Species Status Assessment Report Holiday Darter

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Species Status Assessment Report

for the Holiday Darter (Etheostma brevirostrum)

Version 1.0

Photo from Fishes of Alabama and the Mobile Basin (Mettee et al. 1996, p. 595)

July 2017

U.S. Fish and Wildlife Service

Region 4

Atlanta, GA

Species Status Assessment Report for Holiday Darter (Etheostoma brevirostrum) Prepared by the U.S. Fish and Wildlife Service

EXECUTIVE SUMMARY

This species status assessment (SSA) reports the results of the comprehensive status review for the holiday darter (Etheostoma brevirostrum), documenting the species’ historical condition and providing estimates of current and future condition under a range of different scenarios. The holiday darter is small fish native to the upper Coosa River basin in Georgia and Tennessee that occurs in small rivers with good water quality.

The SSA process can be categorized into three sequential stages. During the first stage, we consider the holiday darter’s life history and use the conservation biology principles of resiliency, redundancy, and representation (together, the 3Rs) to better understand the “needs” of populations and the species to maintain viability. The next stage involved an assessment of the historical and current condition of the species’ demographics and habitat characteristics. The final stage of the SSA involved making predictions about future viability while considering the species’ responses to anthropogenic and environmental influences that are likely to occur within its range. This process used the best available information to characterize viability as the ability of a species to sustain populations in the wild over time.

To evaluate the current and future viability of the holiday darter, we assessed a plausible range of conditions that affect its habitats to allow us to forecast the species’ resiliency, representation, and redundancy. For the purposes of this assessment, populations were delineated using U.S. Geological Survey 10 digit Hydrologic Unit Codes (HUC10s) that are occupied by the species.

Resiliency, assessed at the population level, describes the ability of a population to withstand stochastic events. A species needs multiple resilient populations distributed across its range to persist into the future and avoid extinction. A number of factors, including (but not limited to) water quality, water quantity, habitat connectivity, and instream substrate, may influence whether holiday darter populations will occupy available habitat. As we considered the future viability of the species, more populations with high resiliency distributed across the known range of the species can be associated with higher species viability. As a species, the holiday darter has limited resiliency, with the majority of populations considered to be in low resiliency.

Redundancy describes the ability of a species to withstand catastrophic events. Measured by the number of populations, their resiliency, and their distribution (and connectivity), redundancy gauges the probability that the species has a margin of safety to withstand or can bounce back from catastrophic events (such as a rare destructive natural event or episode involving many populations). Redundancy for the holiday darter is characterized by having multiple, resilient and representative populations distributed throughout its range. Furthermore, these populations

should maintain natural levels of connectivity between them. Connectivity allows for immigration and emigration between populations and increases the likelihood of recolonization should a population become extirpated.

Representation describes the ability of a species to adapt to changing environmental conditions. Representation can be measured by the breadth of genetic or environmental diversity within and among populations and gauges the probability that a species is capable of adapting to environmental changes. The more representation, or diversity, a species has, the more it is capable of adapting to changes (natural or human caused) in its environment. In the absence of species-specific genetic and ecological diversity information, we evaluate representation based on the extent and variability of habitat characteristics across the geographical range.

The analysis of species’ current condition revealed that the extent of occupied habitat has declined, with most populations currently occupying 20-80% of the historically occupied ranges. All historically known populations remain extant; however, there is little connectivity between them and abundances are qualitatively low.

To assess the future condition of the holiday darter, a variety of stressors from urbanization, agriculture, and climate were considered. Populations with low resiliency are considered to be more vulnerable to extirpation, which, in turn would decrease species’ level representation and redundancy. To help address uncertainty associated with the degree and extent of potential future stressors, the 3Rs were assessed using three plausible future scenarios. These scenarios were based, in part, on the results of urbanization and climate models.

An important assumption of the predictive analysis was that future population resiliency is largely dependent on water quality, water flow, and structural habitat conditions. Our assessment predicted that all currently extant holiday darter populations would experience negative changes to these important habitat requisites in the future under the most likely scenario, Status Quo.

Given scenario 1, the “Status Quo” option, loss of resiliency, representation, and redundancy is expected. Under this secenario, we predicted that three populations would likely be extirpated and the remaining populations would have low resiliency. The extirpation of three populations would reduce redundancy by 43%. Representation would be reduced due to the loss of the only known population in the Coosawattee River basin and range contraction out of Piedmont and Ridge and Valley physiographic provinces.

Given scenario 2, the “Best Case” option, we predicted slight improvements to resiliency from the current conditions in one populations although no populations would exhibit “high” resiliency. Two would exhibit “moderate” resiliency, three would exhibit “low” resiliency, and the remaining two populations are expected to become extirpated. Redundancy and representation would decline from current conditions.

Given scenario 3, the “Worst Case” option, loss of resiliency, representation, and redundancy is expected. Under this scenario, we predicted that four populations would likely be extirpated and those that remain would have low resiliency. Redundancy would be reduced overall by 57% in. Representation would be reduced due to the loss of the only known populations in the Coosawattee River basin and Alabama as well as range contraction out of Piedmont and Ridge and Valley physiographic provinces.

Table E1. Current estimated resiliency and predicted future resiliency of populations under multiple scenarios

Population Current Status Quo Best Case Worst Case Conasauga River Low Low Moderate Low Talking Rock Likely Likely Likely Low Creek Extirpated Extirpated Extirpated Mountaintown Likely Likely Low Low Creek Extirpated Extirpated Likely Likely Likely Ellijay River Low Extirpated Extirpated Extirpated Amicalola Creek Moderate Low Moderate Low Etowah River Low Low Low Low Likely Shoal Creek Low Low Low Extirpated

Overall Summary

Currently, the holiday darter continues to occupy all streams where it was historically known to occur. Based on collection records from the last 10 years, all populations occur over shorter overall stream lengths than historical records, indicating range reduction in these seven populations. No population of holiday darter currently exhibits high resiliency due to the reduction in extent of occupied habitat, low abundance of individuals per population, a simple linear spatial arrangement of records, as well as stressors affecting habitat and water quality. Similarly, representation and redundancy is currently low for this species because multiple resilient populations are lacking, connectivity is limited among populations, and this species is increasingly becoming isolated to the upstream limits of its range in the Blue Ridge physiographic province.

Our future scenarios assessment considered the current viability of the species to project likely future viability given plausible scenarios of urban development and climate change. Under no scenario did the species persist in all known populations. Two populations are likely to become extirpated under all scenarios. Under the Status Quo Scenario, a third population is expected to become extirpated and under the Worst Case Scenario, a total of four populations are expected to become extirpated. Resiliency, representation, and redundancy declined in all scenarios due to

4 further range contractions and increased likelihoods for extreme climatic events to impact populations.

Table of Contents

EXECUTIVE SUMMARY……………………………………………………………………….2 CHAPTER 1 – INTRODUCTION………………………………………………………………..8 CHAPTER 2 – SPECIES BIOLOGY AND INDIVIDIUAL NEEDS…………………………..10 Taxonomic History and Uncertainty……………………………………………………..10 Physical Description…………………………………………………………...………...10 Range and Distribution…………………………………………………………………..11 Biology, Life history, Ecology……………………………………………………………11 CHAPTER 3 – POPULATION AND SPECIES NEEDS……………………………………….14 Holiday Darter Resiliency………...……………………………………………………..14 Holiday Darter Representation…………………………………………………………..16

Holiday Darter Redundancy……………………………………………………………..17 CHAPTER 4 – FACTORS INFLUENCING VIABILITY……………………………………...18 Urbanization……………………………………………………………………………..18

Agriculture……………………………………………………………………………….22 Sedimentation…………………………………………………………………………….23

Loss of Riparian Vegetation……………………………………………………………...24

Weather events…………………………………………………………………………...25

Conservation Measures………………………………………………………………….26

CHPATER 5 – CURRENT WATERSHED AND POPULATION……………………………..28 Methods…………………………………………………………………………………..28 Current Condition………………………………………………………………………..33

CHAPTER 6 – FUTURE VIABILITY…………………………………………………………..51

Methods…………………………………………………………………………………..51

Status Quo………………………………………………………………………………..53

Best Case…………………………………………………………………………………69

Worst Case……………………………………………………………………………….75

Summary…………………………………………………………………………………79

LITERATURE CITED…………………………………………………………………………..82

Chapter 1. Introduction

The holiday darter is a freshwater fish found in the Coosa River System in the Piedmont, Blue Ridge and Ridge and Valley physiographic provinces. This species was petitioned for federal listing under the Endangered Species Act of 1973, as amended (Act), as part of the 2010 Petition to List 404 Aquatic, Riparian and Wetland Species from the Southeastern United States by the Center for Biological Diversity (CBD 2010, p. 425).

The Species Status Assessment (SSA) framework (USFWS 2016, entire) is intended to be an in- depth review of the species’ biology and threats, an evaluation of its biological status, and an assessment of the resources and conditions needed to maintain long-term viability. The intent is for the SSA Report to be easily updated as new information becomes available and to support all functions of the Endangered Species Program from Candidate Assessment to Listing to Consultations to Recovery. As such, the SSA Report will be a living document upon which other documents, such as listing rules, recovery plans, and 5-year reviews, would be based if the species warrants listing under the Act.

The SSA Report is not a decisional document by the U.S. Fish and Wildlife Service (Service); rather it provides a review of available information strictly related to the biological status of the holiday darter. The listing decision will be made by the Service after reviewing this document and all relevant laws, regulations, and policies, and the results of a proposed decisions will be announced in the Federal Register, with appropriate opportunities for public input.

For the purpose of this assessment, we generally define viability as the ability of the holiday darter to sustain populations in natural river systems over time. Using the SSA framework (Figure 1.1), we consider what the species needs to maintain viability by characterizing the status of the species in terms of its resiliency, redundancy, and representation (Wolf et al. 2015, entire).

Figure 1. Species Status Assessment Framework

• Resiliency describes the ability of populations to withstand stochastic events (arising from random factors). We can measure resiliency based on metrics of population health; for example,

species abundance and complexity of their spatial occurrences. Highly resilient populations are better able to withstand disturbances such as random fluctuations in birth rates (demographic stochasticity), variations in rainfall (environmental stochasticity), or the effects of anthropogenic activities.

• Representation describes the ability of a species to adapt to changing environmental conditions. Representation can be measured by the breadth of genetic or environmental diversity within and among populations and gauges the probability that a species is capable of adapting to environmental changes. The more representation, or diversity, a species has, the more it is capable of adapting to changes (natural or human caused) in its environment. In the absence of species-specific genetic and ecological diversity information, we evaluate representation based on the extent and variability of habitat characteristics across the geographical range.

• Redundancy describes the ability of a species to withstand catastrophic events. Measured by the number of populations, their resiliency, and their distribution (and connectivity), redundancy gauges the probability that the species has a margin of safety to withstand or can bounce back from catastrophic events (such as a rare destructive natural event or episode involving many populations).

To evaluate the biological status of the holiday darter both currently and into the future, we assessed a range of conditions to allow us to consider the species’ resiliency, redundancy, and representation (together, the 3Rs). This SSA Report provides a thorough assessment of biology and natural history and assesses demographic risks, stressors, and limiting factors in the context of determining the viability and risks of extinction for the species. This document is a compilation of the best available scientific and commercial information and a description of past, present, and likely future risk factors to the holiday darter.

Chapter 2: Species Biology and Individual Needs

Taxonomic History and Uncertainty

The holiday darter (Etheostoma brevirostrum) was formally described in 1991 by Royal Suttkus and David Etnier (Suttkus and Etnier 1991, entire). The type locality was designated as Shoal Creek at Pine Glen Recreation Area in Cleburne County, Alabama. At the time of description the species was considered endemic to the Coosa River system, above the fall line in the Blue Ridge and Valley and Ridge physiographic provinces. The distribution was defined as disjunct with populations occurring in Shoal Creek, Alabama, the upper Conasauga River system in Georgia and Tennessee, and the Coosawattee River and Etowah River systems in Georgia.

In 1997, researchers noted that the populations of holiday darter in the upper Etowah River and Amicalola Creek each likely represented distinct species from the holiday darter described from Shoal Creek based on subtle differences in the breeding coloration of males (Burkhead et al. 1997, p. 393). In 2005, Freeman and others distinguished the populations of Holiday darter in the Conasauga and Coosawattee Rivers as two additional distinct and undescribed species, for a total of five species in the Holiday darter species complex within the Coosa River system (Freeman et al. 2005, p. 584). The most current status of imperiled North American freshwater and diadromous fishes published by the American Fisheries Society (Jelks et al. 2008, p. 404) evaluates each population in this hypothesized species complex separately, an approach that was further advocated by Anderson and others (2012, p. 38) in their analysis of holiday darter occupancy in the upper Etowah River system. In the absence of a published species description, that formally splits the holiday darter into distinct species, we consider the holiday darter as a single species with a disjunct distribution consisting of seven populations (Conasauga River, Talking Rock Creek, Mountatintown Creek, Ellijay River, Amicalola Creek, Etowah River, and Shoal Creek) in this document and are within the boundaries of USGS Hydrologic Unit Code ten digit (HUC 10) watersheds. Each of these populations is considered an important component to representation as they represent irreplaceable variation within the species.

