlands in the Munson Sandhills is likely to continue to decrease over time as human population continues to increase. Therefore, striped newt breeding opportunities will likely decrease.

Striped Newt Biological Status Review Report 10 Climate change may be affecting the amount of winter precipitation in peninsular Florida (Seager 2009, IPPC 2013, Enge et al. 2014a, Dahl et al. 2019) and could potentially affect breeding pond hydrology (hydroperiod, timing, and water temperature) and upland habitat conditions (fire return and intensity) (FWC 2011, Zhu et al. 2017). Long-term droughts may already have caused some populations to disappear because of insufficient population recruitment. Experimental use of rubber EPDM pond liners to mitigate impacted hydroperiods within historical striped newt breeding ponds has proven extremely successful at 3 ponds in the Munson Sandhills (Means and Means 2019). However, this management technique likely would be logistically and financially impractical to use range wide across Florida. The introduction of fish may play a role in population declines. Eastern mosquitofish (Gambusia holbrooki) are sometimes introduced into isolated wetlands for mosquito control purposes. A far more serious threat, however, is the stocking of game fish (Lepomis spp. and Micropterus spp.) into ponds used by striped newts, or the introduction of predaceous fish into formerly fish-free wetlands during natural flooding events. Invertebrates may significantly impact population recruitment by feeding on eggs and larvae, and wading birds and snakes are potential predators while newts are in wetlands. Predators in uplands have not been identified, but mammals and snakes are the most likely. In xeric pinelands with a deep water table, red imported fire ants (Solenopsis invicta) are primarily confined to the margins of ephemeral wetlands, where they may kill striped newts (Means et al. 2015, Ryan Means, pers. commun. 2020). Droughts are naturally occurring events in the ecology of the striped newt, but prolonged droughts can threaten already small populations and exacerbate habitat degradation and fragmentation. Droughts lasting >4 years affect reproductive success, resulting in population declines of striped newts (Dodd 1992, Dodd and Johnson 2007). Changes in precipitation patterns and increased severity of both droughts and major storm events in the southeastern Coastal Plain have been implicated in population declines of ambystomatid species (Walls et al. 2013, Westervelt et al. 2013, Chandler et al. 2016). Long-term droughts exacerbated by groundwater withdrawal have been implicated in extinctions of striped newt populations in ponds in the Munson Sandhills of ANF and elsewhere (Dodd 1993, Means and Means 2008). In the range of the striped newt, droughts are predicted to become more severe and last longer because of climate change (Seager et al. 2009, USFWS 2011, IPCC 2013, Walls et al. 2013, Dahl et al. 2019). Projected warming trends will affect the amount, seasonal timing, and distribution of rainfall, frequency and intensity of storms, and sea level rise (IPCC 2013). Coastal striped newt populations may be affected by future sea level rise (Barrett et al. 2014). Stressors such as habitat loss and droughts are expected to elevate risks of diseases in newts, as has been observed in similar species (USFWS 2011). A chytridiomycete fungus (Batrachochytrium dendrobatidis), or Bd, has been implicated as a primary or suspected cause of disease epidemics and subsequent population declines of amphibians in many parts of the world, although chytrid fungus is not known to be responsible for any amphibian die-offs in the southeastern United States (Yap et al. 2017). Eastern newts are resistant to infection by various strains of Bd (Longo et al. 2019). Ranaviruses are likely a greater threat to amphibians, particularly tadpoles, than Bd in North America (Gray et al. 2009a, b). A fungus-like protist (Amphibiocystidium viridescens) has been reported as the cause of mortality and morbidity in eastern newt populations farther north (Raffel et al. 2008). Landsberg et al. (2013) tested 43 striped newts for Bd in Florida, and 9.3% tested positive. One of 13 striped newts tested positive

Striped Newt Biological Status Review Report 11 for ranavirus (Landsberg et al. 2013). All newts testing positive for pathogens (Bd and ranavirus) came from ONF and appeared asymptomatic. Dead newts have never been observed during hundreds of dipnet surveys of striped newt breeding ponds in Florida (Kevin Enge, pers. obs.). Die-offs of captive striped newts from Georgia may have resulted from infections by a ciliated protozoan or Mycobacterium; striped newts from ANF did not appear to be affected (Means et al. 2017). The fungal pathogen Batrachochytrium salamandrivorans (Bsal) infects salamanders in Europe and Asia and is predicted to invade the New World because of the presence of competent hosts, high volume import pathways, and lack of effective biosecurity measures (Longo et al. 2019). Experiments with eastern newts found that they were resistant to infection by Bsal, mortality from Bsal was much slower than previously observed, and some wild-caught newts may have innate immunity against Bsal (Longo et al. 2019). However, eastern newts exposed to both Bd, which is widespread in nature, and Bsal experienced high mortality rates from Bsal (Longo et al. 2019). It is possible that the intentional or accidental introduction of fish to newt breeding wetlands could elevate the risk of spreading pathogens to aquatic organisms (Rowley et al. 2013). The small geographic distribution of the striped newt plus its attractiveness and hardiness in captivity make it potentially desirable in the pet trade. No sales were reported during a study of the herpetofaunal pet trade in 1990‒1994 (Enge 2005), but its relative rarity and potential future listing have probably increased its desirability to collectors. Large numbers of larvae and paedomorphs can be collected in some ponds. For captive breeding efforts, 9 newts were collected in 2017 from an ANF pond, and at least a couple dozen newts were collected from Livingston Place. Population Assessment – Findings from the BRG are included in Biological Status Review Information Findings tables. The BRG found the striped newt warrants listing as Threatened at the species level. The genetically distinct panhandle population was assessed separately and also warrants listing as Threatened, which is unnecessary if the species is listed.

LISTING RECOMMENDATION Upon review of the BRG’s findings, FWC staff agrees that the Florida population of the striped newt meets criterion A, population size reduction, and criterion B, geographic range. Statewide, the striped newt is suspected to have had a 34% decline of occupied ponds in the past 15 years, a continued >30% decline of occupied ponds in the next 15 years, and a >30% decline of occupied ponds between 2010 and 2025. Additionally, Maxent modeling of potential habitat estimates that the area of occupancy for the species in Florida is 1,587 km2, and that the population is in continued decline. Striped newts are also vulnerable to extreme fluctuations in the number of adults. Further review of the Panhandle population found that the striped newt meets criterion A, population size reduction; criterion B, geographic range; criterion C, population size and trend; and criterion D, population very small or restricted. The BRG concluded that the Florida population of striped newt makes up over half of the global population, which only occurs elsewhere in Georgia. Because the striped newt meets multiple criteria for state listing, including a 34% decline of occupied ponds over 3 generations, limited range, and extreme population fluctuations, FWC staff recommends the statewide population of the striped newt be listed as a Threatened species.

