Species Status Assessment (SSA) Report for the Purpledisk Honeycombhead (Balduina Atropurpurea)

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Species Status Assessment Report for the Balduina atropurpurea (Purpledisk honeycombhead)

Version 1.0

Photo by Dee Mincey, Department of Defense, Fort Stewart, Georgia

August 2019

U.S. Fish and Wildlife Service Southeast Region Atlanta, GA

ACKNOWLEDGEMENTS

This document was prepared by the U.S. Fish and Wildlife Service’s Balduina atropurpurea Species Status Assessment Team (April Punsalan, Nicole Rankin, Caroline Krom, and Erin Rivenbark). We also received substantial assistance from Melanie Olds (USFWS – South Carolina Ecological Services Office), Todd Jones-Farrand (Region 4 – Science Applications), and Carlos Ramirez (Mississippi State University). Habitat suitability and future projection modeling analyses were performed by Mississippi State University.

We would also like to recognize and thank the following individuals who provided substantive information, photos, and/or insights for our species status assessment. Thank you to Tom Patrick, Richard Porcher, Dee Mincey, Jacob Thompson, Stella Osborn, Bobby Hattaway, Scott Wiggers, David Lincicome, Jennifer Ceska, Linda Chafin, Wilson Baker, Amy Jenkins, Keith Bradley, Nicole Hawkins, Herrick Brown, Anna Smith, Jennifer Walls, Al Schotz, Frank Price, Ron Determann, Bruce Sorrie, and Lawrence Carlile.

Valuable peer review of a draft of this document were provided by Laura Robinson. In addition, Lisa Kruse, Michael Jenkins, and Lesley Stark provided partner review of the same draft of this document. We appreciate their input and comments, which resulted in a more robust status assessment and final report.

Suggested reference:

U.S. Fish and Wildlife Service. 2019. Species status assessment report for the Balduina atropurpurea (purpledisk honeycombhead), Version 1.0. August 2019. Atlanta, GA.

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VERSION UPDATES

The changes from Version 0.1 (April 2019) to 0.2 (May 2019) are mostly minor grammar, editorial, and subject matter clarifications. Two substantive changes were made to reflect peer reviewer comments: 1) Changed the current condition resiliency score from low resiliency to very low resilency for the Rosindale population located in North Carolina. 2) Added an explanation of status codes including extant, extirpated, historical, occurrence.

The changes from Version 0.2 (May 2019) to 1.0 (July 2019) are minor and do not change the SSA analysis for Balduina atropurpurea. The changes were: 1) Updated Figure 4.2 to display the map title without the draft date. 2) Updated Figure 4.3 to display the map title without the draft date. 3) Updated Figures 4.5, 4.6, 4.7, 4.8, 4.9, 4.10, and 4.11 to reflect changes in three resiliency scores. 4) Updated Figures 5.3 (a,b), 5.4 (a,b), and 5.5 (a,b) to reflect changes in the future projections of three resiliency scores.

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EXECUTIVE SUMMARY

This report summarizes the results of a Species Status Assessment completed for Balduina atropurpurea (purpledisk honeycombhead), hereafter called Balduina, to assess the species’ overall viability. We considered what the species needs to maintain viability by characterizing the status of the species in terms of its resiliency, representation, and redundancy (3 R’s). We provide a thorough assessment of the biology and ecological needs of the species, followed by a description of the factors influencing viability, the current condition of the species, and predicted future conditions.

Balduina is a Southeastern Coastal Plain endemic found in fire-adapted pine savanna and flatwood ecosystems of Florida, Georgia, South Carolina, North Carolina, and historically in Alabama. The distinguishing characteristics of Balduina include a honeycomb receptacle, small basal rosette, and purple-to-marron disk flowers. Balduina is a facultative wetland species flowering from mid-August to mid-October and fruiting from October to November. The four life stages of Balduina include seeds, basal rosette, basal rosette with vegetative stalk, and reproductive individuals. The life history and resource needs of Balduina appears tightly interlinked with high relative light levels (little to no shading at or near the ground surface) and moisture availability. Balduina requires cross-pollination for viable seed production (Parker and Jones 1975, p. 358).