Physical Description

The Holiday darter is a member of the Ulocentra (snubnose darter) subgenus and was placed in the duryi group (Suttkus and Etnier 1991, p. 17). Adults reach a maximum standard length (SL) of 1.8 in (47 mm) in females to 2 in (53 mm) in males (Suttkus and Etnier 1991, p. 17). Male holiday darters are colorful fish, with notable red blotches surrounded by white or yellow halos on the lower side of the body. Unique from similar species with which it co-occurs, the holiday darter has a distinct median red band across the generally blue-green anal fin in males in spawning color. Eight saddles are along the back with eight to nine blotches along the side. Blotches from the origin of the anal fin to the caudle peduncle (base of tail fin) have a green ventral extension in males (Suttkus and Etnier 1991, p. 18; Etnier and Starnes 1993, p. 554).

Range and Distribution

Historical Distribution

The Holiday darter is known to have a disjunct distribution within the upper Coosa River basin. Occurrence records for this species are above the fall line in the Ridge and Valley and Blue Ridge provinces of Alabama, Georgia, and Tennessee. Historically, records for the Holiday darter existed for Shoal Creek (tributary to Choccolocco Creek in Alabama) and from three large springs in the Chocolocco Creek system within Talladega National Forest, the Conasauga River in Tennessee and Georgia, tributaries to the Coosawattee River in Georgia, and the upper Etowah River (inclusive of Amicalola Creek) in Georgia.

Current Distribution

Currently the Holiday darter is known to occur in parts of Shoal Creek, Conasauga River, Talkingrock Creek, Mountaintown Creek, tributaries of the Ellijay River, Amicalola Creek, and the Etowah River. The Holiday darter has not been collected from the spring tributaries of upper Choccolocco Creek since 1987 (Sizemore and Howell 1990). There are no records for the Holiday darter within the Coosawattee River mainstream.

Biology, Life history, Ecology

An in-depth life history study has not been undertaken for the holiday darter. Therefore we must infer some life history characteristics from studies conducted on other species within Ulocentra subgenus.

Like other species of snubnose darter, the holiday darter likely has an approximate three year lifespan and is a benthic omnivore that primarily feeds on aquatic insect larvae and microcrustaceans (Etnier and Starnes 1993, p. 554). Studies have been undertaken that describe habitat types and spawning and are discussed below.

Habitat

The holiday darter is found in small creeks to moderate sized rivers (Etnier and Starnes 1993, p. 554). This pattern can certainly be seen in Alabama, where the species at least historically occurred in small spring-fed tributaries of Choccolocco Creek as well as parts of Shoal Creek that range from 32 ft - 65 ft (10 - 20 m) wide. Dominant substrate types associated with the species are boulders, cobble, and gravel. American water-willow (Justicia americana) and hornleaf riverweed (Podostemum ceratophyllum), two aquatic plants, are commonly associated with holiday darters across its range. The holiday darter prefers riffles and shallow areas of rivers at depths ranging from 8.6 in – 13.3 in (22 - 34 cm) deep in moderate to fast currents (1.77 – 2.66 ft/second; 0.54-0.81 meters/second; Johnston and Shute 1997, p. 2). All populations occur in generally clear streams with low turbidity and cool water temperatures with populations in

Tennessee and Georgia found to occur in cooler water temperatures than observed in Shoal Creek (Etnier and Starnes 1993, p. 554; Boschung and Mayden 2005, p. 516).

Reproduction

Spawning takes place from April - May at temperatures of 51.1˚ - 64.2 ˚ F (10.6 - 17.9° C) based on the collection of males in breeding coloration and direct observation of spawning activity and breeding behavior (aggression and courtship) has been observed from April - June at temperatures of 50.9˚ - 68.5˚ F (10.5 - 20.3° C; Anderson 2009, p. 42). Spawning begins with a period of courtship where a male follows a female. Often secondary males may try to disrupt the courting pair. Holiday darters utilize an egg-attaching spawning behavior. The male follows the female until suitable spawning substrate is selected by the female. Suitable spawning substrate is typically coarse material ranging from gravel to rock and sometimes woody material. Females position themselves vertically on the spawning substrate and are mounted by the male. A period of intense quivering ensues during which a single egg and sperm are released. Breeding is promiscuous and spawning sites may or may not be reused. Spawning habitats are generally similar to typical adult habitat. Spawning depth ranges from 0.5 – 2.5 feet (15-76 cm) at a velocity of 0.32 feet/second (0.1 meter/second; Johnston and Shute 1997, entire; Anderson 2009, p. 42)

Fecundity ranged from 50-150 mature eggs in four females from the Amicalola Creek system, with egg size ranging from 1.2-1.5 mm (Burkhead, unknown date).

Table 1. Overview of needs of an individual holiday darter based on our knowledge of the species’ biology, ecology, and life history summarized above. *Based on the biology of another Ulocentra in the duryi group: Etheostoma sp. cf. E. zonistium (Blueface darter)

Life Stage Resources Needed Information Source Fertilized Egg Clean hard substrate such as Anderson 2009, p. 40; rocks or woody debris, Boschung and Mayden 2007, flowing water p.516 Larvae Clear flowing water, Douglas 2013, p. 10 connectivity to downstream habitat to accommodate a pelagic larval duration of 60 days*

Juveniles Clear flowing water, adequate Anderson 2009, p. 34 food availability, presence of instream structures (woody debris, large rocks)

Adult Clear flowing water, adequate Anderson 2009, p. 34 food availability, presence of instream structures (woody debris, large rocks)

Chapter 3: Population and Species Needs

Holiday Darter Resiliency

Each population of the holiday darter needs to be able to withstand or be resilient to stochastic events or disturbances. These are events that are reasonably likely to occur and can drastically alter the ecosystem. Examples of stochastic events that may affect holiday darters include drought, major storms and flooding, or accidental discharge of pollutants into streams.

To be resilient to stochastic events populations of holiday darter need to have an adequate number of individuals (abundance). They should cover a large enough area such that small localized events do not cause extirpation (spatial extent). Finally the area occupied by the population needs to exist in multiple tributaries (spatial complexity) so a single event (such as a spill of toxic chemicals) cannot eliminate an entire population as it propagates downstream.

Population level characteristics (abundance, spatial extent, and spatial complexity) that influence resiliency are controlled by environmental conditions present in the upper Coosa River system. Population abundance, spatial extent and spatial complexity are results of successful spawning, recruitment (survival of young individuals to maturity and spawning), adult survival, and dispersal. Spawning, survival, and dispersal are all influenced by aspects of water quality, water quantity, structural habitat, and connectivity (as indicated by Table 1 and Figure 4).

Water Quality

Holiday darters, like other benthic species are sensitive to poor water quality (Warren et al. 1997, p. 125). Broadly, good water quality for the holiday darter consists of high amounts of dissolved oxygen (DO), moderate pH (slightly acidic in the Blue Ridge to slight basic in the Ridge and Valley physiographic province), unaltered temperature regimes, and little to no pollutants present. Degraded water quality has the potential to induce stress on individuals, reduce spawning success, or cause direct mortality.

Water Quantity (Flow Regime)

In addition to water quality, holiday darters have adapted to the perennial flows and seasonal predictability of their preferred habitat in small rivers and streams. Low flows can negatively affect water quality parameters (temperature and DO), prevent fine sediments, contaminants, and excess nutrients from being flushed downstream, and lead to temporary eutrophication (thereby reducing DO). Low flows can also concentrate holiday darters into pools exposing them to predation or stress resulting from high density of organisms concentrated in wetted pools. Conversely, extremely high flows can also negatively affect populations of holiday darters. Extensive scour and loss of habitat along river margins after extreme storms and flooding can cause localized declines in abundance of fish species (USFWS 2011, p. 9). Finally, altered flow regimes seen downstream of dams and around urban areas can negatively alter structural habitat

through scouring, decouple timing of life history patterns from requisite flows, and alter temperature regimes, DO concentrations, and sedimentation (Bunn and Arthington 2002, entire).

Structural Habitat

Structural habitat provides the necessary physical features for holiday darters to spawn, forage, and seek refuge from predators or high flows. Structural habitat refers to substrate (gravel, cobble, and sand), boulders, and large woody debris utilized by holiday darters (see Table 1). Clean substrates (free of fine sediment) ensure adequate DO levels are available for developing eggs. Additionally, a lack of fine sediments maintains adequate interstitial spaces (voids underneath and between rocks and logs) for the holiday darter to feed and shelter. The small rivers where the holiday darter is found receive woody material from riparian areas that provide additional structural habitat that is used by holiday darter for foraging. Woody material and other terrestrial plant material, such as leaves, that enters streams and rivers are the primary source of nutrients and form the base of the food web for the river ecosystem where the holiday darter occurs. Therefore, terrestrial plant material provides structure for shelter and foraging and directly influences food resources on which the holiday darter depends (Vannote et al. 1980, p. 132).

Connectivity

Connectivity, for the purpose of this assessment, refers to a species’ ability to disperse to and from habitat patches (Gido et al. 2010, p 293) and is influenced by natural or artificial features present on a river or stream (e.g. waterfalls or dams). We consider that longitudinal connectivity (movement parallel with the stream flow in an upstream or downstream direction) to be the most relevant for the holiday darter. The holiday darter has adapted to an ecosystem that would naturally provide connectivity (with the exception of isolation as indicated by geographic variation). Therefore, the holiday darter has adapted to an ecosystem that would allow it to repopulate areas where localized extirpation during natural stochastic events may have occurred. Additionally, natural connectivity would allow the holiday darter to maintain gene flow across the multiple occupied rivers. Therefore, factors that reduce connectivity (dams or culverts that act as barriers) would limit dispersal, reduce the species’ ability to recolonize after disturbance events, limit gene flow, increase genetic drift, reduce genetic variability within populations, increase genetic variability among populations, and culminate in an increased risk of extirpation and extinction (Gido et al 2010, p. 296; Gaggiotti 2003, p. 160).

Figure 2. A simple influence diagram illustrating habitat factors that influence breeding, feeding, and sheltering factors, which in in turn affect demographic factors that ultimately drive fish population growth and maintenance.

Holiday Darter Representation

Representation describes the ability of a species to adapt to changing environmental conditions over time and encompasses the “ecological and evolutionary patterns and processes that not only maintain but also generate species” (Shaffer and Stein, p. 308). It is characterized by the breadth of genetic and environmental diversity within and among populations. For the holiday darter to exhibit adequate representation, resilient populations should occur in all physiographic provinces to which it is native (Blue Ridge, Ridge and Valley, Piedmont). These occupied physiographic provinces represent the ecological setting in which the holiday darter has evolved. Additionally, evolutionary patterns that are exhibited by the morphological, genetic, and behavioral variation that exists within the species should be maintained. For the holiday darter, this variation has currently been identified as morphological differences between Shoal Creek, Amicalola Creek, Upper Coosawattee River, Etowah River, and Conasauga River populations. Finally, natural levels of connectivity are important to be maintained between representative populations because it allows for the exchange of novel and beneficial adaptations where connectivity is high or is the mechanism for localized adaption and variation where connectivity is lower and the species is naturally more isolated.

Holiday Darter Redundancy

Redundancy describes the ability of a species to withstand catastrophic events. It “guards against irreplaceable loss of representation” (Redford et al. 2011 p. 42; Tear et al. 2005 p. 841) and minimizes the effect of localized extirpation on the range-wide persistence of a species (Shaffer and Stein, p. 308). Redundancy for the holiday darter is characterized by having multiple, resilient and representative populations distributed throughout its range. Furthermore, these populations should maintain natural levels of connectivity between them. Connectivity allows for immigration and emigration between populations and increases the likelihood of recolonization should a population become extirpated.

Figure 3. Resiliency, representation, and redundancy are interrelated conservation biology principles that can be used to evaluate the current and future condition of a species. Some components that influence resiliency, representation, and redundancy are provided.

Chapter 4: Factors Influencing Viability

The upper Coosa River basin has been identified as a priority region for conservation by numerous agencies and researchers (Service, Georgia Department of Natural Resources, University of Georgia). Efforts have been made in parts of this region to implement watershed wide conservation plans (Etowah Habitat Conservation Plan and Conasauga Working Lands for Wildlife (WLFW)). Because of the high level of interest by researchers and management agencies in the area, factors that affect aquatic habitats in the upper Coosa River basin have been identified and described (Freeman et al. 2002, entire; Wenger and Freeman 2007, entire). The current and potential future effects of these factors, along with information about populations help to inform species viability and vulnerability to extinction.

Urbanization

Urbanization refers to a change in land cover and land use from forests or agriculture to increased density of residential and commercial infrastructure. Urbanization is expected to affect the holiday darter across its range due to the majority of known localities occurring in close vicinity to the Atlanta metropolitan area and areas with growing populations and increasing development that exist between Chattanooga and Atlanta. Urbanization introduces a multitude of stressors into lotic systems that co-vary and have synergistic effects that are difficult to disentangle (Matthaei and Lang 2016, p. 180). Streams affected by urbanization have been described to exhibit an “urban stream syndrome” (Matthaei and Lang 2016, p. 180; Wenger et al. 2009, entire; Walsh et al. 2005, p. 207). The urban stream syndrome consistently includes “a flashier hydrograph, elevated concentrations of nutrients and contaminants, altered channel morphology and stability, and reduced biotic richness, with an increased dominance of species tolerant to poor water quality and variably includes reduced baseflow and increased suspended solids” (Walsh et al. 2005, p. 207; Paul and Meyer 2001, entire). Therefore, where urbanization occurs, it is anticipated to increase the magnitude of nearly all stressors present within the holiday darter’s occupied range.