Striped Newt Biological Status Review Report 12

SUMMARY OF THE INDEPENDENT REVIEW Independent scientific review was sought and received from 3 scientists. All 3 peer reviewers agreed that the Biological Status Review report was thorough, complete, and used appropriate data whenever possible. All 3 peer reviewers agreed that the striped newt met criteria for listing throughout their range in Florida. No major questions were raised by peer reviewers, and therefore no formal staff responses are provided in this report. Peer reviewers provided comments, suggestions, and citations that have been incorporated throughout the document, where appropriate. The complete scientific reviews are provided in Appendix 2. Staff of the FWC gratefully acknowledges the assistance of the independent reviewers.

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Zhu, J., G. Sun, W. Li, Y. Zhang, G. Miao, A. Noormets, S. G. McNulty, J. S. King, M Kumar, and X. Wang. 2017. Modeling the potential impacts of climate change on the water table level of selected forested wetlands in the southeastern United States. Hydrology and Earth System Sciences 21:6289‒6305.

Striped Newt Biological Status Review Report 20

Species/taxon: Striped newt (Florida population) Biological Status Review Date: 6/26/20 Information Findings Assessors: Enge, Greene, Johnson, Means, Stevenson

Generation length: 5 years

Sub-Criterion Criterion/Listing Measure Data/Information Data Type* References Met? *Data Types - observed (O), estimated (E), inferred (I), suspected (S), or projected (P). Sub-Criterion met - yes (Y) or no (N).

(A) Population Size Reduction, ANY of (a)1. An observed, estimated, inferred or 34% decline of occupied ponds; S N See Table 1 suspected population size reduction of at least causes of reduction are not 50% over the last 10 years or 3 generations, clearly reversible, understood, whichever is longer, where the causes of the and ceased reduction are clearly reversible and understood and ceased1 (a)2. An observed, estimated, inferred or 34% decline of occupied ponds S Y See Table 1 suspected population size reduction of at least in past 15 years decline in 30% over the last 10 years or 3 generations, occupied ponds in past 15 years whichever is longer, where the reduction or its based on an index of abundance, causes may not have ceased or may not be decline in AOO and habitat understood or may not be reversible1 quality, and effects of fire ants and pathogens; causes of the reduction are not clearly reversible, understood, and ceased (a)3. A population size reduction of at least 30% Expect >30% decline of occupied S Y projected or suspected to be met within the next ponds in next 15 years based on 10 years or 3 generations, whichever is longer past trends and a decline in AOO (up to a maximum of 100 years) 1 and habitat quality and effects of fire ants and pathogens

Striped Newt Biological Status Review Report 21 (a)4. An observed, estimated, inferred, projected Expect >30% decline of occupied S Y See Table 1 or suspected population size reduction of at least ponds in 2010-2025 based on an 30% over any 10 year or 3 generation period, index of abundance, decline in whichever is longer (up to a maximum of 100 AOO and habitat quality, and years in the future), where the time period must effects of fire ants and pathogens; include both the past and the future, and where causes of the reduction are not the reduction or its causes may not have ceased clearly reversible, understood, or may not be understood or may not be and ceased reversible.1 1 based on (and specifying) any of the following: (a) direct observation; (b) an index of abundance appropriate to the taxon; (c) a decline in area of occupancy, extent of occurrence and/or quality of habitat; (d) actual or potential levels of exploitation; (e) the effects of introduced taxa, hybridization, pathogens, pollutants, competitors or parasites. (B) Geographic Range, EITHER (b)1. Extent of occurrence < 20,000 km2 (7,722 31,484 km2 E N Geographical range map (Enge mi2 ) OR 2019) (b)2. Area of occupancy < 2,000 km2 (772 mi2 263.5 km2 around ponds; E Y Mapping of possibly extant ) 1,587 km2 of potential habitat subpopulations using a 1-km buffer around ponds (Brittany Bankovich, pers. commun.); Maxent modeling of potential habitat (Enge et al. 2014a) AND at least 2 of the following: a. Severely fragmented or exist in ≤ 10 Occurs in >10 locations and not S N Farmer et al. (2017) locations severely fragmented b. Continuing decline, observed, inferred or Based on previous declines I Y Farmer et al. (2017) projected in any of the following: (i) extent of (some of which occurred for occurrence; (ii) area of occupancy; (iii) area, unknown reasons), future extent, and/or quality of habitat; (iv) number of declines in i-v are inferred locations or subpopulations; (v) number of mature individuals c. Extreme fluctuations in any of the Extreme fluctuations in number I Y Pechmann et al. (1991), Dodd following: (i) extent of occurrence; (ii) area of of adults can occur in amphibian (1993) occupancy; (iii) number of locations or species breeding in ephemeral subpopulations; (iv) number of mature wetlands individuals (C) Population Size and Trend Population size estimate to number fewer than Population size >10,000 adults; E N Johnson (2002b); Table 1 10,000 mature individuals AND EITHER 53 ponds with records in the past 5 years, and 76 ponds with records in the past 10 years; 1 pond had >2,000 adults

Striped Newt Biological Status Review Report 22 (c)1. An estimated continuing decline of at least Expect >10% decline of occupied I Y Farmer et al. (2017) 10% in 10 years or 3 generations, whichever is ponds in next 15 years based on longer (up to a maximum of 100 years in the past trends and continued threats future) OR (c)2. A continuing decline, observed, projected, Expect a continued decline of I Y Farmer et al. (2017) or inferred in numbers of mature individuals occupied ponds in next 15 years AND at least one of the following: based on past trends and continued threats a. Population structure in the form of EITHER (i) No subpopulation estimated to contain One subpopulation contains 11 S N Johnson (2002); Kevin. Enge more than 1000 mature individuals; OR ponds, and 1 pond had >2,000 (unpubl. data) adults (ii) All mature individuals are in one 46 subpopulations are present S N Kevin Enge (unpubl. data) subpopulation b. Extreme fluctuations in number of mature Extreme fluctuations in number I Y Pechmann et al. (1991), Dodd individuals of adults can occur in amphibian (1993) species breeding in ephemeral wetlands (D) Population Very Small or Restricted, EITHER (d)1. Population estimated to number fewer than Population size >10,000 adults E N Johnson (2002b) 1,000 mature individuals; OR (d)2. Population with a very restricted area of 263.5 km2 around ponds; N Mapping of possibly extant occupancy (typically less than 20 km2 [8 mi2]) or 1,587 km2 of potential habitat; >5 E subpopulations using a 1-km number of locations (typically 5 or fewer) such locations buffer around ponds (Brittany that it is prone to the effects of human activities Bankovich, pers. commun.); or stochastic events within a short time period in Maxent modeling of potential an uncertain future habitat (Enge et al. 2014a) (E) Quantitative Analyses e1. Showing the probability of extinction in the wild is at least 10% within 100 years No PVA N

Initial Finding (Meets at least one of the criteria/sub- Reason (which criteria/sub-criteria are criteria OR Does not meet any of the criteria/sub-criteria) met) Meets at least one of the criteria A2(b,c,e), A3(c,e), A4(b,c,e), B2b(i,ii,iii,iv,v)c(iv)