The primary factors impacting the viability of Balduina are habitat based: habitat loss due to development or land conversion (e.g. agriculture, pine plantations, etc.) and degradation due to fire suppression. Across Balduina’s range, the transition zone between longleaf pine uplands and aquatic wetlands has been heavily impacted by habitat destruction and modification. Large tracts of land, containing both uplands and aquatic wetlands, are needed to protect these transitions zones. Further, Balduina and its habitat requires frequent fire prescription to maintain the open conditions in these mesic transition zones to abate woody encroachment and facilitate nutrient releases. Other potential factors influencing the viability of Balduina include non-native invasive species (i.e. feral hogs) and climate change.

For the purpose of this assessment, we defined viability as the ability of Balduina to sustain populations in pine savanna and flatwood ecosystems over time. Using the SSA framework, we describe the viability of Balduina by defining populations, assessing current conditions, and predicting the future condition using the metrics of the 3Rs.

We delineated Balduina populations using NatureServe’s Habitat-based Plant Element Occurrence Delimitation Guidance (NatureServe Explorer 2018, unpaginated). For each Balduina element occurrence, we used a 2-kilometer (km; 1.24 miles (mi)) separation distance rule to separate populations. When occurrences within 2 km were disjunct from one another for more than 1 km due to agriculture fields or pine plantations, we used a 1 km separation distance rule to separate populations. Overall, we delineated 79 Balduina populations from 140 element occurrence records.

To assess the current condition and resiliency of Balduina, we revised a habitat and population metrics datasheet. The habitat metrics included five metric categories that were designed to

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assess the vegetation, hydrology, and fire management of each Balduina population. In addition to the habitat metrics, we used two population metrics (number of individuals per population and number of occurrences within a population) to assess resiliency of Balduina populations. For Balduina populations (28) without a completed habitat and population metrics datasheet, we compiled state heritage data and used the two population metrics (number of individuals per population and number occurrences within a population) to compute population resiliency. Six Balduina populations did not have completed metric datasheets or state heritage data that included the number of individuals. Therefore, we used the best available science to assign a resiliency score based on the time since last seen, land protection status, and number of occurrences per population. For representation and redundancy, we assessed the distribution of Balduina populations across 6 representation units.

Out of the 79 Balduina populations, 38 remain extant, 39 are historical, and two are extirpated. The majority (64) of Balduina populations occur in Georgia. There are five extant populations in Florida while one extant population remains in North Carolina and one in South Carolina. Thirty-nine Balduina populations are considered in historical status and two are extirpated. Of the 38 extant Balduina populations, five currently have high resiliency and four have moderate resiliency. Four of the highly resilient and two moderate resilient populations occur on protected land. Across the range, the remaining extant populations (29) have low (7) to very low (22) resiliency. Overall, Balduina has experienced a decline in the number of resilient populations within each of the representation units and across the range. The distribution of resilient populations is clustered in three representation units in Georgia and Florida. In addition, the current redundancy has been reduced from historical conditions.

To determine future projections, we used a habitat suitability model, SLEUTH model, and three management scenarios. We selected a high urbanization SLEUTH model to determine the projected percent of suitable habitat (determined from habitat suitability model) lost due to development at two time steps: 2040 and 2060. If a population was projected to lose greater than 25% of suitable habitat under the high urbanization scenario, it was assessed to have a high risk of development. Conversely, it had a low risk of development, if it was projected to lose 25% or less of suitable habitat. Then, we used the high and low risk of development to determine what would happen to the future resiliency of populations if management stayed the same (Scenario 1 – Status Quo), decreased (Scenario 2 – Decreased Management), or increased (Scenario 3 – Increased Management). The interaction between risk of development and management for the three scenarios was used to determine future resiliency at 2040 and 2060 for all extant Balduina populations.