Water Quantity

A major feature of urbanized areas is an overall increase in impervious surfaces. Impervious surfaces can be defined as hard surfaces that preclude water infiltration such as paved roads, parking lots, roofs, and even highly compacted soil like sports fields. Runoff from impervious surfaces directly affects stream flows and water quantity by altering the natural hydrologic cycle of streams and rivers and introduces more flow variability (flashy flows). In a natural forested system, most rainfall soaks in to the soil and is carried into nearby streams via subsurface flow. Some evaporates or transpires, and a relatively small amount travels to streams via overland flow. Generally, in a natural, forested Southeastern watershed, there is a time-lag between a rainfall event and peak flows after a rain event, peak flows are moderated due to evaporation,

transpiration, and percolation into groundwater, and baseflows are maintained by groundwater input that is consistently recharged due to percolation of rainwater (Figure 6). However, in an urbanized system with a large proportion of impervious cover, most stormwater is rapidly conveyed into streams due to a lack of infiltration and diverted quickly to streams via stormwater drain pipes or ditches. As a result, storm flows in the receiving stream are higher and more frequent, although shorter in duration, and base flows are lower due to a lack of groundwater recharge in an urban stream (see Figure 6). The storm discharge of urban streams can be twice that of rural streams draining watershed of similar size (Pizzuto et al. 2000, p. 81, Rose and Peters 2000, p. 1454). The holiday darter can be directly affected by flashy flows patterns induced by urbanization. The flashy flows as a result of urbanization may cause stress, displacement, or mortality (Konrad and Booth 2005, pp. 160-161) and it can decouple life- history cues and reduce spawning success (Bunn and Arthington 2002, p. 497).

Figure 4. A comparison of a forested hydrograph and urban hydrograph. Black bars represent a rain event, the solid line represents stream discharge in a forested watershed, and the dotted line represented discharge in an urbanized watershed. From Walsh et al. 2005.

In addition to direct effects on holiday darters, flashy stream flows and frequent, smaller high- flow events seen in urban streams negatively affects structural habitat on which the species depends. “Reduction in channel complexity, and thus instream habitat, appears an almost universal symptom of the urban stream syndrome” (Walsh et al. 2005, p. 711). This decrease in channel complexity can be seen as channel straightening (either through engineering or hydrologic processes) and a reduction in in stream cover and natural substrates like boulders, cobble, and gravel. Another result of frequent, smaller high-flow events is the low retention time (i.e. it is removed from the system quickly) of large woody structure and other terrestrial plant material (Walsh et al. 2005, p. 711). As a result, urban streams have lower amounts of plant

19 material and holiday darter food resources are negatively affected as well as habitat used for foraging and shelter.

Water Quality

“Increased concentrations of loads of several chemical pollutants in stream water appear universal in urban streams” (Walsh et al. 2005, p. 710). Pollutants, including metals, hydrocarbons, pesticides and other potentially harmful organic and inorganic compounds, are common in urban streams. Pesticides also are heavily used in urban and suburban areas, and many of these find their way into streams and groundwater. A comparison of agricultural and urban groundwater quality in the Mobile Basin (which includes the Coosa River basin) found a large variety and frequency of pesticide compounds in the urban groundwater (Robinson 2003, p. 27). Chlordane and other now-banned organochlorine pesticides are still common in urban streams, including those in the Mobile Basin (Zappia 2002, p. 53). Streets and parking lots can contribute large quantities of heavy metals that are largely derived from automobiles (Van Hassel et al. 1980, p. 642; Bannerman et al. 1993, p. 46). Oil and other hydrocarbons are also common constituent in urban runoff (Fam et al. 1987, p. 1045).These pollutants tend to accumulate on impervious surfaces due to the lack of infiltration and are subsequently washed into streams during rain events. In addition to pollutants that accumulate on impervious surfaces, wastewater discharges and leaky septic systems can input excess nutrients, fecal coliforms, household chemicals, and pharmaceuticals in to urban streams. Other declines in water quality parameters are observed as stream temperatures elevating to stressful levels due to stormwater becoming superheated on impervious surfaces and lower dissolved oxygen. This overall degradation of water quality makes urban streams inhospitable to aquatic species that have adapted to habitats in cool, clear, flowing water, like the holiday darter.

Infrastructure Acting as Movement Barriers

As discussed earlier, increases in infrastructure is current and ongoing within the range of the holiday darter. This increase in infrastructure due to growing human populations has been generally identified as increases in housing density, commercial space, and associated impervious surface cover. However, other aspects of infrastructure can directly affect stream and river ecosystems. These are engineered structures that cross or are placed directly within streams and rivers (i.e. bridges, culverts, dams, pipelines, and telecommunication lines). Structures installed at road crossings (bridges and culverts), dams, and pipelines all have the potential to act as barriers to fish movement and reduce connectivity. Fishes can be particularly susceptible to a loss of connectivity resulting from movement and dispersal barriers because their movement is restricted to the stream network. In contrast, terrestrial organisms may be less susceptible to the introduction of a single barrier on the landscape because multiple routes between suitable patches may be available. In general, barriers to fish movement can limit drift of pelagic larvae to downstream reaches, block exchange of genetic material between populations, and increase a

population’s vulnerability to local extinction and prevent recolonization after extirpation has occurred.

Structures like dams, not only reduce connectivity within a watershed. They can substantially alter hydrology downstream, especially when operated for hydroelectric power generation (Freeman et al. 2001, p. 183, Power et al. 1996, p. 893). These dams are usually operated through hydropeaking, which only produce high flows when power generation is needed. Increases in flow frequency or intensity during hydropeaking operations can result in channel widening through bank erosion or deepening to accommodate the additional discharge unless the channel is physically constrained (Wolman 1967, p. 392; Arnold et al. 1982, p. 160; Booth 1990, p. 409). This results in increased downstream sedimentation and unstable beds, both of which degrade spawning habitat, channel complexity, and feeding and shelter habitat for riffle-dwelling species that rely on sediment-free gravel such as the holiday darter.

Non-hydropeaking reservoirs (farm ponds, amenity lakes, and other impoundments) may also substantially alter hydrologic regimes by storing water during low flow periods, effectively dampening moderate to high flows and in some cases augmenting flows. The demand for these reservoirs will increase as human populations increase to accommodate water needs. For instance, estimated water withdrawal for the Metro Atlanta Water District (which includes a portion of the holiday darter range) was 687.0 mgd while it was permitted to use up to 882 mgd and current water use budgets anticipate permitting 1,140 mgd by 2035 (Metropolitan North Georgia Planning District 2009, p. 2 - 7). Surface water has and will continue to be the major supply of fresh water in this region due to the bedrock geology (Metropolitan North Georgia Planning District 2009, p. 2 - 7).Therefore, to provide for the increase in water demand, storage reservoirs have been proposed in the upper Coosa River basin. Depending on the location of drinking water supply reservoirs, holiday darter habitat may be disconnected or destroyed.

Infrastructure placed within occupied habitats has the potential to directly destroy physical habitat within the footprint of the structure. It can reduce connectivity which limits dispersal, reduces the species’ ability to recolonize after disturbance event, limits gene flow, increases genetic drift, reduces genetic variability (within populations), increases genetic variability among populations, and culminates in increased risk of extirpation and extinction (Gido et al 2010, p. 296; Gaggiotti 2003, p. 160). In addition to the above effects, dams can also substantially alter flows and negatively affect suitable downstream habitat and temperatures and the reservoirs formed upstream of dams converts suitable flowing water habitat into lake-like habitat. Currently, the holiday darter is exposed to these stressors from numerous road crossings, two large hydropower dams, and small run-of-river dams.

Agriculture

Agriculture is another predominant land use within the range of holiday darter. Within the Ridge and Valley province, row crop agriculture is prevalent and poultry farming is common throughout all northern Georgia. The effects of these agricultural practices on streams and rivers and the holiday darter are explained below.

Poultry Farming

Poultry production that occurs within the range of the holiday darter is undertaken primarily in poultry houses. In addition to chickens each poultry house has an estimated ability to produce approximately 100 tons of litter a year (assuming a 20,000 square foot poultry house stocked at one bird/square foot and six flocks produced/year, a probable underestimate of litter production per broiler house; Coufal et al. 2006, p. 403). Poultry litter is a mixture of chicken manure, feathers, spilled food, and bedding material that is used to fertilize pastureland or row crops that frequently occur adjacent to rivers and streams. Runoff from heavy rains carries excess nutrients from manure into nearby streams as a result of surface-spreading of litter. Litter can also contain arsenic, which is formed from a chemical routinely used as a feed additive to prevent disease and stimulate growth (Stolze et al. 2007, p. 821) and enters streams through runoff. Other substances often found in poultry litter included fecal coliform, salmonella, and other pathogens, pesticide residue, and other heavy metals (Bolan et al. 2010, pp. 676-683). In general, the input of compounds from poultry litter into rivers and streams can diminish water quality on which the holiday darter depends and cause physiological stress.

Estrogens, a type of endocrine disruptor that can be found in poultry litter, have been identified as a threat to the Conasauga River system (Jacobs 2015, entire). Increased levels of estrogens have been found to have numerous effects of fishes including: intersex individuals and testicular oocytes (Yonkos et al. 2010, p. 2338), decreased competitive behavior (Martinovic et al. 2007, p. 275), decreased sperm concentrations and decreased sperm mobility, and delayed spermatogenesis (Aravindakshan et al. 2004, p. 161). All of these effects lead to decreases in spawning success and potentially population collapse within short time frames (Kidd et al. 2007, p. 8899-8900). In a recent study of endocrine disruptors on fishes in the Conasauga River, approximately 7.5% of male fishes surveyed were found to have testicular oocytes (Jacobs 2015, p. 39). Studies have not been conducted to clarify the effects of endocrine disruptors on holiday darters; however, instances of intersex, testicular oocytes, and decreased reproductive health attributed to higher concentrations of endocrine disruptors has been observed in blackbanded (Percina nigrofasciata), speckled (Etheostma stigmaeum), rainbow (Etheostma caeruleum), and greenside (Etheostma blenniodes) darters (Jacobs et al 2015, p. 65; Tetreault et al. 2011, p. 287; Fuzzen et al. 2015, p. 111). High levels of estrogens are expected in other drainages that have similarly high numbers of poultry houses such as the Coosawattee River basin and the Amicalola River basin. Therefore, it is reasonable to infer that other rivers, besides the Conasauga River,

within the Coosa basin are affected by estrogens from poultry litter and the holiday darter is exposed to their negative effects in multiple portions of its range.

Other Contaminants

Pesticides and herbicides are frequently found in streams draining agricultural lands, with herbicides being the most commonly detected (McPherson et al. 2003, p. 44). Many agricultural streams still contain DDT and its degradation products (Zappia 2002, p. 50). Agricultural lands that surround occupied holiday darter habitat in Georgia have adopted “Roundup Ready” crops extensively. These GMOs were developed to survive applications of the herbicide Roundup and their prevalent use in agriculture corresponds with an increased use of the proprietary herbicide. “Roundup’s active ingredient is glyphosate, which impedes photosynthesis. Glyphosate is non- toxic to slightly toxic to most fish, although toxicity appears to be higher in several important sport or food fish, including brown trout, rainbow trout, channel catfish, bluegill, and tilapia (Kegley et al. 2016, entire). Roundup is commonly used in salt form (isopropylamine salt). This salt, as well as the surfactant normally found in Roundup (polyethoxylated tallowamine; POEA) and/or other ‘inert’ ingredients in the Roundup formulation appear more toxic to fish (Mitchell et al. 1987, p. 1032) and mussels than glyphosate alone, causing death of mussel glochidia (Bringolf et al. 2007, p. 2099) and subcellular and DNA changes in fish that may affect survival (Szarek et al. 2000, p. 439; Cavalcante et al. 2008, p. 4; Langiano and Martinex 2008, p. 229). Temperature, pH, suspended sediment, and other water quality parameters may affect glyphosate and Roundup’s effects on aquatic species. Holiday darters exposed to agricultural chemicals likely experience stress and reduced fitness.

Livestock Access to Streams

Livestock (primarily cattle) is produced in all counties occupied by the holiday darter. In Georgia, the mean number of cattle farms among six counties with holiday darter records is 115 (USDA 2014, p. 464). In Tennessee, the holiday darter occurs in Polk and Bradley counties which have 146 and 527 cattle farms respectively (USDA 2014, p. 371). Within the range of the holiday darter, land that can be grazed for cattle is largely confined to river valleys. As a result, livestock is grazed on pastures adjacent to streams and rivers occupied by holiday darters. In some cases, livestock is allowed free access to the water. Access to streams by cattle has a negative effect on water quality and habitat destruction. Of particular relevance, livestock accessing streams can introduce excess nutrients into streams which degrades water quality, destabilize stream banks which creates increased sediment loads within streams (discussed below), and overgrazing can destroy natural riparian vegetation (discussed below).

Sedimentation

A wide range of current activities and land uses can lead to sedimentation within streams that can include: agricultural practices, construction activities, stormwater runoff, unpaved roads, forestry activities, utility crossings, and mining. Fine sediments are not only input into streams during

23 presently ongoing activities, historical land use practices may have substantially altered hydrological and geological processes such that sediments continue to be input into streams for several decades after those activities cease (Harding et al. 1998, p. 14846).