Is species/taxon endemic to Florida? (Y/N) N If Yes, your initial finding is your final finding. Copy the initial finding and reason to the final finding space below. If No, complete the regional assessment sheet and copy the final finding from that sheet to the space below. Striped Newt Biological Status Review Report 23

Final Finding (Meets at least one of the criteria/sub- Reason (which criteria/sub-criteria are criteria OR Does not meet any of the criteria/sub-criteria) met) Meets at least one of the criteria A2(b,c,e), A3(c,e), A4(b,c,e), B2b(i,ii,iii,iv,v)c(iv)

Striped Newt Biological Status Review Report 24 1 Species/taxon: Striped newt (Florida population) 2 Biological Status Review Information Date: 6/26/20 3 Regional Assessment Assessors: Enge, Greene, Johnson, Means, 4 Stevenson 5 6 7 8 Initial finding 9 2a. Is the species/taxon a non-breeding visitor? (Y/N/DK). If 2a is YES, go to line 18. If 2a is NO or DO NOT No 10 KNOW, go to line 11. 2b. Does the Florida population experience any significant immigration of propagules capable of No 11 reproducing in Florida? (Y/N/DK). If 2b is YES, go to line 12. If 2b is NO or DO NOT KNOW, go to line 17. 2c. Is the immigration expected to decrease? (Y/N/DK). If 2c is YES or DO NOT KNOW, go to line

12 13. If 2c is NO go to line 16. 2d. Is the regional population a sink? (Y/N/DK). If 2d is YES, go to line 14. If 2d is NO or DO

13 NOT KNOW, go to line 15. 14 If 2d is YES - Upgrade from initial finding (more imperiled) 15 If 2d is NO or DO NOT KNOW - No change from initial finding 16 If 2c is NO or DO NOT KNOW- Downgrade from initial finding (less imperiled) 17 If 2b is NO or DO NOT KNOW - No change from initial finding

2e. Are the conditions outside Florida deteriorating? (Y/N/DK). If 2e is YES or DO NOT 18 KNOW, go to line 24. If 2e is NO go to line 19. 2f. Are the conditions within Florida deteriorating? (Y/N/DK). If 2f is YES or DO NOT 19 KNOW, go to line 23. If 2f is NO, go to line 20. 2g. Can the breeding population rescue the Florida population should it decline? 20 (Y/N/DK). If 2g is YES, go to line 21. If 2g is NO or DO NOT KNOW, go to line 22. 21 If 2g is YES - Downgrade from initial finding (less imperiled) 22 If 2g is NO or DO NOT KNOW - No change from initial finding 23 If 2f is YES or DO NOT KNOW - No change from initial finding 24 If 2e is YES or DO NOT KNOW - No change from initial finding 25 26 Final finding Change

Striped Newt Biological Status Review Report 25 Species/taxon: Striped newt (panhandle population) Biological Status Review Date: 6/26/20 Information Findings Assessors: Enge, Greene, Johnson, Means, Stevenson

Generation length: 5 years

Sub- Criterion/Listing Measure Data/Information Data Type* Criterion References Met? *Data Types - observed (O), estimated (E), inferred (I), suspected (S), or projected (P). Sub-Criterion met - yes (Y) or no (N).

(A) Population Size Reduction, ANY of (a)1. An observed, estimated, inferred or 70% decline, but causes are not clearly S N See Table 1 suspected population size reduction of at least reversible, understood, and ceased 50% over the last 10 years or 3 generations, whichever is longer, where the causes of the reduction are clearly reversible and understood and ceased1 (a)2. An observed, estimated, inferred or 70% decline in occupied ponds in past 15 S Y See Table 1 suspected population size reduction of at least years based on an index of abundance, 30% over the last 10 years or 3 generations, decline in AOO and habitat quality, and whichever is longer, where the reduction or its effects of fire ants and pathogens; causes causes may not have ceased or may not be of the reduction are not fully understood understood or may not be reversible1 and probably have not ceased (a)3. A population size reduction of at least 30% Unsure if populations will go extinct in at P ? projected or suspected to be met within the next least 2 of the 6 known occupied ponds in 10 years or 3 generations, whichever is longer (up the next 15 years to a maximum of 100 years) 1

(a)4. An observed, estimated, inferred, projected or suspected population size reduction of at least 30% over any 10 year or 3 generation period, whichever is longer (up to a maximum of 100 years in the future), where the time period must include both the past and the future, and where the reduction or its causes may not have ceased or may not be understood or may not be reversible.1

Striped Newt Biological Status Review Report 26 1 based on (and specifying) any of the following: (a) direct observation; (b) an index of abundance appropriate to the taxon; (c) a decline in area of occupancy, extent of occurrence and/or quality of habitat; (d) actual or potential levels of exploitation; (e) the effects of introduced taxa, hybridization, pathogens, pollutants, competitors or parasites. (B) Geographic Range, EITHER (b)1. Extent of occurrence < 20,000 km2 (7,722 3,043 km2 E Y Geographical range map mi2 ) OR (Enge 2019) (b)2. Area of occupancy < 2,000 km2 (772 mi2 ) 27.6 km2 around ponds; 1,587 km2 of E Y 1-km buffer around all potential habitat throughout Florida, so it is known breeding ponds; considerably less in just the panhandle Maxent model of potential habitat (Enge et al. 2014a) AND at least 2 of the following: a. Severely fragmented or exist in ≤ 10 Severely fragmented and 2 locations O Y ANF and Livingston Place locations are the 2 locations, which are 60 km apart b. Continuing decline, observed, inferred or Based on previous declines in ANF and I Y projected in any of the following: (i) extent of increased groundwater withdrawal for occurrence; (ii) area of occupancy; (iii) area, Tallahassee, future declines in i-v are extent, and/or quality of habitat; (iv) number of inferred locations or subpopulations; (v) number of mature individuals c. Extreme fluctuations in any of the following: Extreme fluctuations in number of adults S Y Pechmann et al. (1991), (i) extent of occurrence; (ii) area of occupancy; can occur in amphibian species breeding in Dodd (1993) (iii) number of locations or subpopulations; (iv) ephemeral wetlands number of mature individuals (C) Population Size and Trend Population size estimate to number fewer than A mean of 1,667 adults would need to be S Y 10,000 mature individuals AND EITHER present in each of the 6 ponds for there to be 10,000 adults, which is unlikely (c)1. An estimated continuing decline of at least Unsure if a population will go extinct in at S N 10% in 10 years or 3 generations, whichever is least 1 of the 6 known occupied ponds in longer (up to a maximum of 100 years in the the next 15 years future) OR (c)2. A continuing decline, observed, projected, Based on past declines and continued I Y or inferred in numbers of mature individuals threats in ANF, a continuing decline in the AND at least one of the following: number of adults can be inferred a. Population structure in the form of EITHER (i) No subpopulation estimated to contain more than 1000 mature individuals; OR Livingston Place may have >1,000 adults S N