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populations are predicted to have an increase in resiliency, resulting in 20 and 18 populations with low resiliency at 2040 and 2060, respectively. If management efforts decrease in the future as under Scenario 2, over half (22) of the currently extant Balduina populations (38) are predicted to be extirpated in 2040 and 2060. Currently, many of the Balduina populations (22) have very low resiliency and occur in fragmented habitat. As such, if management efforts decrease in the future, they will be lost from the landscape. Further, with a decrease in management, no high resiliency populations are predicted to occur in 2040 and 2060.

When assessing representation under Scenario 1, at 2040, Balduina is predicted to have 36 extant populations (38 populations currently) in varying levels of resiliency occurring in five ecoregions. At 2060, the species is predicted to have 34 extant populations occurring in five ecoregions. Therefore, the species has some level of adaptive capacity, but given the low to very low resiliency to extirpated status of 31 Balduina populations at 2040 and 32 populations at 2060, the future representation is similar to current levels of representation. In addition, when assessing redundancy under Scenario 1, the future species redundancy is predicted to remain similar to current redundancy.

Under Scenario 2, at 2040 and 2060, Balduina is predicted to have 16 extant populations (38 populations currently) in varying levels of resiliency occurring in three ecoregions. Given the low to very low resiliency to extirpated status of 35 Balduina populations at 2040 and 2060, the future representation is predicted to be greatly reduced under Scenario 2. Under Scenario 2, future redundancy is predicted to be further reduced from current conditions. Given the lack of connectivity and amount of isolated populations under this scenario, without human intervention and cultivated plants, it may be difficult to re-establish occurrences affected by a catastrophic event.

Under Scenario 3, at 2040 and 2060, Balduina is predicted to have 38 extant populations (38 populations currently) in varying levels of resiliency occurring in five ecoregions. Given the low to very low resiliency of 22 Balduina populations at 2040 and 2060, the future representation under Scenario 3 is predicted to increase from current representation levels. In addition, future redundancy is predicted to increase for Balduina. No currently extant populations are predicted to be extirpated under this scenario. Having higher resiliency populations spread across the range and representation units may allow occurrences within a population and low to very low resiliency populations to become re-established following a catastrophic event.

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TABLE OF CONTENTS

ACKNOWLEDGEMENTS ...... ii VERSION UPDATES ...... iii EXECUTIVE SUMMARY ...... iv LIST OF TABLES ...... ix LIST OF FIGURES ...... xi CHAPTER 1 – INTRODUCTION ...... 1 CHAPTER 2 – SPECIES BIOLOGY, HABITAT, AND RESOURCE NEEDS ...... 4 2.1 Taxonomy ...... 4 2.2 Species Description ...... 4 2.3 Habitat ...... 6 2.4 Range and Distribution ...... 6 2.4.1 Explanation of Status Codes ...... 11 2.5 Life History and Individual Resource Needs ...... 11 2.6 Population and Species Level Needs ...... 14 CHAPTER 3 – FACTORS INFLUENCING VIABILITY ...... 16 3.1 Habitat Destruction and Modification ...... 16 3.2 Lack of Fire Management ...... 18 3.3 Hydrological Impacts ...... 20 3.4 Non-Native Invasive Species ...... 20 3.5 Climate Change ...... 21 3.6 Conservation Measures ...... 23 3.7 State Protections...... 24 3.8 Synergistic Effects ...... 24 3.9 Summary of Factors Influencing Viability ...... 25 CHAPTER 4 – POPULATION AND SPECIES NEEDS AND CURRENT CONDITION ...... 27 4.1 Delineating Populations ...... 27 4.2 Methods for Estimating Current Condition ...... 27 4.2.1 Population Resiliency ...... 27 4.2.2 Species Representation and Redundancy...... 31