The negative effects of increased sedimentation are well understood for aquatic species (Newcombe and MacDonald 1991, p.72; Burkhead et al. 1997, p 411; Burkhead and Jelks 2001, p. 964). Sedimentation can affect fish species by degrading physical habitat used for foraging, sheltering and spawning (Burkhead and Jelks 2001, p. 964; Sutherland 2005, p. 90), alter food webs and stream productivity (Schofield et al. 2004, p. 907), force altered behaviors (Sweka and Hartman 2003, p. 346), and even have sub-lethal effects and mortality on individual fish (Sutherland 2005, p. 94; Wenger and Freeman 2007, p. 7). Chronic exposure to sediment has been shown to have negative impacts to fish gills, causing gill damage, stress, and may reduce growth rates (Sutherland and Meyer 2007, p. 401). Holiday darters may experience detrimental effects of sedimentation in the form of gill damage, reduced visibility for feeding and communication, decreased availability of suitable spawning habitats, and reduced spawning success as a result of fine sediments smothering and killing eggs.

Loss of Riparian Vegetation

For this assessment, loss of riparian vegetation refers to removal of natural plant communities from the riparian zone of rivers and streams. Where the holiday darter occurs, reduced or lost riparian vegetation is typically associated with lands used for agriculture or rapidly urbanizing lands. In these cases, the natural riparian plant community is often replaced with lawns, pasture land, or even impervious surfaces.

Removal of riparian vegetation can destabilize stream banks, increase stream sedimentation and turbidity (Barling and Moore 1994, p. 544; Beeson and Doyle 1995, p. 989); reduce the stream’s capacity for trapping and removing contaminants and nutrients from runoff (Barling and Moore 1994, p. 555; Peterjohn and Correll 1984, p. 1473; Osborne and Kovacic 1993, p. 255; Vought et al. 1994, p. 346); increase water temperature (Brazier and Grown 1973, p. 4; Barton et al. 1985, p. 373; Pusey and Arthington 2003, p. 4); and increase light penetration to streams. In turn, this increases algae production (decreasing water quality) (Noel et al. 1986, p. 667; Pusey and Arthington 2003, p. 6); reduces woody debris inputs, removing a source of aquatic habitat (Karr and Schlosser 1978, p. 231); and reduces leaf litter, therefore, decreasing overall stream production (Nakanao et al. 1999, p. 2440; Wallace et al. 1999, p. 429). Aquatic food webs are largely driven by inputs of organic matter from plant and other coarse material that is blown into the stream (Allan 1995, pg. 109), primarily from riparian zones.

Development practices associated with urbanization may remove all vegetation within the riparian zone. However, voluntary best management practices (BMPs) often involve maintaining riparian buffers (a width of land adjacent to a waterbody where vegetation is left in place). Buffers are an essential component of an overall program of stream ecosystem protections,

however, studies that compared open and forested reaches in the Etowah basin along five small streams in suburban catchments (Roy et al. 2005, entire) concluded that riparian buffers – although necessary for protecting fish assemblages –were insufficient alone to maintain healthy assemblages in an urban setting where much stormwater runoff is transported to the stream in pipes, bypassing the buffer. Similarly, agricultural ditches that rapidly drain water from farm fields effectively bypass the filtering provided by a riparian zone with intact vegetation allowing for increased agricultural chemicals and nutrients to enter rivers and streams.

In summary, the holiday darter has adapted to occupy habitats that are surrounded by vegetation, in particular, overstory tree canopy. This riparian vegetation moderates temperature by blocking solar radiation, provides a source for terrestrial plant material that forms the base of the food web and provided shelter and foraging habitat for the holiday darter, and helps to maintain clear, clean water and substrate through filtration. Loss of riparian vegetation is expected to decrease habitat suitability for the holiday darter.

Weather Events

Weather events that affect stream flows are considered to be most relevant to the resiliency of holiday darter populations in this assessment. Broadly, these events include extreme storms and droughts. Increased flows can cause physical washout of eggs and larval fishes, stress on adults (Freeman et al. 2001, p. 187; Power et al. 1996, p. 893), and alter the water quantity and quality and influence primary and secondary production in an stream (Bunn and Arthington 2002, entire), indirectly affecting many fish species. Within the range of the holiday darter, extreme flows associated with hurricanes have been reported to have negative effects on stream fish populations (USFWS 2011, p. 9). Reduced baseflows due to droughts can cause population declines, habitat loss, reduced water quality (decreased dissolved oxygen and temperature alteration) leading to death, crowding of individuals leading to stress, and decreased reproduction in stream fish populations (Mathews and Mathews 2003 , p. 1237). Climate models for the southeastern United States project that average annual temperatures will increase, cold days will become less frequent, the freeze-free season will lengthen by up to a month, temperatures exceeding 95 degrees will increase, heat waves will become longer, and the number of category 5 hurricanes will increase (Ingram et al. 2013, p. 32). While these climate models predict variability into the future, they suggest that the region will be subjected to more frequent large storms (hurricanes) and subsequent storm flows as well as low flows from droughts.

Figure 5. Influence diagram depicting how resource needs of the holiday darter are affected by land use practices and environmental conditions.

Conservation Measures

The holiday darter is recognized by Alabama, Georgia, and Tennessee as a species of concern. It is listed as Priority 1/Highest Conservation Concern by the State of Alabama, Endangered by the State of Georgia, and Threatened by the State of Tennessee. In general, protections accorded to the holiday darter by the States prohibit direct exploitation of the species.

Within the Conasauga River basin, the Natural Resource Conservation Service (NRCS) has begun a Working Lands for Wildlife (WLFW) project that provides technical and financial assistance to help landowners improve water quality and help producers plan and implement a variety of conservation activities, or practices that benefit aquatic species. Holiday darter may benefit from water quality improvements in portions of the Conasauga River that are affected by agricultural practices as a result of the WLFW project.

Priority watersheds within the range of the holiday darter have been designated as Stategic Habitat Units (SHUs) by the Alabama Rivers and Streams Network (ARSN). The SHU project was developed for species restoration and enhancement. Watersheds occupied by holiday darter that have been designated as SHUs are Choccolocco Creek watershed (which includes the Shoal Creek populations) and the Oostanaula River watershed (which includes the Conasauga and Coosawattee River populations).

Some populations of holiday darter are known from watersheds in which a substantial percentage of lands are owned and managed by the U.S. National Forest Service (NFS). These populations are found in the Conasauga River, upper Etowah River, and Shoal Creek. In the Conasauga River and Shoal Creek, the majority of current records for the holiday darter are within the proclamation boundary of NFS lands. Cherokee National Forest in Tennessee, Chattahoochee National Forest in Georgia, and Talladega National Forest in Alabama own and manage natural resources in occupied watersheds in those portions of the holiday darter’s range. Management prescriptions implemented by the NFS in areas that overlap with the range of the holiday darter are expected to benefit the species. Specifically, a 4.5 mile reach of the Conasuaga River is eligible for Congressional Wild River designation and managed to protect and perpetuate the features that led to the eligibility status. The river is also recognized for its aquatic biodiversity by NFS and management strategies employed by both Cherokee and Chattahoochee National Forests within the watershed involve designated wilderness areas, recommended wild river, recommended recreational river, black bear habitat management, restoration and maintenance of rare communities, restoration/management of old growth characteristics, and scenic corridors and sensitive viewsheds among others. These management strategies that emphasize natural forest communities and water quality are expected to benefit holiday darter within the Conasauga River watershed. The Chattahoochee National Forest management prescriptions within the upper Etowah River also broadly emphasize and promote natural plant communities and as such are expected to benefit holiday darter within this watershed. Standards outlined in the Revised Land and Management Plan for National Forests in Alabama (2004) generally protect water and habitat quality in streams. Direct observations of Shoal Creek have found the stream to have good water quality with high levels of dissolved oxygen, stable pH levels, and low sedimentation and confirms the benefits of NFS management strategies to holiday darter habitat.

Within the Amicalola Creek watershed approximately 13.6 miles of the stream are protected by lands owned and managed by the State of Georgia. Among the stated management goals for these lands, maintenance or enhancement of populations of sensitive species and management of riparian areas to benefit water quality, aquatic resources, and aesthetics are expected to benefit holiday darters. Additionally, approximately 488 acres of the aforementioned state owned lands were purchased with the assistance of a Recovery Land Acquisition Grant that prioritized the conservation of aquatic resources and species. Therefore, it is anticipated that State ownership and management within the Amicalola Creek watershed will benefit the long-term survival of holiday darters.

Chapter 5: Current Watershed and Population Conditions

Current habitat and population conditions are described below. This section details specific stressors acting within the occupied watersheds (populations). Additionally, collection history and qualitative abundance is provided. Current population resiliency is assessed for each watershed specifically, followed by a summary of range wide redundancy and representation.

To qualitatively assess current viability we considered five components that broadly relate to either the physical environment (“Habitat Elements”) or characteristics about the population specifically (“Population Elements”). Habitat elements consisted of an assessment of physical habitat, connectivity, water quality, and hydrologic regime. Population elements consisted of an estimation of approximate abundance, the extent of occurrence (total length of occupied streams), and an assessment of occurrence complexity. We further defined how each of these components might vary in terms of qualitative condition (see Table 2). An overall resiliency condition was estimated by combing habitat and population elements. Population elements were weighted 2X higher than habitat elements because they are considered direct indicators of population condition.

Resiliency conditions were classified as “Low”, “Moderate”, or “High”. To help visualize variability that exists within each condition category, we utilized a simple color ramp when displaying overall population resiliency. This color ramp is only intended as a visual aid to highlight populations that vary in resiliency even though the final resiliency condition categories (low, moderate, high) are equivalent.

Low Moderate High

Figure 6. Example color ramp that visualizes variability found within each resiliency condition category.

Methods

Population Elements

To assess population elements, collection records from natural history museums and field notes (when available) were evaluated. Collection records were compiled and provided to the Service by state partners funded under a concurrent Section 6 status assessment for the holiday darter. The dataset used in this analysis is not considered to be exhaustive, but represents the best data accessible in the public domain. Each collection record is georeferenced with geographic coordinates. These records were considered “current” if they represented a collection within the last 10 years (2007 or more recent) and historical if they represented a collection prior to 2007.

Holiday darter occurrence records came from multiple sources and represented a diversity of sampling techniques and methods and therefore, did not exhibit standardization. The number of individuals collected was inconsistently recorded and sampling methods varied among records. Therefore, we did not analyze numbers collected for each record. Instead, abundance was estimated for each record categorically. Two categories were used to assess whether a record represented a “rare” (1-10 individuals) collection or a “common” (10-100 individuals) collection. The percentage of rare and common collections within a population was used to assess population level abundance. In some cases, “present” was the only value associated with collection number. These records were not considered in our qualitative abundance estimations.

Occurrence extent for the holiday darter was evaluated by measuring the distance between the furthest upstream record and the furthest downstream record. Historical and current records were assessed separately to quantify any range reduction that may have occurred.

Occurrence complexity is a measure of the spatial complexity of the occupied habitat. For aquatic species that inhabit rivers, complex spatial occurrence would relate to a species occupying multiple tributaries and the main-stem river as opposed to only inhabiting the main- stem river. If good connectivity is assumed, than the more complex and dendritic (tree-like) spatial arrangement of occupied habitat will be more resilient against extinction (Fagan 2002, p. 3244). We considered high occurrence complexity to exist when individuals occupied the main- stem river in addition to more than three major tributaries. Low occurrence complexity would exist if a species only inhabited the main-stem river and up to two tributaries.

When assessing components of resiliency, we consider population elements to be a more direct indicator of resiliency condition than habitat elements. Therefore, population elements are weighted 2x higher than habitat elements in our resiliency assessment.

Habitat Elements

Population abundance, spatial extent and spatial complexity are results of successful spawning, recruitment (survival of young individuals to maturity and spawning), adult survival, and dispersal. These population dynamics are related to the “needs” of a species which correspond to water quality, water quantity, structural habitat, and connectivity for the holiday darter (discussed in more detail in Chapter 3 of this document). Therefore, it is useful to consider habitat elements when assessing the condition of a population.

Physical (structural) habitat was assessed, in part, with spatial data. Specifically, land cover and use was evaluated within the Active River Area (ARA) of each occupied watershed. The ARA was developed to provide a more meaningful method for assessing riparian zones than a simple buffer. It encompasses the dynamic processes within the aquatic and riparian zones that interact with a lotic system and create the associated habitats and habitat conditions (Smith et al. 2008, pg. 1). The percentage of natural vegetation within the ARA was used as an indicator of within stream habitat quality. Land cover types considered as natural include open water, deciduous

29 forest, mixed forest, shrub, wooded wetland and herbaceous wetland. For this analysis, the evergreen forest land cover type was considered as altered vegetation because this likely represents silviculture as dominate natural plant communities are hardwood and mixed hardwood forests within the range of the holiday darter based on the 2011 GAP/LANDFIRE (USGS GAP Analysis Program 2011). In addition to vegetation types within the ARA, we assessed other land use practices that may affect in stream habitat quality (e.g. cattle access to streams, extent of urban areas outside of ARA, channelization, etc.).

Connectivity was assessed at two scales within this assessment. Connectivity, as it relates to population resiliency, was assessed by evaluating the number and placement of small and large dams and road crossings within populations of holiday darter. Specifically, the presence of barriers that fragment known occupied stream reaches within a population was the metric used to evaluate connectivity as it relates to resiliency.

Connectivity is also important across the range of the species and was considered when assessing redundancy and representation. At this scale, long reaches of unsuitable habitat or barriers located at the lower reaches of occupied watersheds can eliminate connectivity and movement between populations thus affecting redundancy and representation, but not necessarily resiliency.