Striped Newt Biological Status Review Report 27 (ii) All mature individuals are in one 2 subpopulations S N subpopulation b. Extreme fluctuations in number of mature Extreme fluctuations in number of adults I Y Pechmann et al. (1991), individuals can occur in amphibian species breeding in Dodd (1993) ephemeral wetlands (D) Population Very Small or Restricted, EITHER (d)1. Population estimated to number fewer than >1,000 adults S N 1,000 mature individuals; OR (d)2. Population with a very restricted area of 2 locations O Y ANF already experienced a occupancy (typically less than 20 km2 [8 mi2]) or severe population decline, number of locations (typically 5 or fewer) such and Livingston Place may that it is prone to the effects of human activities have deleterious management or stochastic events within a short time period in practices, St. Marks NWR an uncertain future population extirpated (E) Quantitative Analyses e1. Showing the probability of extinction in the wild is at least 10% within 100 years No PVA N

Initial Finding (Meets at least one of the criteria/sub- Reason (which criteria/sub-criteria are met) criteria OR Does not meet any of the criteria/sub-criteria) Meets at least one of the criteria A2(b,c,e), B1ab(i,ii,iii,iv,v)c(iv), B2ab(i,ii,iii,iv,v)c(iv), C2b, D2

Is species/taxon endemic to Florida? (Y/N) N If Yes, your initial finding is your final finding. Copy the initial finding and reason to the final finding space below. If No, complete the regional assessment sheet and copy the final finding from that sheet to the space below.

Final Finding (Meets at least one of the criteria/sub- Reason (which criteria/sub-criteria are met) criteria OR Does not meet any of the criteria/sub-criteria) Meets at least one of the criteria A2(b,c,e), B1ab(i,ii,iii,iv,v)c(iv), B2ab(i,ii,iii,iv,v)c(iv), C2b, D2

Striped Newt Biological Status Review Report 28 1 Species/taxon: Striped newt (panhandle population) 2 Biological Status Review Information Date: 6/26/20 3 Regional Assessment Assessors: Enge, Greene, Johnson, Means, 4 Stevenson 5 6 7 8 Initial finding 9 2a. Is the species/taxon a non-breeding visitor? (Y/N/DK). If 2a is YES, go to line 18. If 2a is NO or DO No 10 NOT KNOW, go to line 11. 2b. Does the Florida population experience any significant immigration of propagules capable of reproducing in Florida? (Y/N/DK). If 2b is YES, go to line 12. If 2b is NO or DO NOT KNOW, go to line No 11 17. 2c. Is the immigration expected to decrease? (Y/N/DK). If 2c is YES or DO NOT KNOW, go to

12 line 13. If 2c is NO go to line 16. 2d. Is the regional population a sink? (Y/N/DK). If 2d is YES, go to line 14. If 2d is NO or

13 DO NOT KNOW, go to line 15. 14 If 2d is YES - Upgrade from initial finding (more imperiled) 15 If 2d is NO or DO NOT KNOW - No change from initial finding 16 If 2c is NO or DO NOT KNOW- Downgrade from initial finding (less imperiled) 17 If 2b is NO or DO NOT KNOW - No change from initial finding

2e. Are the conditions outside Florida deteriorating? (Y/N/DK). If 2e is YES or DO 18 NOT KNOW, go to line 24. If 2e is NO go to line 19.

2f. Are the conditions within Florida deteriorating? (Y/N/DK). If 2f is YES or 19 DO NOT KNOW, go to line 23. If 2f is NO, go to line 20.

2g. Can the breeding population rescue the Florida population should it 20 decline? (Y/N/DK). If 2g is YES, go to line 21. If 2g is NO or DO NOT KNOW, go to line 22. 21 If 2g is YES - Downgrade from initial finding (less imperiled) 22 If 2g is NO or DO NOT KNOW - No change from initial finding 23 If 2f is YES or DO NOT KNOW - No change from initial finding 24 If 2e is YES or DO NOT KNOW - No change from initial finding

Striped Newt Biological Status Review Report 29 25 26 Final finding Change

Striped Newt Biological Status Review Report 30 Additional notes – Generation length is defined as the average age of parents of the current cohort, which is greater than the age at first breeding and less than the age of the oldest breeding individual. We infer a mean generation length of 5 years. Paedomorphs may become sexually mature in 1 year (Johnson 2002b), but newts that become efts require longer to reach sexual maturity. A larva requires ca. 6 months before metamorphosing into an eft (Johnson 2002b), and our best guess is that the eft stage in Florida lasts 1‒3 years. This means that a terrestrial adult returning to a pond to breed for the first time is 1.5‒3.5 years old if it began as an eft. According to Johnson (2002b), a paedomorph breeds only once before transforming into a terrestrial adult. If this is the case, paedomorphs should be <2 years old. Johnson (2002b) estimated that 25% of the larvae in his study pond became paedomorphs, whereas 75.5% of 1,161 adult newts captured during FWC dipnet surveys were paedomorphs (Kevin Enge, unpubl. data). If paedomorphs breed only once and most adults are paedomorphs, then the average age of parents is relatively young. Notophthalmus typically live 12‒15 years (Forester and Lykens 1991), and a captive striped newt lived 17 years (Wallace et al. 2009). Although striped newts have the potential to live over a decade, we do not know their survival rate in the wild. Striped newts have been documented recolonizing a pond after it was dry for 10 years (Dodd and Johnson 2007), but it is unknown if this was the natal pond of the colonizers. An adult, nongilled newt spends ca. 75% of its time on land in years when its breeding pond fills. Newts in uplands are susceptible to mortality from desiccation, predation, and fire. If paedomorphs inhabiting ponds with permanent water and unsuitable upland habitat can breed multiple times, then the mean generation length would likely be 1 or 2 years longer. These paedomorphic populations would likely have more young than old individuals because of cumulative mortality from many predators. Sub-criterion A1 ‒ Although the panhandle population is suspected of experiencing at least a 50% population size reduction since 2005, all BRG members agreed that it was unnecessary to address this sub-criterion because all the causes of striped newt population declines are not known, and these declines apparently are continuing to occur. These declines are not clearly reversible, although repatriation is being attempted in the Munson Sandhills of Apalachicola National Forest (ANF), along with installation of pond liners to increase hydroperiods (Means et al. 2017). Sub-criterion A2 ‒ A reduction in striped newt population size presumably has occurred as the human population in Florida has increased and converted suitable habitat to urban, agricultural, and other land uses. However, striped newt populations are now almost exclusively restricted to conservation lands, so habitat loss in the past 15 years (3 generations) has probably not been responsible for substantial striped newt population declines. A decrease in the number of occupied ponds (Table 1) was used as an index of population size reduction. Threats potentially responsible for population reductions are (c) declines in habitat quality and possibly area of occupancy and (e) the effects of introduced taxa and pathogens. Habitat degradation associated with unfavorable fire regimes or other habitat management practices on conservation lands have probably contributed to population declines. Prolonged droughts exacerbated by groundwater withdrawal may also have caused population declines. Documented or suspected population declines have occurred on conservation lands, even ones those with habitat that appears to be in decent condition because of prescribed burning and other management practices favorable for striped newts. For example, striped newts have disappeared from the Panacea Unit