Barriers located at the lower limits of an occupied watershed that represents a population as well as length of unsuitable habitat were considered

Water quality conditions within populations of holiday darter were determined by assessing known and reported water quality issues from the Environmental Protection Agency’s Clean Water Act Section 303(d) and Total Maximum Daily Loads (TMDLs) program. When available, watershed reports by non-governmental organizations (NGOs) were also used to assess water quality. Additionally, we inspected aerial imagery to assess the prevalence of land use practices that are known to negatively affect water quality (e.g. poultry houses, urban areas, and cattle access to streams). These land use practices are discussed specifically within each population current condition and resiliency assessment.

To assess hydrologic regime conditions within populations of holiday darter, any land use or resource use practice that is known to affect flows and occurred within the occupied watershed was considered. These practices were related to urbanization, agriculture, and reservoir construction. Any of these practices can cause deviations from a more natural flow regime and have negative effects on holiday darter as discussed in Chapter 3.

Table 2. Definitions of conditions for components used to assess current conditions

High Medium Low 0 Physical No known alteration Known low level Habitat heavily Unable to Habitat occurring within alterations to habitat but altered and support Active River Area not known to be recognized as survival (ARA),> 90% natural negatively affecting impacting species, veg species, natural <50% natural vegetation 50-89% in vegetation in ARA ARA

Connectivity No known barriers to Passage barriers known Passage barriers Unable to fish passage within a but do not impact species identified as support population within a population negatively affecting survival populations within a population

Water Quality Minimal or no known Issues recognized (i.e. WQ issues known to Unable to water quality issues 303d streams, unpaved impact populations support roads, moderate housing survival amounts)

Hydrologic Minimal or no known Issues recognized but Flow issues known to Unable to Regime flow issues low intensity (lower impact species support density suburban (hydropeaking, water survival development, lower avail issues below amounts of reservoirs) channelization)

Approximate >75% of collections 75-50% of collections >50% of collections Extirpated Abundance classified as common classified as common classified as rare (1- (10-100 individuals) (10-100 individuals) 10 individuals)

Occurrence Entire known range <30% decline from >30% decline in Extirpated Extent currently occupied known range known range

Occurrence Occupies main Occupies main channel Occupies main Extirpated Complexity channel and and at least 3 tributaries channel and at least 2 numerous tributaries tributaries

Representation

To evaluate representation for the holiday darter, we considered the variability that has been documented within the species (genetic, morphological, and behavioral). The aforementioned variability has been described in Shoal Creek, upper Coosawattee River, Etowah River, Amicalola Creek, and Conasauga River. We consider that geographic variation identified above is irreplaceable to the holiday darter. Therefore, representation is measured by the presence of resilient populations that represent this known variability within the species. Additionally, we measure representation in terms of the species’ ecological setting in which the species has evolved by assessing occupied physiographic provinces. The holiday darter occupies the Blue Ride, Piedmont, and Ridge and Valley provinces. We assert that holiday darter intrinsically exhibits moderate representation as measured by physiographic province. To clarify, it is helpful to consider representation that is lost if extirpation occurs within a population. If the holiday darter is extirpated from one physiographic province, this represents a 33% reduction in variability of ecological setting. Finally, we consider connectivity when assessing representation. Lack of connectivity between the seven populations would prevent the exchange of novel and beneficial adaptations or prevent migrations to more suitable habitat. Therefore, without connectivity, the species’ ability to adapt to a changing environment is lost even if representative populations persist.

Table 3. Definitions of condition categories for representation

High Medium Low Physiographic * All provinces One or two provinces Province Variability occupied with high occupied high resiliency, resiliency, populations populations Genetic, High resiliency Extant populations Variation is lost due Morphological, populations that that represent all to extirpation in at Behavioral represent all known known genetic, least one population Variability genetic, morphological, and that represents known morphological, and behavioral variability variability behavioral variability

Redundancy

Redundancy for the holiday darter is characterized by having multiple, resilient and representative populations distributed throughout its range. Similar to representation, we consider the small range of this endemic species when evaluating redundancy. The holiday darter is effectively known from seven populations (Conasauga River, Talking Rock Creek, Mountaintown Creek, Ellijay River, Amicalola Creek, Etowah River, and Shoal Creek). Therefore, we make the assertion that the holiday darter would not naturally display “High” redundancy because the loss of any one population would represent an approximate 14% reduction in total redundancy. Furthermore, redundancy is necessary to protect against “irreplaceable loss of representation”; therefore, multiple populations are that represent known variation are needed for the holiday darter to exhibit “high” redundancy.

In this assessment, we consider the holiday darter to intrinsically have “Medium” redundancy and that this condition can be maintained if five of the seven populations are found to have a “High” resiliency. We also consider connectivity when evaluating redundancy. Connectivity would allow for dispersal and recolonization events to occur should a population become extirpated within a watershed (rescue effect). Without connectivity the relative importance of localized stochastic events is increased. In other words, as isolated populations become extirpated, the species as a whole is less robust to smaller, more probable, and potentially more frequent stochastic events.

Current Condittion

Conasauga River

Current records (those made since 2007) for the holiday darter exist in the Conasauga River from the southern end of the Alaculsy Valley in Georgia to the mouth of Minnewauga Creek and within downstream reaches of Jacks River. The total currently occupied reach length in the Conasauga River watershed is approximately 10 river miles (~16 river kilometers). This reach length represents 60% of the historical known range of the holiday darter in the Conasauga River watershed. Additionally, 32% of collection records found the holiday darter to be “rare” (less than 10 individuals were collected) while 24% found the holiday darter to be common (10-50 individuals collected). Uncertainty exists when assessing abundance for the holiday darter in the Conasauga River because 44% of collections in our database did not provide numeric data for number of individuals collected. The spatial arrangement of occurrence records is generally a simple linear arrangement, representing low spatial complexity.

Figure 7. Current and historical range of the holiday darter in the Conasauga River watershed.

The majority of current records for this species are upstream of the US 411 bridge in Polk County, TN. Much of the surrounding land in this portion of the watershed is owned by the U.S. Forest Service (USFS) and is noted for having good water quality and suitable habitat, although unpaved forestry roads are a source of increased levels of sedimentation and the Jacks River is on the Georgia 303(d) list because it is known to have elevated fecal coliform counts. Due to a decline in extent of habitat occupied, a simple spatial arrangement of extent occupied, and low qualitative abundance, the holiday darter is considered to be susceptible to stochastic events and have a low resiliency.

Low Moderate High X Figure 8. Final resiliency condition for the Conasauga River population

Talking Rock Creek

Within the Talking Rock Creek watershed, the holiday darter has been collected from an approximately 1.25 river mile (2 km) reach near the GA Hwy 136 crossing. One collection has been made since 2007 where four individuals were collected. The spatial arrangement of occurrence records is linear from the upstream to downstream records, indicating low complexity for this population’s distribution.

Two limestone quarries are present along Talona Creek and have the potential to increase sedimentation and alter water chemistry within Talona Creek and Talking Rock Creek. The Georgia Environmental Protection Department (EPD) has identified portions of Talking Rock Creek and its tributaries to be "non-supporting" for fishing and named negative effects of nonpoint source pollution on fish communities as the source. The nonpoint source pollution potentially originates from low level developments and agricultural fields and pastures. Poultry houses are present in this watershed and pastures are spread with chicken litter likely increasing nutrient and endocrine disruptor concentrations. Upper portions of Talking Rock Creek that flow through pasture land have little or no natural vegetation in the riparian zone and experience bank erosion and increased levels of sedimentation. Sedimentation, nutrient pollution, riparian alteration from forestry and agriculture, and urbanization has been identified to negatively affect water quality (Albanese et al. 2013).

Fish populations in Talking Rock creek are completely isolated from the Coosawattee and the rest of the Upper Coosa River basin by a reregulation dam and reservoir associated with Carters Dam. Water impounded from the reregulation dam inundates lower reaches of Talking Rock Creek and eliminates suitable habitat for the Holiday Darter from the lower reaches of the stream. Small reservoirs upstream of the town of Talking Rock prevent upstream movement.

Two hypotheses exist to explain the apparent rarity of sensitive fish species in Talking Rock Creek (Albanese et al. 2013). Talking Rock Creek begins in the Blue Ridge geographic province and rapidly moves into Ridge and Valley where it has a lower gradient than the other streams in

the region that are occupied by Holiday Darter. The rapid change in geology and elevation suggest that Talking Rock Creek may have historically provided less suitable habitat and behaved as a "sink" population. Alternatively, river and land use alteration may have degraded water quality and habitat and led to the apparent decline of holiday darter and other sensitive fish species (blue shiner and goldline dater) in Talking Rock Creek. Lack of connectivity between this population and others prevents any natural immigration into Talking Rock Creek from occurring and increases the risk of extirpation.

Due to identified water quality issues, the low abundance, extremely localized extent of occurrence records, and lack of connectivity for holiday darters in this system, this population has a very low resiliency to stochastic events and is currently considered at risk of extirpation by GA DNR (Albanese and Abouhamdan 2017, p. 10).

Low Moderate High X Figure 9. Final resiliency condition for the Talking Rock Creek population

Figure 10. Current and historical range of the holiday darter in the Talking Rock Creek watershed.

Ellijay River

Despite an appreciable number of fish surveys having been undertaken in the entire Ellijay River watershed (37 since 2007), no records for Holiday darter are known from the main-stem of the Ellijay River. Records exist for this species historically in tributaries to the Ellijay River including: Cherry Log Creek, Rock Creek, Briar Creek, and Turniptown Creek. Current (2007 – 2017) records for the holiday darter exist at one point in Rock Creek upstream of a small impoundment and within an approximately 0.6 river mile (1 km) stretch of Turniptown Creek between Northcutt Road and GA Hwy 515. The current range of holiday darter within the Ellijay River basin represents 18% of the assumed historical range. Of these three records, two were noted as being rare and no abundance data was collected for the third.

The Town of Ellijay is adjacent to the Ellijay River at its lower reaches and introduces urban stressors to the river. Upstream of the Town of Ellijay, numerous agricultural pastures are present and adjacent to the river. These pastures have limited natural vegetation in the riparian zone and in some cases provide access to the river for cattle, causing erosion and sedimentation. These pastures are likely spread with chicken litter from the numerous chicken farms present in the Ellijay River watershed, leading to an increase in excess nutrients and endocrine disruptors in the water. The Ellijay River is within Gilmer County, which has been noted as producing chickens at the highest density of any county in Georgia (USDA 2014a). Small impoundments exist throughout the watershed. One small impoundment isolates holiday darters in Rock Creek from the rest of the Ellijay River. A small impoundment at the lower reaches of Cherry Log Creek fragments this historically occupied stream from the rest of the Ellijay River.

Due to very few occurrences, small extent of occupied habitat, relatively simple spatial complexity of occupied habitat, fragmentation from small impoundments, and stressors introduced form urbanization and agriculture the holiday darter is expected to have low resiliency to stochastic events and is considered at risk of extirpation by GA DNR (Albanese and Abouhamdan 2017, p. 10).

Low Moderate High X Figure 11. Final resiliency condition for the Ellijay River population

Figure 12. Current and historical range of the holiday darter in the Ellijay River watershed. The dot depicts the Rock Creek collection record and the arrow shows the location of a small reservoir that isolates darters from the rest of this watershed.

Mountaintown Creek

Collections of the holiday darter have been made in Mountaintown and East (Little) Mountaintown creeks over a stream length of approximately 12 river miles (20 km) since 2007. This represents approximately 74% of the holiday darter’s historical range in the Mountaintown Creek watershed. Of these records, 90% found the holiday darter to be “rare” (less than 10 individuals collected). Because occurrences are from Mountaintown Creek and a single tributary, spatial occurrence complexity is considered low.

Similar to the Elijay River, numerous agricultural pastures are present and adjacent to the river. These pastures have little or no natural vegetation in the riparian zone and in some cases provide access to the river for cattle, causing erosion and sedimentation. These pastures are likely spread with chicken litter from the numerous chicken farms present in the Mountaintown Creek watershed, leading to an increase in excess nutrients and endocrine disruptors. Mountaintown Creek is within Gilmer County, which has been noted as producing chickens at the highest density of any county in Georgia (USDA 2012). Additional stressors are introduced into lower reaches of the Mountaintown Creek watershed through increased levels of low density development associated with development.

Due to the small and simple spatial extent of occupied habitat, low qualitative abundance, and stressors particularly form agricultural practices, the holiday darter in the Mountaintown Creek watershed is expected to have a low resiliency to stochastic events.

Low Moderate High X Figure 13. Final resiliency condition for the Mountaintown Creek population.

Figure 14. Current and historical range of the holiday darter in the Mountaintown Creek watershed.

Amicalola Creek

Collections of the holiday darter since 2007 occur in 28 river miles (45 km) within the watershed which represents approximately 80% of its historical range. Because the holiday darter is known to occur in Amicalola Creek, Little Amicalola Creek, Cochran Creek, and Gab Creek, it has a moderate spatial occurrence complexity. Uncertainty exists regarding abundance of holiday darters within this watershed. Collections that found the species to be “rare” (less than 10 individuals) made up 43% of the total records, “common” (10-100 individuals) collections made 37% of the total records, and 20% of records did not provide numerical count data.

Many of the same stressors that are present in other North Georgia streams exist in the Amicalola watershed. Some major tributaries have been channelized during past agricultural practices and experienced a reduced amount of suitable habitat and decreased stream residence times (increased flashiness) as a result. Chicken farms are present in this watershed and some pasturelands adjacent to occupied streams receive chicken litter from poultry farming practices. Small impoundments at the headwaters of occupied streams prevent upstream movement of holiday darter. Approximately 13.6 miles (22 km) of the stream are protected by lands owned and managed by the State of Georgia. Among the stated management goals for these lands, maintenance or enhancement of populations of sensitive species and management of riparian areas to benefit water quality, aquatic resources, and aesthetics are expected to benefit holiday darters.

Conservation actions, as well as relatively intact forest cover through-out the watershed and along the active river area (~74% natural vegetation) have led to overall good water quality and high amounts of suitable habitat. These attributes of the watershed in addition to a moderate spatial complexity are expected to provide for a moderate resiliency to stochastic events despite uncertainty regarding abundance.

Low Moderate High X Figure 15. Final resiliency condition for the Amicalola Creek population

Figure 16. Current and historical range of the holiday darter in the Amicalola Creek watershed.

Etowah River

Within the Etowah River basin, the holiday darter has been collected from an upstream-most site in the Etowah River at Hightower Church Rd (FS 28-1), near the mouth of Ward Creek, downstream to about 0.9 river miles (~1.4 km) downstream of the mouth of Castleberry Bridge Road on the Etowah River. Current records exist for the species from the Etowah River at the upper Hightower Church Road (FS 28-1) crossing to the Etowah River at Castleberry Bridge Road, a distance of about 21 river miles (~33 km) which represents approximately 49% of its historical range in the watershed. This decline in occurrence within the Etowah River is primarily due to a lack of current records in Nimblewill Creek. Because current records are only known from the Etowah River, this population exhibits low spatial complexity. Holiday darters have been collected 37 times in the last ten years in the Etowah River and 90% of these records found the species to be rare (<10 individuals per collection).

The upper portions of the watershed are within Chattahoochee National Forest and wildlife management areas managed by Georgia Department of Natural Resources. Additionally, this part of the Etowah River basin is largely forest cover (~75% natural vegetation in the active river area). Low intensity developments in some parts of the watershed can negatively affect water quality from leaky septic systems and storm water. As with other parts of North Georgia, chicken farms are present and spreading occurs on pastures adjacent to the Etowah River. However, there are fewer open agricultural lands here than in other watersheds occupied by holiday darter. Sedimentation from non-paved roads is present. In general, this watershed has good water quality, adequate flows, and suitable habitat. However the Etowah River population is expected to have a low resiliency to stochastic events because, although water quality and habitat is generally good, the abundance is qualitatively low, the spatial complexity of the occupied habitat is low, and this population has experienced a substantial reduction in extent of occurrence.

Low Moderate High X Figure 17. Final resiliency condition for the Etowah River population

Figure 18. Current and historical range of the holiday darter in the Etowah River watershed.

Shoal Creek

The nominative population of Holiday darter is the only known population within Alabama. Recent studies of Holiday darter within Shoal Creek delineated its range between Whiteside Mill Lake and Sweetwater Lake in the Talladega National Forest (Meade 2009, p. 15). High Rock Lake effectively divides the Shoal Creek population into three isolated subpopulations. Records for these isolated subpopulations consist of approximately 9.3 river miles (15 km) of Shoal Creek and approximately 0.19 river miles (300 meters) of Little Shoal Creek. The current range of holiday darter within Shoal Creek represents 57% of its historically known range within Shoal Creek. Of the current records for the holiday darter in Shoal Creek, 80% considered the species rare (less than 10 individuals collected).

Shoal Creek is a tributary to Choccolocco Creek. Historically, spring tributaries in upper Choccolocco Creek and downstream of the mouth of Shoal Creek were occupied by Holiday darters but have not been collected since the early 1990s. This distributional pattern suggests that the species historically occupied portions of Choccolocco Creek and Shoal Creek watersheds but the range has since contracted to only Shoal Creek.

Impoundments are the primary stressor in this watershed. Whitesides Mill Lake, Highrock Lake, and Sweetwater Lake eliminated the holiday darter from portions of this watershed and fragmented occupied reaches. These impoundments alter flows and have been found to decrease suitable habitat and increase levels of sedimentation (Mead 2009, p. 10). Current records for the holiday darter are primarily within land owned and managed by the U.S. Forest Service. Habitat and water quality were primarily found to be in good condition within the National Forest likely as benefit of high amounts of natural vegetation present in the riparian zone (Mead 2009 pg. 10).

While good habitat and water quality exists within Shoal Creek, this population of holiday darter is expected to have a low resiliency to stochastic events due to the small and simple extent of occurrence, low qualitative abundance, and the lack of connectivity within the watershed.

Low Moderate High X Figure 19. Final resiliency condition for the Shoal Creek population.

Figure 20. Current and historical range of the holiday darter in the Shoal Creek watershed in Alabama. Arrows show the location of three small reservoirs that contain and fragment this population, from left to right: Whiteside Mill Lake, Highrock Lake, and Sweetwater Lake.

Table 4. Estimated resilience factors and overall resiliency condition for holiday darter populations

Approximate Occurrence Occurrence Physical Connectivity Water Hydrologic Overall Abundance Extent Complexity Habitat (within Quality Regime Condition population) Conasuaga Low Low Low Moderate High Moderate Moderate Low River (25 current records) Talking Rock Low Low Low Moderate High Low Moderate Low Creek (1 current record) Elijay River Low Low Low Moderate Moderate Low Low Low (3 current records) Mountaintown Low Low Low Moderate Moderate Moderate Moderate Low Creek (10 current records) Amicalola Moderate Moderate Low Moderate Moderate Moderate Moderate Moderate Creek (70 current records) Etowah River Low Low Low Moderate High Moderate High Low (50 current records) Shoal Creek Low Low Low Moderate Low High Moderate Low (14 current records)

Current Species Representation

Currently, all historically occupied physiographic provinces are occupied, albeit primarily with populations that exhibit low resiliency, as described under the current condition assessment above. Populations that exhibit low resiliency are susceptible to stochastic events and have a higher risk of extirpation than populations with high resiliency; therefore, populations with low resiliency have a lower relative importance to representation than populations with high resiliency. Because our assessment considers six populations of holiday darter to have a low resiliency and one population to have high resiliency, we considered physiographic province representation to be low. Similarly, although populations that exhibit the known genetic, morphological, and behavioral variability are currently extant, they do not exhibit high resiliency. Therefore, genetic, morphological, and behavioral representation is considered low.

Connectivity is an important aspect of representation because it provides for the exchange of novel and beneficial adaptations and migration to more suitable habitat (should it be necessary). Connectivity is reduced for the species, range-wide. Dams have completely isolated the seven populations into four groups. The upper Etowah River-Amicalola Creek populations are isolated by Alatoona Dam, the Talking Rock Creek population is isolated by Carters Re-regulation dam and the Elijay River and Mountaintown Creek populations are isolated by Carters Dam. The Conasauga River population is prevented from dispersing in to the other populations by all dams mentioned. The Shoal Creek population is isolated by large dams on the Coosa River. Where dams do not fragment habitat, long reaches of unoccupied habitat are present between populations; indicating that migration between populations is uncommon or unlikely.

Finally, all populations of holiday darter exist on the periphery of the Coosa River basin and have likely reached the upstream limits for the species. It is unlikely that individuals within a population will be able to migrate further upstream if necessitated by changes in environmental conditions, further decreasing the ability of the species to adapt to changing environmental conditions.

We estimate that the holiday darter currently has low adaptive potential due to limited representation in six occupied watersheds, lack of connectivity, and confinement to upper reaches of occupied watershed. Overall representation for the holiday darter is low.

Current Species Redundancy

Redundancy describes the ability of a species to withstand catastrophic events. It “guards against irreplaceable loss of representation” (Redford et al. 2011 p. 42; Tear et al. 2005 p. 841)

and minimizes the effect of localized extirpation on the range-wide persistence of a species (Shaffer and Stein, p. 308)

Redundancy for the holiday darter is characterized by having multiple, resilient and representative populations distributed throughout its range. Because all but one populations of holiday darter exhibit low resiliency, as determined under our current resiliency assessment, the species is considered to also have low redundancy. The current spatial extent of each population is smaller than the historically known spatial extent; therefore, we infer that all populations have experienced declines in amount of occupied habitat. Additionally, all populations have qualitatively low numbers with most records consisting of ten or fewer individuals observed as well as low spatial complexity as seen by the simple linear arrangement of populations. Furthermore, redundancy is only present within the Coosawattee River, with two populations that are at risk of extirpation and one low resiliency population. Therefore, a key component of redundancy – protection from irreplaceable loss of representation – is not met.

Connectivity is also reduced for the species, range-wide. Dams have completely isolated the seven populations into four groups. The upper Etowah River-Amicalola Creek populations are isolated by Alatoona Dam, the Talking Rock Creek population is isolated by Carters Re- regulation dam and the Elijay River and Mountaintown Creek populations are isolated by Carters Dam. The Conasauga River populations is prevented from dispersing in to the other populations by those same dams. The Shoal Creek population is isolated by large dams on the Coosa River in addition to Whiteside Mill Dam on Shoal Creek. Where dams do not fragment habitat, long reaches of unoccupied habitat are present between populations; indicating that migration between populations is uncommon or unlikely. As mentioned earlier, the overall lack of connectivity between populations increases the effects of localized stochastic events and the species as a whole is less robust to smaller, more probable, and potentially more frequent stochastic events. Therefore, a key component of resiliency – the minimization of the effect of localized extirpation – is not exhibited in this species.

Chapter 6: Future Viability

In this chapter, we describe how current viability of the holiday darter may change over a period of 50 years. Like current conditions, we evaluate species viability in terms of resiliency at the population scale, and representation and redundancy at the species scale (3Rs). Here we describe three plausible future scenarios and whether there will be a change, from current conditions, to any of the 3Rs under each scenario. Our future scenarios differ by considering variations that are predicted in two main elements of change, urbanization and climate. These scenarios capture the range of likely viability outcomes that the holiday darter will exhibit by the end of 2070.

The human population in the southeastern United States has grown at an average rate of 36.7% since 2000, making it the fastest growing region in the country (U.S. Census 2016, p. 1-4). The holiday darter has been exposed to this reality because it inhabits small rivers of the upper Coosa River basin that are in close proximity to Atlanta, one of the largest metropolitan areas in the Southeast. As a result, urbanization has consistently been identified as a major stressor that affects streams within the range of the holiday darter (Freeman et al. 2002, entire; Wenger and Freeman 2007, entire). Growth will continue at a rapid pace within Atlanta and the surrounding areas. Therefore, development and urban sprawl is expected to expand and influence areas that previously were unaffected by urbanization. Rapid growth in the Atlanta area and across the southeastern U.S. as a whole is expected to be a major driver of change and an important consideration when evaluating future viability of the holiday darter. In this section, we consider how land use across the holiday darter’s range is predicted to change and develop. Further, we assess how this increase in developed areas affects populations and the species as a whole.

Methods

To forecast future urbanization, we consider future scenarios that incorporate the SLEUTH (Slope, Land use, Excluded area, Urban area, Transportation, Hillside area) model. This model simulates patterns of urban expansion that are consistent with spatial observations of past urban growth and transportation networks, including the sprawling, fragmented, “leapfrog” development that has been dominant in the southeastern U.S. (Terando et al. 2014, p. 2). The extent of urbanized areas has been predicted to increase across the southeastern U.S. by approximately 100 – 192 % based on the “business-as-usual” (BAU) scenario that expects future development to match current development rates (Terando et al. 2014, p. 1). We use this range of percent change in urbanization to develop our future scenarios.

The Fifth Assessment Report (AR5) by the Intergovernmental Panel on Climate Change (IPCC) found that “continued emission of greenhouse gases will cause further warming and long-lasting changes in all components of the climate system, increasing the likelihood of severe, pervasive and irreversible impacts for people and ecosystems” (IPCC 2013, p. 8). Therefore, we expect climate change to be a driver of change that should be addressed when evaluating future viability of the holiday darter.

The IPCC utilized a suite of alternative scenarios in the AR5 to make near-term and long-term climate projections. These scenarios, called “representative concentration pathways” (RCPs) are plausible pathways toward reaching a target radiative forcing (the change in energy in the atmosphere due to greenhouse gases) by the year 2100 (Moss et al. 2010, p. 752). RCPs help scientists capture the most plausible range of outcomes for climate futures based on uncertainties inherent in the natural and socio-economic environment. In this assessment, we used RCPs to help understand how climate may change in the future and what effects may be observed that impact the holiday darter in the upper Coosa River basin.

There are four RCPs (Table 2) that have been utilized by the IPCC. These RCPs correspond to a high radiative forcing trajectory pathway (RCP8.5), a low radiative forcing trajectory (RC2.6), and two intermediate radiative forcing trajectories (RCP6.0 and RCP4.5). We used these RCPs as the basis for developing future scenarios with low, moderate, and high probabilities of extreme weather events as a result of low levels of climate change (RCP2.6), moderate level of climate change (RCP6.0-RCP4.5), and extremely altered climate conditions (RCP8.5), respectively.

Table 5. description of the four RCPs from Moss et al. 2010 pg. 753, please consult Moss et al. 2010 and Collins et al. 2013 for detailed descriptions of future climate scenarios.