Striped Newt Biological Status Review Report 31 of St. Marks National Wildlife Refuge (Dodd et al. 2007) and most of ANF (Means et al. 2008), despite suitable upland habitat and wetlands remaining. Apparent population declines or extirpations may be related to several years of unsuitable weather for breeding or population recruitment (or to unknown factors), but populations may recolonize ponds (Dodd and Johnson 2007). Means et al. (2008) speculated that some unknown disease pathogen contributed to the serious population decline in ANF. Introduced red imported fire ants kill emerging striped newts along pond margins in ANF (Ryan Means, pers. commun. 2020). Actual estimates of striped newt population size do not exist, and the number of populations or percentage of habitat impacted by habitat degradation or prolonged droughts is difficult to quantify. However, the striped newt population decline in ANF is well documented (Means and Means 2008). Dipnet surveys of 17 known ANF breeding ponds detected striped newts in 12 ponds in 1986‒1995, 12 ponds in 1996‒2000 (newts were found in 7 ponds during both time periods), 0 ponds in 2001‒2005 (1 survey year), 2 ponds in 2006‒2010 (2 survey years), 0 ponds in 2011‒2015, and 1 pond in 2016‒2020 (Rebecca Means, Coastal Plains Institute, pers. commun. 2020). At a pond completely encircled by a drift fence in ANF, 2,023 striped newts were trapped in 1995‒2000 but only 10 newts in 2001‒2005 (Rebecca Means, pers. commun. 2020). An estimate of the percent population decline during the past 15 years can be derived based on the last year newts were detected in known breeding ponds (Table 1). Several assumptions must be made for the number of ponds to be an accurate index of population size: 1) population size must be equivalent among ponds, 2) ponds must be surveyed frequently and intensively enough to provide meaningful results, and 3) newts must be present in ponds during surveys of extant populations. These assumptions cannot be met with existing information, so there is uncertainty associated with the calculated percent population decline. Newt detection rates from dipnetting surveys are low, particularly in large ponds with low newt densities, and are affected by many variables that differ among ponds. Means et al. (2017) estimated detectability of 10‒20% in ponds with low newt densities. As of 2005 (15 years ago), striped newts were known to occupy 82 ponds in Florida, 18 of which were in the panhandle in ANF. This number includes ponds with suitable habitat where striped newts were last documented in the 1990s. A prolonged drought occurred in much of Florida in the late 1990s, and many ponds dried and could not be surveyed. Striped newts were found in only 2 ANF ponds during a 2007 dipnet survey, but populations could still have been extant in the other ponds but at such low densities that they were not detected. Even if ponds were dry and could not be surveyed, long- lived adults could have remained in the surrounding uplands. Striped newts were thought to be extinct in ANF (Means et al. 2008) but were found in 1 ANF pond in 2016, where they went undetected for 9 years (Farmer et al. 2017). Because of an expansion of survey effort by FWC staff, 70 additional breeding ponds have been discovered since 2005 (Kevin Enge, unpubl. data), including 5 ponds in Jefferson County in the panhandle (Hill and Sash 2015). If we add these new ponds to the totals, a 34% decline in occupied ponds (N = 152) has occurred since 2005. Including newly discovered ponds, the panhandle population has experienced a 70% decrease in occupied ponds (N = 23) since 2005.

This suspected population size reduction of 34% for the species (i.e., Florida population) slightly exceeds the 30% threshold. Three BRG members suspected a population size reduction of at least 30% in the Florida population since 2005, but 2 BRG members did not feel comfortable in concluding this because of the uncertainty of the data. Populations in undiscovered ponds probably also went extinct since 2005, which would increase the population size reduction, but this could have been counteracted by failure to detect newts in ponds where populations were thought to have gone extinct, such as occurred in 1 ANF pond. Some ponds on private property, where populations are assumed to be extinct because of poor habitat conditions, have not been surveyed. Recent finds of new breeding ponds in poor upland habitat (i.e., xeric hammock) in Ocala National Forest and Seminole State Forest have shown that populations can persist in sandhill habitat that has not burned in decades. All BRG members suspected a population size reduction of at least 50% since 2005 for the panhandle population based upon a 70% decrease in occupied ponds.

Table 1. The number of known ponds where striped newts were last observed by time period (NS = not surveyed). Each number represents a unique pond, and the total number of ponds at each site is the same as appears in Table 2.

1986- 1996- 2001- 2006- 2011- 2016- Site 1995 2000 2005 2010 2015 2020 Apalachicola National Forest 5 10 1 1 1 Big Bend Wildlife Management Area 1 1 Camp Blanding Military Reservation 6 2 8 Faver-Dykes State Park 2 1 NS Goethe State Forest 5 Guana River Wildlife Management Area 1 2 GTMNERR 1 NS NS NS Half Moon Wildlife Management Area 2 Jennings State Forest 5 9 4 Livingston Place 5 Merritt Island National Wildlife Refuge 1 Ocala National Forest 3 3 11 3 23 Ordway-Swisher Biological Station 4 5 NS 6 Pumpkin Hill Creek Preserve State Park 1 NS Rainbow Springs State Park 1 NS NS NS Rock Springs Run State Reserve 1 NS 3 Seminole State Forest 1 NS 1 Triple N Ranch Wildlife Management Area 9 DeeDot Ranch (St. Johns Co.) 1 NS NS NS NS Plum Creek Timberland (Putnam Co.) 3 NS NS NS NS

Sub-criterion A3. – Three generations from 2020 would be 2035. Rayer and Wang (2020) projected an increase of 3,942,000 people from 2020 to 2035 based on the most likely medium population growth series. According to Carr and Zwick (2016), Florida’s population is expected to increase by more than 6 million people in the next 20 years (by 2040), requiring about 2.5 million more acres. The exact relationship between human population increase and habitat loss is unknown. Approximately 40% of the land that will be converted to new urban use is in native habitat or pine plantation (Carr and Zwick 2016). Almost all known striped newt populations occur on protected lands, so the impact of population growth and associated development may be minimal. However, undiscovered striped newt populations could occur on private land, particularly in Clay and Putnam counties. The Livingston Place population was recently discovered (Hill and Sash 2015). In the next 20 years, population growth and transformation of formerly undeveloped lands to urban use is predicted to be explosive in central Florida, including Marion, Lake, and Seminole counties (Carr and Zwick 2016). All known extant striped newt populations in these counties are on public lands, primarily Ocala National Forest (ONF). Centers of considerable growth include Keystone Heights in Putnam County and almost all of St. Johns County (Carr and Zwick 2016); these areas have striped newt populations, possibly even on private land. Flagler and Osceola counties are the 2 counties with striped newt populations that will undergo the greatest land-use transformation (Carr and Zwick 2016). Very few known striped newt breeding ponds occur outside of conservation lands, however. Populations on unprotected lands will likely decrease or go extinct in the future because of development or habitat degradation from altered fire regimes. On public lands, there is an increasing trend of backlogged, fire-suppressed communities (FWC 2011), and the ability of striped newt populations to continue to persist on these protected properties is unknown. The ability to properly manage striped newt habitat by prescribed burning will be further compromised by habitat fragmentation by roads and by increased development around newt populations and the risk of property destruction and of smoke causing traffic accidents and health issues. Fortunately, efforts are being made to restore degraded sandhill habitat (FWC 2011), but the ability of striped newts to survive mechanical treatment is unknown. Inexplicable population declines or extinctions of striped newts on conservation lands are a cause for concern and indicate that such events could occur elsewhere in the future. If the fungal pathogen Bsal is introduced into North America, it could have devastating impacts on populations of newts, particularly those already exposed to Bd (Longo et al. 2019). Three BRG members suspected a population size reduction of at least 30% in the Florida population in the next 15 years, but 2 BRG members did not feel comfortable reaching this conclusion. Confidence might be higher for a future than for a past population decline of at least 30% for the Florida population. A conclusion regarding a future population size reduction in the panhandle population could not be reached: 2 BRG members voted yes, 2 voted no, and 1 abstained. To reach the 30% threshold, at least 2 of the 6 ponds would need to have populations go extinct. Sub-criterion A4. – Despite the uncertainty of the data, all 5 BRG members suspected a population decline of at least 30% for the Florida population during period from 2010 through 2025. Population declines have been documented in the past 10 years (Table 1), and there is no reason to expect the trend to change in the next 5 years, particularly because of problems with burning, groundwater withdrawal, and introduced taxa or pathogens. The panhandle population was not discussed. Sub-criterion B1. – The extent of occurrence (EOO) was estimated by using ArcGIS to calculate the area of the geographical range delineated in the book Amphibians and Reptiles of Florida (Enge 2019). The EOO in Florida totals 31,484 km2, whereas that in the panhandle totals 3,043 km2 (Brittany Bankovich, FWC, pers. commun. 2020). All 5 BRG members agreed that the panhandle population met the sub-criterion threshold of <20,000 km2. See Sub-criterion B2 for a discussion of the other conditions that must be met. Sub-criterion B2. – The area of occupancy (AOO) was estimated using 2 methods. The first method drew 1-km buffers around each breeding pond (3.14 km2) with a newt record since 1998 and suitable wetland and circumjacent upland habitats (Figs. 2 and 3). Based on extrapolation of migration distances revealed by upland drift fences, Johnson (2003) estimated a protected area of “core habitat” extending ca. 1 km from the study pond at Ordway-Swisher Biological Station would likely be needed to encompass almost all of the striped newts that breed there. A circle with a 1-km radius has an area of 3.14 km2, but population buffers overlap because of the proximity of ponds. Buffer zones intersected if ponds were within 2 km of each other, and the area of merged buffers was calculated instead of individual buffers. Buffers terminated along major highways, large streams, and lake borders. The calculated AOO should be considered maximal because newt dispersal distances are likely shorter in suboptimal circumjacent habitat, such as scrub, hammocks, and mesic flatwoods. Upland dispersal may not occur from ponds in scrub where adult populations consist solely of paedomorphs. The AOO in Florida is 263.5 km2 and in the panhandle is 27.6 km2 (Brittany Bankovich, pers. commun. 2020). Both the Florida and panhandle populations easily meet the sub-criterion threshold of <2,000 km2. The second method used Maxent (maximum entropy) modeling to determine that amount of potential habitat for the striped newt and was less restrictive than the first method. Using this method, the AOO in Florida is 1,587 km2 (Enge et al. 2014a), which means the panhandle population also meets this sub-criterion. In order to meet Criterion B, 2 of the 3 following conditions must be met: a) severely fragmented or exists in ≤10 locations; b) continuing decline in (i) extent of occurrence, (ii) area of occupancy, (ii) habitat, (iv) number of locations or (v) subpopulations, or number of mature individuals; or c) extreme fluctuations in (i) extent of occurrence, (ii) area of occupancy, (iii) number of locations or subpopulations, or (iv) number of mature individuals. “Subpopulations” are defined as geographically or otherwise distinct groups in the population between which there is little demographic or genetic exchange (typically 1 successful migrant individual or gamete per year or less) (IUCN 2019). The term subpopulation is equivalent to metapopulation (IUCN 2019). The definition of “severely fragmented” is most individuals are found in small and relatively isolated subpopulations (IUCN 2019). A “location” is defined as a geographically or ecologically distinct area in which a single threatening event can rapidly affect all individuals present (IUCN 2019). For the striped newt, such a threatening event could be a disease outbreak. Because the species is long-lived, a prolonged drought that eliminated population recruitment for several years would not rapidly affect all individuals. A flood that introduced predatory fish species into all ponds would immediately affect individuals present in ponds but not individuals residing in uplands. The panhandle population probably consists of 2 locations, ANF and Livingston Place. Throughout the range of the striped newt in Florida, the 150 known breeding ponds are fragmented into 46 subpopulations (Table 2, Fig. 2). Subpopulations appear to be dispersed widely across the landscape, although clumping occurs in population strongholds (Fig. 2). Striped newts have limited dispersal capabilities, and most subpopulations are isolated by distance. One BRG member voted that these subpopulations are severely fragmented, except possibly for ONF, because 98% of the original longleaf pine ecosystem has been lost and genetic exchange is no longer possible between most subpopulations. The other BRG members did not feel that the remaining subpopulations met the IUCN definition of severely fragmented, because 46% of the ponds are located in 8 subpopulations containing 7‒11 ponds each at 5 sites. Several of the ponds in these 8 subpopulations are known to have robust striped newt populations, so they do not feel that most individuals are found in small and relatively isolated subpopulations (the definition of severely fragmented). Small, isolated subpopulations likely have relatively few individuals compared to subpopulations containing many ponds. Ocala National Forest contains 29% of all known ponds and 23% of all presumably extant, viable subpopulations. The discovery of additional breeding ponds between subpopulations in ONF could reduce the number of identified subpopulations (N = 11) and increase their size, because most subpopulations are isolated by distance and not by barriers to movement. In contrast, 4 of the 5 subpopulations in Jennings State Forest are separated by streams with hardwood corridors 0.15‒1.0 km wide. All 5 BRG members agreed that the panhandle population met the definition of severely fragmented, because it is now restricted to only 2 subpopulations (Table 2). The Leon County (ANF) subpopulation apparently now consists of only 1 pond with a self-sustaining population (Means et al. 2017). One of the ANF repatriation ponds possibly could be considered a third subpopulation, but until a large number of F1 and later generation terrestrial adults immigrate into the pond and breed successfully, it should not be considered to have a long-term, self- sustaining population (Ryan Means, pers. commun. 2020). The 18 historical ANF ponds occurred in 4 subpopulations, 2 of which consisted of only a single pond (Fig. 3). The westernmost ANF pond is situated 2.25 km from the nearest known breeding pond. Another ANF pond is isolated from other breeding ponds by 2 major highways to the west and Munson Slough to the east (Fig. 23.) The Jefferson County subpopulation on Livingston Place consists of 5 known ponds (Table 2). The Leon and Jefferson County subpopulations are separated by >60 km of mostly unsuitable habitat and several major highways (Fig. 3). This species has limited mobility, so these subpopulations are isolated. In addition, the panhandle population contains a maximum of 3 known locations if the repatriated subpopulation is included. All 5 BRG members considered both the Florida and panhandle populations to meet the sub-criterion of an inferred continuing decline in (i) extent of occurrence, (ii) area of occupancy, (iii) habitat, (iv) number of locations or subpopulations, or (v) number of mature individuals. The striped newt likely meets all 5 of these conditions unless remedial measures are taken. Looking at the last year striped newts were observed in various ponds (Table 1), there has been a trend of population extinction, even on conservation lands with appropriate habitat management (Farmer et al. 2017). Some of these ponds may still have extant populations because detection rates can be low, and some ponds were not surveyed sufficiently. However, other ponds, such as in ANF, have been surveyed intensively enough to conclude that population extinctions have occurred.