Name Radiative forcing Concentration (ppm) Pathway -2 RCP8.5 >8.5 W m in 2100 >1,370 CO2-equiv in 2100 Rising without stabilization

-2 RCP6.0 ~6 W m at ~850 CO2-equiv (at Stabilization without stabilization after 2100 stabilization after 2100) overshoot

-2 RCP4.5 ~4.5 W m at ~650 CO2-equiv (at Stabilization without stabilization after 2100 stabilization after 2100) overshoot

-2 RCP2.6 Peak at ~3 W m Peak at ~490 CO2-equiv Peak and decline before 2100 and then before 2100 and then declines declines

In the Southeast through the 21st century, climate models project that average annual temperatures will increase, cold days will become less frequent, the freeze-free season will lengthen by up to a month, days with temperatures exceeding 95 F will increase, heat waves will become longer, sea levels will rise an average of 3 feet, the number of category 5 hurricanes will increase, and air quality will decline (Ingram et al. 2013, p. 32). Aquatic systems will be negatively affected by increasing water temperatures, decreasing dissolved oxygen levels, altered streamflow patterns, increased demand for water storage and conveyance structures, and increasing toxicity of pollutants (Ficke 2007, p. 585, 586, 589; Rahel and Olden 2008, p. 522 and

526). Reduced spring/summer rainfall, coupled with increased evapotranspiration and water demand, could lead to local extirpations if streams dry out more frequently. Regardless of RCP and scenario evaluated, we anticipate some or all of events to occur. We attempt to capture the range of plausible climate outcomes by considering that the frequency and probability of extreme climate scenarios will be greatest under RCP8.5, minimal under RCP2.6, and intermediate under RCP6.0 and RCP4.5. For instance, we assume there will be a greater likelihood of a category 5 hurricane negatively impacting the holiday darter under RCP8.5 than under RCP2.6, RCP4.5, or RCP6.0.

Status Quo

In the Status Quo Scenario, current environmental regulations and policy, land use management techniques, and conservations measures remain the same over the next 50 years. We anticipate the current trend in greenhouse gas emissions to continue and moderate impacts from extreme weather events including intense drought, floods, and storm events to occur. Rapid urbanization will continue at the current estimated rate for the piedmont region of the southeastern U.S. (~165 % growth, Terando et al. pg 5) which will increase demand for water resources and introduce multiple additional stressors into local streams and rivers. Despite an overall growth in population and increases in developed areas, some regions will remain predominantly in agriculture and experience associated water quality declines. In pace with current trends we anticipate declines in habitat and water quantity and quality as a result of rapid urbanization, climate change, agricultural practices, and an overall lack of voluntary conservation measures being implemented.

Conasauga River

According to the SLEUTH model, urbanization is expected to increase slightly within the upper Conasauga River but substantially more around Dalton, Georgia. Development will begin to merge the cities of Dalton and Chatsworth, Georgia. Areas that are not developed will remain in agriculture. Under the Status Quo scenario, agricultural practices are expected to remain consistent with those of today. Therefore, we expect continued inputs of pollutants from herbicide applications, excess nutrients and endocrine disruptors from chicken farming and chicken litter spreading on pastures and fields adjacent to the Conasauga River, and increases in sedimentation. Due to water quality and physical habitat that has further degraded from current levels in the Conasauga River adjacent to private lands, we expect the holiday darter to be extirpated from lower reaches of the Conasauga River but persist in upper reaches owned and managed by the U.S. Forest Service. This substantial loss of occupied range and exposure to slightly more extreme climate events anticipated in this scnenario will lead holiday darter to have low resiliency in the Conasauga River by the end of 2070.

Low Moderate High X Figure 21. Final resiliency condition for the Conasauga River population

Figure 22. Predicted urban growth based on the SLEUTH model in the Conasauga River watershed.

Talking Rock Creek

Given the currently very low abundance of the population and extremely small range extent in Talking Rock Creek, further habitat degradation from urbanization and moderate increases in extreme climatic events that are expected to occur under the Status Quo scenario will likely cause extirpation of this already at risk population of holiday darter under this scenario.

Figure 23. Predicted urban growth based on the SLEUTH model in the Talking Rock Creek watershed.

Elijay River

Elijay River will see substantially more development around the town of Elijay, Georgia. The added pressures of urbanization will continue to degrade water quality and alter flows in this watershed. Agricultural lands that remain will add to degraded water quality. It is unlikely that this currently small population with low abundances will be able to persist in the future under the Status Quo scenario, this population will likely be extirpated.

Figure 24. Predicted urban growth based on the SLEUTH model in the Ellijay River watershed.

Mountaintown Creek

According to the SLEUTH model, urbanization is expected to increase in the downstream portions of the Mountaintown Creek watershed. The resulting degradation of habitat and water quality will likely cause the holiday darter to be extirpated from these reaches. Upstream of developed areas current agricultural practices are expected to continue under the Status Quo scenario. Runoff containing chicken litter will continue to enter into the stream and a lack of natural vegetation in the riparian zone will further degrade habitat and water quality. Small impoundments upstream of occupied stream reaches will limit the ability of holiday darters to move upstream and seek refugia from altered environments downstream. Because of the currently low abundance within this population and the small extent of occurrence, it is not likely that this population will persist with increasing urbanization and continual input of agricultural pollution. Under the Status Quo scenario, this population will likely be extirpated.

Figure 25. Predicted urban growth based on the SLEUTH model in the Mountaintown Creek watershed.

Amicalola Creek

Development is expected to increase within the watershed primarily along Cochran Creek according to the SLEUTH model. Increases in development will reduce water and habitat quality. The effects of urbanization here will likely cause range reduction in this population. Agricultural practices are likely to contribute to reductions in water and habitat quality. Because much of the Amicalola Creek corridor is owned and managed by the State of Georgia some effects from both urbanization and agriculture are anticipated to be minimized, allowing holiday darter to persist within this watershed. Therefore, under the status quo scenario, this population is expected to persist and maintain a low resiliency to stochastic events due to a reduced distribution and spatially simple extent of occurrence.

Low Moderate High X Figure 27. Final resiliency condition for the Amicalola Creek population

Figure 26. Predicted urban growth based on the SLEUTH model in the Amicalola Creek watershed.

Etowah River

Urbanization is expected to expand eastward from Dawson and westward from Dahlonega, Georgia towards the upper Etowah River. Northward expansion from Cumming, Georgia along Hwy 19 and Hwy 53 is also expected. In all, this watershed is anticipated to be 20% developed by 2070. As a result from increased levels of urbanization in the status quo scenario, portions to the Etowah River are expected to exhibit characteristics of the urban stream syndrome (flashiness and degraded water quality). Under the Status Quo scenario, we anticipate the range of holiday darter within the Etowah River to contract further into upstream reaches that are managed by the U.S. Forest Service. Therefore, as a result of declines in spatial extent this population is expected to have low resiliency to stochastic events.

Low Moderate High X Figure 29. Final resiliency condition for the Etowah River population

Figure 28. Predicted urban growth based on the SLEUTH model in the Etowah River watershed.

Shoal Creek

Because Shoal Creek is primarily contained within U.S. Forest Service lands, minimal alteration is expected due to urbanization. Under the Status Quo scenario, isolation caused by Highrock Lake will likely cause genetic drift or genetic bottle necks to occur. Additionally, these small, fragmented subpopulations will be susceptible to extreme weather events anticipated in the Status Quo scenario as well as other stochastic events. The holiday darter is expected to persist at low resiliency and at risk of extirpation under the Status Quo scenario.

Low Moderate High X Figure 31. Final resiliency condition for the Shoal Creek population

Figure 30. Predicted urban growth based on the SLEUTH model near the Shoal Creek watershed.

Table 6. Estimated resilience factors and overall resiliency condition for holiday darter populations under the Status Quo scenario

Approximate Occurrence Occurrence Physical Connectivity Water Hydrologic Overall Abundance Extent Complexity Habitat Quality Regime Condition Conasuaga Low Low Low Moderate High Moderate Moderate Low River Talking Rock 0 0 0 Moderate High Low Moderate Likely Creek Extirpated Elijay River 0 0 0 Low Low Low Low Likely Extirpated Mountaintown 0 0 0 Low Low Low Low Likely Creek Extirpated Amicalola Low Low Low Moderate Moderate Moderate Moderate Low Creek Etowah River Low Low Low Moderate High Moderate High Low

Shoal Creek Low Low Low Moderate Low High Moderate Low

Representation

Representation is expected to decline under the Status Quo scenario. Total known morphological variability is expected to be reduced by approximately 20% with the loss of all populations within the Coosawattee River. Physiographic province variability will be exhibited by holiday darters occupying the Blue Ridge in Tennessee and Georgia and the Ridge and Valley in Alabama. Due to degraded habitat and water quality in downstream reaches of occupied rivers, connectivity is unlikely to be restored among populations and will continue to limit the exchange of novel and beneficial adaptations and migration to more suitable habitat. This limited representation will reduce the adaptive potential of the holiday darter by 2070.

Redundancy

Redundancy for the holiday darter is characterized by having multiple, resilient and representative populations distributed throughout its range. Under the Status Quo scenario, overall redundancy will be reduced by 43% due to the extirpation of three populations. All remaining extant populations are expected to have low resiliency. Under the Status Quo scenario, the holiday darter is expected to have low redundancy. Extant population have experienced declines in extent of occupied habitat, lowered abundance, and lowered spatial complexity.

Connectivity is not expected to improve for the species, range-wide under the Status Quo scenario. Dams have completely isolated the six populations into three groups. As mentioned earlier, the overall lack of connectivity between populations increases the significance of localized stochastic events and the species as a whole is less robust to smaller, more probable, and potentially more frequent stochastic events. Therefore, a key component of resiliency – minimization of the effect of localized extirpation– is not met.

Best Case

In the Best Case scenario we predict wider adoption of conservation measures and policies which involves watershed scale conservation plans (WLFW and watershed HCPs) and enacting a water policy for Alabama. Rapid urban growth remains expected, albeit, at slower rate than under the status quo and worst case scenarios (~100%, Terando et al. pg. 1). Under the Best Case scenario rapidly growing urban areas address environmental concerns and implement water conservation

measures and green infrastructure. These actions would lessen the demand on water resources (requiring fewer drinking water supply reservoirs) and minimize urban effects on streams. While large numbers of roads will still be constructed, under the Best Case scenario, road crossings will be constructed that allow for fish passage. In this scenario we expect carbon emissions to peak before 2020 (van Vuuren et al. 2007, p. 132) resulting in a lower probability of extreme weather conditions negatively effecting stream fishes. As a result of a diverse array of conservation actions being undertaken across the landscape, we would anticipate the species to persist or experience a slightly positive response.

Conasauga River

Under the Best Case scenario development proceeds around Dalton, Georgia at a slower rate than under the Status Quo scenario, leading to about 7.6% of the watershed becoming developed. However, urban planners incorporate green infrastructure and manage stormwater. These actions reduce the negative effects development typically has on streams. Much of the river upstream of Dalton, is maintained for agriculture. Conservation programs (Working Lands for Wildlife) have been successfully implemented and because of wide adoption of best management practices (BMPs) by farmers, pollution from herbicide application and poultry litter are reduced. These actions lead to improved water quality in the Conasauga River. These actions increase the potential for the holiday darter to persist in the Conasauga River outside of U.S. Forest Service lands. Range expansion, if it occurs, will be limited due to the legacy effects from current practices. Under the Best Case Scenario, Conasauga River will have a moderate resilience to stochastic events because its range will still be small and simple (confined to the main stem of the Conasauga River).

Low Moderate High X Figure 31. Final resiliency condition for the Conasauga River population

Talking Rock Creek

Under the Best Case scenario, urbanization that is likely to occur in the Talking Rock Creek watershed will have reduced negative effects to the stream and improvement in agricultural practices will likely create an improvement in the water quality from current conditions. However, under this scenario translocation of fish into Talking Rock Creek is not assumed. Given the currently very low abundance of the population and extremely small range extent in Talking Rock Creek, conservation actions as describe under the Best Case scenario will not likely benefit the holiday darter. Under the Best Case scenario, the holiday darter is expected to become extirpated from Talking Rock Creek.

Elijay River

The Elijay River watershed will see substantially more development around the town of Elijay, Georgia. Even with conservation measures being implemented, the added pressures of urbanization will negatively affect water quality in this watershed to the detriment of the holiday darter population. Due to the very low qualitative abundance, few records, and the small, simple extent of occurrence this population is not expected to persist under the Best Case scenario and will likely be extirpated by 2070.

Mountaintown Creek

According to the SLEUTH model, urbanization is expected to increase in the downstream portions of the Mountaintown Creek watershed. Under the Best Case scenario, conservation measures anticipated to be implemented will likely decrease the negative effects associated with this increase in development. These conservation measures, however, are not expected to improve habitat and water quality from currently observed conditions. Therefore, the holiday darter is expected to persist in Mountaintown Creek and maintain a low resiliency to stochastic events.

Low Moderate High X Figure 36. Final resiliency condition for the Mountaintown Creek population

Amicalola Creek

Urbanization is expected to increase within the watershed; however, its effects will be reduced from conservation measures, green infrastructure, and slower rates of development. Reduced effects of stressors from urbanization and because adjacent lands to Amicalola Creek are owned and managed as a WMA by the State of Georgia will allow this population to persist in all known locations currently occupied. Therefore, it is expected have a moderate resiliency to stochastic events due to an increase in spatial extent occupied.

Low Moderate High X Figure 35. Final resiliency condition for the Amicalola Creek population

Etowah River

Under the Best Case scenario, urbanization is expected to increase by 100% in this watershed. While conservation measures will reduce negative effects to the Etowah River, this rapidly developing area will still see some impacts from increased levels of development and will likely negatively affect holiday dater abundance. Under this scenario, we do not anticipate appreciable declines nor do we expect to see expansion to the range of holiday darter within the Etowah

River. Therefore, it is expected this population will maintain a low resiliency to stochastic events.