Fig. 2. Subpopulations of the striped newt in Florida, with each breeding pond surrounded by a 1-km radius of core upland habitat.

Because striped newts are long-lived, populations can survive unfavorable stochastic environmental events, such as periods of less-than-average rainfall, that affect reproduction (Dodd 1993, Dodd et al. 2005). However, terrestrial amphibian species that breed in ephemeral wetlands often experience extreme fluctuations in population size greater than 1 order of magnitude based on the number of adults using breeding ponds (Pechmann et al. 1991). At a pond encircled by a drift fence in northern peninsular Florida, adult striped newt numbers fluctuated annually during a period of drought from 29 to 920 (Dodd 1993), which is greater than 1 order of magnitude. The number of adult females of 4 terrestrial amphibian species using an ephemeral wetland in South Carolina fluctuated annually by over 3 orders of magnitude, and the number of adults fluctuated by at least 1 order of magnitude (Pechmann et al. 1991). Fluctuations in breeding populations were not correlated among the 4 species, and no declines in these populations could not readily be explained as natural fluctuations in part related to drought (Pechmann et al. 1991). Two of these amphibian species, the mole salamander and ornate chorus frog (Pseudacris ornata), still breed in ANF ponds where striped newt populations have gone extinct. All 5 BRG members agreed that the Florida and panhandle populations likely experience extreme fluctuations in (iv) number of mature individuals.

Fig. 3. Subpopulations of the striped newt in the Florida panhandle, showing 1-km buffers around breeding ponds and barriers to movement that separate subpopulations. The 5 ponds in Jefferson County are in the same subpopulation, whereas the 18 ponds in Leon County are in 4 subpopulations.

All 5 BRG members agreed that the panhandle population met Criterion B2 because it has a small extent of occurrence and area of occupancy, plus it is severely fragmented and has an inferred continuing population decline unless remedial measures are taken. Remedial measures are being undertaken in ANF, where 2,277 captive-reared striped newt larvae and aquatic adults have been released into 6 ponds from 2013 through 2020. Repatriation efforts have had some early signs of success, primarily in 1 pond. Success benchmarks observed include the production of F1 and F2 generations at the 1 pond from 2016 through 2018. However, fewer than 50 terrestrial individuals recruited into the uplands from the most successful pond, and only 3 potential breeders have returned in successive years. No breeding was observed by returning adults. Drift fences at 4 recipient ponds captured 77 (4%) released individuals leaving ponds to circumjacent uplands. Five individuals were observed in later years returning to breed in their release pond, but breeding apparently did not occur. Researchers must conclude at this time that repatriated populations are not yet robust enough for long-term viability, and therefore repatriation results do not yet positively affect the status of panhandle populations (Means et al. 2017; Ryan Means, pers. commun. 2020).

Table 2. The number of known ponds with records since 1997 and the number of presumably extant subpopulations of the striped newt by site, plus the last year the species was observed at a site.

No. No. Site Ponds Subpopulations Last Year Apalachicola National Forest 18 1 2018 Big Bend Wildlife Management Area 2 1 2019 Camp Blanding Military Reservation 16 8 2018 Faver-Dykes State Park 3 1 2014 Goethe State Forest 5 2 2010 Guana River Wildlife Management Area 3 1 2019 GTMNERR 1 1 2002 Jennings State Forest 18 5 2018 Livingston Place 5 1 2019 Merritt Island National Wildlife Refuge 1 1 2016 Ocala National Forest 44 11 2020 Ordway-Swisher Biological Station 13 3 2018 Pumpkin Hill Creek Preserve State Park 1 1 2006 Rock Springs Run State Reserve 4 3 2010 Seminole State Forest 2 2 2018 Triple N Ranch Wildlife Management Area 9 2 2017 Private land = Dee Dot Ranch 2 1 1999 Private land = Plum Creek Timberland 3 1 1998 Total 150 46

Some habitat practices may be detrimental to the Jefferson County population, where swaths of the uplands and dry wetland basins are roller chopped in fall to create walkable lanes for bobwhite quail (Colinus virginianus) hunters and their dogs and to reduce understory thickets. This practice disturbs and compacts soil, causes temporary siltation in wetlands when they fill, and possibly results in mortality of newts. Sub-criterion C1. – Johnson (2002b) trapped 2,072 adult striped newts during 1 immigration event at a pond at Ordway-Swisher Biological Station, Putnam County. This number does not include paedomorphs that may already have been present in the pond. Johnson (2002b) estimated that 25% of the population became paedomorphs. If this is the case, then the population in this pond contained >2,500 adults. In contrast, the most striped newts that Means (1999) trapped entering a pond in ANF during a 1-year period was 88 (presumably terrestrial adults). The most aquatic, nonpaedomorphic adults dipnetted was 23 in 90 person-minutes from a pond in Big Bend WMA–Spring Creek Unit, Taylor County (Kevin Enge, unpubl. data). The most paedomorphs dipnetted was 143 in 90 person-minutes from a permanent scrub pond in ONF, Marion County (Kevin Enge, unpubl. data). Hundreds of more paedomorphs were likely present in the latter pond. According to Farmer et al. (2017), there are 106 known breeding ponds in Florida with apparently extant populations, and 4 additional ponds have been discovered since then (Kevin Enge, unpubl. data). Therefore, the number of adult striped newts present in Florida far exceeds 10,000 individuals. All 5 BRG members agreed that the Florida population had >10,000 adults and therefore did not meet Criterion C1, but they considered the other sub-criteria anyway. All 5 BRG members agreed that the panhandle population did not have >10,000 adults but were uncertain if a population will go extinct in at least 1 of the 6 known occupied ponds (i.e., >10% decline) in the panhandle in the next 15 years; thus, Criterion C1 was not met. For example, 1 subpopulation in Jennings State Forest contains 11 known ponds and likely has >1,000 adults. Sub-criterion C2. – All 5 BRG members agreed that the panhandle population will experience a continuing decline in the number of adults, which is inferred from the decline experienced in ANF. The only natural population remaining in ANF may go extinct, and repatriation efforts there have not yet proven successful. Some habitat management practices at Livingston Place may be deleterious to striped newts. Climate change will likely result in more frequent and prolonged droughts, reducing population size. Sub-criterion C2a(i) is not met, because Livingston Place is suspected of having >1,000 adults; 1 large pond there appears to contain a robust population. Sub-criterion C2a(ii) is not met, because there are 2 subpopulations. However, all 5 BRG members agreed that Sub-criterion C2b is met regarding extreme fluctuations in the number of adults for reasons already discussed for Sub-criterion B2c. Sub-criterion D1. ‒ All 5 BRG members agreed that >1,000 adult striped newts occur in both the Florida and panhandle populations. Johnson (2002b) documented >2,000 adults in 1 peninsular pond, but this might not be representative of panhandle ponds. However, the 6 panhandle ponds with self-sustaining populations would only need to average 167 adults to have a population size of 1,000 adults, and this does not include newts released in the ANF repatriation ponds. As discussed for Sub-criterion C2, Livingston Place is suspected of having >1,000 adults. Sub-criterion D2. ‒ As discussed for Sub-criterion B2, the estimated AOO for the Florida population far exceeds 20 km2. The Florida population also has >5 locations; thus, it does not meet Sub-criterion D2. The 5 BRG members agreed that the panhandle population has ≤5 locations. One location is at Livingston Place in Jefferson County, and ANF has 1 or 2 locations, depending upon whether the repatriated subpopulation is counted as a separate location. The distance between the ponds with a natural and a repatriated population in ANF are separated by 5.6 km and a major highway. There was no need to determine the AOO. The other condition necessary is that the population is prone to the effects of human activities or stochastic events within a short time period in an uncertain future. The ANF population has already experienced a drastic population decline, and a prolonged drought exacerbated by groundwater withdrawal could eliminate the remaining population. In 2013, Tall Timbers Research and Land Conservancy acquired Livingston Place, which will continue to host the Continental Field Trial for pointing dogs and to manage for bobwhite quail. It is possible that human activities could deleteriously impact striped newt population in the future.

Criterion E. ‒ A population viability analysis (PVA) does not exist for the striped newt. An attempt at conducting a PVA was unsuccessful, so a potential habitat model was created instead for Florida (Endries et al. 2009). APPENDIX 1. Abbreviated biographies of the Striped Newt Biological Review Group members.

Kevin M. Enge received an M.S. in Wildlife Ecology and Conservation from the University of Florida and B.S. degrees in Wildlife and Biology from the University of Wisconsin–Stevens Point. He is an Associate Research Scientist in the Reptile and Amphibian Subsection of FWC’s Fish and Wildlife Research Institute. Kevin has worked for FWC since 1989, serving previously as a nongame survey and monitoring biologist and the Herp Taxa Coordinator. He has conducted surveys of many native and exotic amphibians and reptiles, including the striped newt. He has authored 10 publications on the striped newt and coauthored a 2019 book entitled Amphibians and Reptiles of Florida. Kevin served as the lead on biological status reviews of 13 species or populations. Aubrey Greene received an M.S. in Entomology from Florida A&M University and B.S. in Animal Ecology with an emphasis on wildlife from Iowa State University. She is a Research Associate in the Reptile and Amphibian Subsection of FWC’s Fish and Wildlife Research Institute. Aubrey has spent the past 14 years working on research and the conservation of a variety of Southeast taxa including reptiles, amphibians, birds, aquatic invertebrates, and small mammals. Since joining FWRI in Fall 2016, Aubrey has worked primarily with Florida’s declining amphibian species, including the striped newt, gopher frog, flatwoods salamander, and ornate chorus frog.

Steve A. Johnson received his Ph.D. from the University of Florida and M.S. and B.S. degrees from the University of Central Florida. He is an Associate Professor of Wildlife Ecology at the University of Florida. Previously, he worked as a Research Wildlife Biologist for the U.S. Geological Survey’s Amphibian Research and Monitoring Initiative. Steve’s area of expertise is natural history and conservation of amphibians and reptiles. His dissertation research was conducted on ecology of the striped newt, and he has 14 publications on this species. He, his students, and collaborators conduct research on numerous species of reptiles and amphibians, including pond-breeding amphibians such as the striped newt. In 2018, he received the Paul Moler Herpetological Conservation Award for his career-long commitment to conservation of amphibians and reptiles.

Ryan C. Means received an M.S. in Wildlife Ecology and Conservation and B.S. in Zoology from the University of Florida. He is the President of the Coastal Plains Institute, where he has worked as a conservation biologist since 1997. Ryan’s scientific research has focused primarily in herpetology, ephemeral wetland ecology and conservation, and southeastern ecosystem conservation. He is the principal investigator of the striped newt repatriation project, a unique salamander conservation effort in the panhandle that includes many partners united in a common goal to repatriate an extirpated species back into historically occupied ephemeral wetlands. Ryan has helped monitor striped newt breeding ponds in the Apalachicola National Forest for over 20 years, and he co-petitioned the U.S. Fish and Wildlife Service to list the striped newt.

Dirk J. Stevenson received a B.S. in Zoology from Southern Illinois University. He is the Lead Biologist and Owner of Altamaha Environmental Consulting. Dirk has worked in the Southeast. for over 25 years, primarily conducting field studies of reptiles and amphibians, including surveying striped newt breeding ponds in Georgia. He has worked for The Nature Conservancy, the Department of Defense, FWC, and The Orianne Society. Dirk has published many technical and popular articles on the eastern indigo snake, turtles, salamanders, dragonflies, and arachnids. Many of his photographs appeared in the 2008 book entitled Amphibians and Reptiles of Georgia, and his popular articles have appeared in Wildlife Conservation, Alabama Wildlife, South Carolina Wildlife, Illinois Audubon, and Georgia Backroads. He is a regular nature columnist for The Savannah Morning News.

APPENDIX 2. Peer Reviews of Independent Scientists.

Peer review #1 from Dr. Kenneth Dodd, University of Florida

Peer Review #2 from Sarah May

Peer Review #3 from Lora Smith, Jones Center at Ichauway

APPENDIX 3. Summary of letters and emails received during the solicitation of information from the public.

Pursuant to subsubparagraph 68A-27.0012(2)(c)2.b., F.A.C., the Florida Fish and Wildlife Conservation Commission requested information from the public pertaining to the biological status of the striped newt, including information and data on: population size and trends; distribution and range; threats to the species; published population viability models; and specific aspects of the species’ life history that may influence the status of the species.

This solicitation was noticed in the Florida Administrative Register, and two periods were held for the public to submit information on the biological status of the striped newt: the first from July 2nd, 2019 through July 15th, 2019, and the second from December 17th, 2020 through February 10th, 2021.

No emails or letters were received during either solicitation period.