Low Moderate High X Figure 36. Final resiliency condition for the Etowah River population

Shoal Creek

Because Shoal Creek is primarily contained within U.S. Forest Service lands, minimal alteration is expected due to urbanization. Under the Best Case scenario, isolation caused by Highrock Lake will remain and likely cause genetic drift or genetic bottle necks to occur. Because extreme weather events are less likely to occur in the scenario, the holiday darter is expected to persist in the Shoal Creek system, and retain a low resiliency to stochastic events.

Low Moderate High X Figure 36. Final resiliency condition for the Etowah River population

Table 7. Estimated resilience factors and overall resiliency condition for holiday darter populations under the Best Case scenario

Approximate Occurrence Occurrence Physical Connectivity Water Hydrologic Overall Abundance Extent Complexity Habitat Quality Regime Condition Conasuaga Moderate Moderate Low Moderate High Moderate Moderate Moderate River Talking Rock 0 0 0 Moderate High Low Moderate Likely Creek Extirpated Elijay River 0 0 0 Low Low Low Low Likely Extirpated Mountaintown Low Low Low Low Low Low Low Low Creek Amicalola Moderate Moderate Moderate Moderate Moderate Moderate Moderate Moderate Creek Etowah River Moderate Low Low Moderate High Moderate High Low

Shoal Creek Low Low Low Moderate Low High Moderate Low

Representation

Representation is not expected to change from current conditions under the Best Case scenario. Total known morphological variability is expected to be maintained even with the loss of two populations within the Coosawattee River. Physiographic province variability will be exhibited by holiday darters occupying the Blue Ridge and Ridge and Valley in Tennessee and Georgia, the Ridge and Valley in Alabama, and some portions of the Piedmont in Georgia. Due to degraded habitat and water quality in downstream reaches of occupied rivers, connectivity is unlikely to be restored among populations and will continue to limit the exchange of novel and beneficial adaptations and migration to more suitable habitat. This limited representation will further reduce the adaptive potential of the holiday darter by 2070.

We estimate that the holiday darter will continue to have low adaptive potential due to limited representation in five occupied watersheds, lack of connectivity, and confinement to upper reaches of occupied watershed. Overall representation for the holiday darter is low.

Redundancy

Redundancy for the holiday darter is characterized by having multiple, resilient and representative populations distributed throughout its range. Under the Best Case scenario, overall redundancy will be reduced by 28% due to the extirpation of two populations. Three remaining extant populations are expected to have low resiliency and two are expected to have moderate resiliency. Under the Best Case scenario, the holiday darter is expected to have low redundancy. Most extant populations will have experienced declines, low numbers, and/or have low spatial complexity.

Worst Case

In the Worst Case scenario we anticipate negative effects to aquatic ecosystems as a result of rapid urbanization (~250% increase in urban areas; Terando 2013, pg. 5). In conjunction with rapid urban growth, we predict that there will be a general lack of conservation measures and policies being implemented at the local, regional, or national levels. Water demand will increase with population and new reservoir construction will take place. In addition to rapid urbanization, carbon emissions are projected to continue to increase above the current levels in this scenario, resulting in a higher probability of extreme weather events that can negatively affect fish species. In areas that remain predominately used for agricultural, there will be an increased amount of herbicide and poultry litter spreading and no protective measures implemented to address water quality issues. Under this scenario, we anticipate a general decline in available suitable habitat, population size, and abundance.

Conasauga River

Under the Worst Case scenario, increased rates of pollutants, nutrients, and endocrine disruptors originating from agricultural practices will be observed within the river. This increase in the rate of pollutant input into the river will result in their higher concentrations than observed under the Status Quo scenario. Rapid development in downstream reaches will further degrade habitat and water quality. Due to degraded water quality in the Conasauga River adjacent to private lands, we expect the holiday darter to be extirpated from these reaches. The holiday darter will likely persist in upper reaches owned and managed by the U.S. Forest Service. This small and simple range will be exposed to extreme climate events that are anticipated to increase under the Worst Case scenario. In general, due to a loss of historical occupied range and increased likelihood for extreme climate events, we expect to it to have low resiliency in the Conasauga River by the end of 2070.

Low Moderate High X Figure 37. Final resiliency condition for the Conasauga River population

Talking Rock Creek

Given the currently very low abundance of the population and extremely small range extent in Talking Rock Creek, further habitat degradation from urbanization and increases in extreme climatic events that are expected to occur under the Worst Case scenario will likely cause extirpation of this already at risk population of holiday darter under this scenario.

Elijay River

Elijay River will see substantially more development around the town of Elijay, Georgia. The added pressures of urbanization will continue to degrade water quality and alter flows in this

watershed. Agricultural lands that remain will add to degraded water quality. It is unlikely that this currently small population with low abundances will be able to persist in the future under the Worst Case scenario, this population will likely be extirpated

Mountaintown Creek

According to the SLEUTH model, urbanization is expected to increase in the downstream portions of the Mountaintown Creek watershed. The resulting degradation of habitat and water quality will likely cause the holiday darter to be extirpated from these reaches. Upstream of developed areas current agricultural practices are expected to continue under the Worst Case scenario. Continued input of chicken litter into the stream and a lack of natural vegetation in the riparian zone will further degrade habitat and water quality. Small impoundments upstream of occupied stream reaches will limit the ability of holiday darters to move upstream and seek refuge from altered environments downstream. Because of the currently low abundance within this population and the small extent of occurrence, it is not likely that this population will persist with increasing urbanization and continual input of agricultural pollution. Under the Worst Case scenario, this population will likely be extirpated.

Amicalola Creek

Development is expected to increase within the watershed primarily along Cochran Creek according to the SLEUTH model. Increases in development will reduce water and habitat quality. The effects of urbanization here will likely cause range reduction and simplification of the occupied range in this population. Agricultural practices are likely to contribute to reductions in water and habitat quality. Because much of the Amicalola Creek corridor is owned and managed by the State of Georgia effects from both urbanization and agriculture are anticipated to be minimized. Therefore, under the Worst Case scenario, this population is expected to persist but maintain a low resiliency to stochastic events due to the small and spatially simple range of occurrence.

Low Moderate High X Figure 38. Final resiliency condition for the Amicalola Creek population

Etowah River

Due to rapid urbanization outward from Atlanta the Etowah River is anticipated to see declines in water quality. Like the Status Quo scenario, we anticipate the range of holiday darter within the Etowah River to contract into upstream reaches that are managed by the U.S. Forest Service. As a result of declines in spatial extent, this population will be susceptible to extreme climate events that are anticipated in this scenario. Under the Worst Case scenario, the holiday darter is expected to persist but exhibit low resiliency within the Etowah River.

Low Moderate High X Figure 38. Final resiliency condition for the Amicalola Creek population

Shoal Creek

Because Shoal Creek is primarily contained within U.S. Forest Service lands, minimal alteration is expected due to urbanization. Under the Worst Case scenario, isolation caused by Highrock Lake will likely cause genetic drift or genetic bottle necks to occur. Additionally, these small, fragmented subpopulations will be susceptible to extreme weather events anticipated to occur in the Worst Case scenario as well as other stochastic events. Under the Worst Case scenario, the holiday darter is not expected to persist and will likely be extirpated.

Table 8. Estimated resilience factors and overall resiliency condition for holiday darter populations under the Worst Case scenario

Approximate Occurrence Occurrence Physical Connectivity Water Hydrologic Overall Abundance Extent Complexity Habitat Quality Regime Condition Conasuaga Low Low Low Moderate High Moderate Moderate Low River Talking Rock 0 0 0 Moderate High Low Moderate Likely Creek Extirpated Elijay River 0 0 0 Low Low Low Low Likely Extirpated Mountaintown 0 0 0 Low Low Low Low Likely Creek Extirpated Amicalola Low Low Low Moderate Low Moderate Moderate Low Creek Etowah River Low Low Low Moderate High Low Moderate Low

Shoal Creek 0 0 0 Moderate Low Moderate Low Likely Extirpated

Representation

Representation is expected to decline under the Worst Case scenario. Total known morphological variability is expected to be reduced by approximately 40% with the loss of all populations within the Coosawattee River and the Shoal Creek population. Physiographic province variability will be exhibited solely by holiday darters occupying the Blue Ridge in Tennessee and Georgia. Due to degraded habitat and water quality in downstream reaches of occupied rivers, connectivity is unlikely to be restored among populations and will continue to limit the exchange of novel and beneficial adaptations and migration to more suitable habitat. This limited representation will reduce the adaptive potential of the holiday darter by 2070.

Redundancy

Redundancy for the holiday darter is characterized by having multiple, resilient and representative populations distributed throughout its range. Under the Worst Case scenario, overall redundancy will be reduced by 57% due to the extirpation of four populations. All remaining extant populations are expect to have low resiliency. Under the Status Quo scenario, the holiday darter is expected to have low redundancy. Extant population will have experienced range reductions, low abundances, and/or have low spatial complexity.

Connectivity is not expected to improve for the species, range-wide under the Worst Case scenario. Dams have completely isolated the six populations into three groups. As mentioned earlier, the overall lack of connectivity between populations increases the importance of localized stochastic events and the species as a whole is less robust to smaller, more probable, and potentially more frequent stochastic events. Therefore, a key component of resiliency – minimization of the effect of localized extirpation– is not met.

Summary

The future scenario assessment has sought to understand how viability of the holiday darter may change over the course of 50 years in the terms of resiliency, representation, and redundancy. To account for considerable uncertainty associated with future projections, we defined three scenarios that would capture the breadth of changes likely to be observed in the upper Coosa River basin to which the holiday darter will be exposed. These scenarios considered two primary

79 elements of change: urbanization (Terando et al. 2014, entire) and climate change (IPCC 2013, entire). While we consider these scenarios plausible, we acknowledge that each scenario has a different probability of materializing at different time steps. To account for this difference in probability, a discretized range of probabilities (Table 4) was used to describe the likelihood a scenario will occur based on professional judgment (Table 5).

Table 9. Explanation of confidence terminologies used to estimate the likelihood of a scenario (after IPCC guidance, Mastrandrea et al. 2011)]

Confidence Terminology Explanation Very likely Greater than 90% certain Likely 70-90% certain As likely as not 40-70% certain Unlikely 10-40% certain Very unlikely Less than 10% certain

Table 10. Likelihood of a scenario occurring at 10 and 50 years.

Status Quo Best Case Worst Case 10 years Very likely Unlikely As likely as not 50 years Likely Unlikely As likely as not

In the Status Quo scenario three extant populations of holiday darter are expected to become extirpated. This will decrease overall redundancy for the species as well as representation (the Coosawattee River will no longer be represented with the extirpation of the Talking Rock Creek, Elijay River, and Mountaintown Creek populations). Physiographic representation will decline in the future because the holiday darter’s range is expected to contract out of the Piedmont and Ridge and Valley of Georgia and Tennessee to upstream stream reaches that are owned and managed by state and federal agencies within the Blue Ridge physiographic province. Representation will remain within the Ridge and Valley of Alabama. This scenario is very likely and likely within 10 and 50 years, respectively (Table 4).

The Best Case scenario assumed slightly slower rates of development and wider implementation of conservation activities that reduced the impact of development on streams. Additionally, this scenario assumed that through broad climate policies greenhouse gas emissions would begin to peak and decline, decreasing the likelihood of pervasive extreme climate events. Under this scenario, two populations of holiday darer (Ellijay River and Talking Rock Cree) would become extirpated, the remaining populations of holiday darter would persist. Due to continued urban growth and delayed response from conservation actions being implemented, we considered range expansions to be unlikely even under this scenario. As a result, the holiday darter is expected to slightly benefit under the Best Case scenario due to increased resiliency of some populations.

This scenario is considered unlikely to occur in both 10 and 50 year projections because it is unlikely that the broad conservation actions and policies will be implemented that require this scenario to materialize.

In the Worst Case scenario urban growth proceeds more quickly than current rates, conservation actions are not implemented widely, and the likelihood of extreme climate events increase as greenhouse gas emissions continue to increase. As a result, four populations become extirpated and three remain extant. The populations that remain decline in resiliency due to range contractions out of privately held land and into publicly held lands in the Blue Ridge and are exposed to extreme climate events. Like the Status Quo scenario, the Worst Case scenario sees an overall decrease in redundancy and representation. This scenario has is “as likely as not” to occur within 10 years and within 50 years.

Overall Summary

Currently, the holiday darter continues to occupy all streams where it was known to occur historically. Our assessment of current range size compared to historical range size (see current population discussions in Chapter 5) found that all populations currently occur over shorter overall stream lengths than historical records, indicating range reduction in these populations. No population of holiday darter currently exhibits high resiliency due to the reduction in extent of occupied habitat, low abundance of individuals per collection record, a simple linear, arrangement of records, as well as stressors affecting habitat and water quality. Similarly, representation and redundancy is currently low for this species because multiple resilient populations are lacking, connectivity is limited among populations, and this species is increasing becoming isolated to the upstream limits of its range in the Blue Ridge physiographic province.

Our future scenarios assessment considered the current viability of the species to project likely future viability given plausible scenarios of urban development and climate change. Only in the Best Case scenario did the species persist in all known populations. However, under this scenario holiday darters were not expected to expand outside of historical range boundaries and resiliency was expected to be moderate at best. Two and three populations were extirpated under the Status Quo and Worst Case scenarios, respectively. Resiliency, representation, and redundancy declined in both these scenario due to further range contractions and increased likelihoods for extreme climatic events to impact populations.

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