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4.3 Current Condition ...... 32 4.3.1 Populations ...... 32 4.3.2 Current Population Resiliency ...... 33 4.3.3 Current Species Representation ...... 37 4.3.4 Current Species Redundancy ...... 45 CHAPTER 5 – FUTURE CONDITIONS AND VIABILITY ...... 47 5.1 Introduction to Methods for Estimating Future Condition ...... 47 5.2 Projections and Modeling ...... 47 5.3 Incorporating Management into the Future Scenarios ...... 50 5.4 Future Condition ...... 52 5.4.1 Future Population Resiliency ...... 52 5.4.2 Future Species Representation ...... 55 5.3.3 Future Species Redundancy ...... 63 LITERATURE CITED ...... 64 APPENDIX A ...... 70 APPENDIX B ...... 72 APPENDIX C ...... 76 APPENDIX D ...... 80

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LIST OF TABLES

Table 2-1. The diagnostic morphological characters across three species within the genus Balduina ...... 6

Table 2-2. Balduina atropurpurea individual resource needs by life stage ...... 13

Table 3-1. Protected lands that provide suitable habitat for extant Balduina atropurpurea populations ...... 24

Table 4-1. Habitat and population metrics used to assess and determine the current condition and population resiliency of Balduina atropurpurea ...... 30

Table 4-2. Current resiliency of Balduina atropurpurea populations across the species’ range ..35

Table 4-3. Current resiliency classes for protected and unprotected Balduina populations across the species’ range ...... 36

Table 4-4. Balduina atropurpurea representation units with the number of records, number of populations, and current population resiliency scores ...... 38

Table 5-1. Environmental variables used in habitat suitability modeling for Balduina atropurpurea ...... 48

Table 5-2. Percent habitat loss due to urbanization scenarios within each Balduina atropurpurea representation unit ...... 50

Table 5-3. Interaction between risk of development and management in three plausible future scenarios to determine future resiliency of Balduina atropurpurea populations ...... 51

Table 5-4. Future resiliency for Balduina atropurpurea populations at 2040 under three future scenarios ...... 52

Table 5-5. Future resiliency for Balduina atropurpurea populations at 2060 under three future scenarios ...... 53

Table 5-6. Balduina atropurpurea population resiliency across for the six representation units under three future scenarios at 2040 and 2060...... 57

Table B-1. Current resiliency computed from completed Habitat and Population Metric Datasheets for extant Balduina atropurpurea populations in 2018 ...... 72

Table B-2. Current resiliency computed using population metric data for Balduina atropurpurea populations from State heritage programs ...... 73

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Table C-1. Future resiliency for extant Balduina atropurpurea populations at 2040 under three future scenarios ...... 76

Table C-2. Future resiliency for extant Balduina atropurpurea populations at 2060 under three future scenarios ...... 78

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LIST OF FIGURES

Figure 1-1. Species Status Assessment Framework ...... 2

Figure 2-1. Balduina atropurpurea basal rosette, flowers/inflorescences, and habitat ...... 5

Figure 2-2. Botanical illustration of Balduina atropurpurea...... 5

Figure 2-3. Balduina atropurpurea range map displaying extant and historical populations across counties within the species range ...... 7

Figure 2-4. Dense vegetation along powerline right-of-way where Balduina atropurpurea was previously found in 2004 at the Rosindale Longleaf Pine Forest in Bladen County, North Carolina ...... 8

Figure 2-5. Balduina atropurpurea population in open conditions at Fort Stewart, Georgia ...... 9

Figure 2-6. Largest Balduina atropurpurea population in Florida located in a powerline right-of- way in Ralph E. Simmons Memorial State Forest, northern Nassau County ...... 10

Figure 2-7. Life history stages of Balduina atropurpurea: seed, basal rosette, basal rosette with vegetative stalk, and reproductive individual ...... 12

Figure 3-1. Presettlement range and major divisions of the longleaf pine ecosystem ...... 16

Figure 3-2. This map displays agriculture production and land conversion on private land and protected land (Fort Stewart) across Long, Liberty, Evans, and Tattnall counties, Georgia ...... 18

Figure 3-3. Historic (1650–1850 CE) mean fire interval estimates for the presence of fire in all or part of an average 1.2 km2 area ...... 19

Figure 3-4. Time series of the seasonal average of maximum air temperature in the South Atlantic region ...... 21

Figure 3-5. Time series of the seasonal average of precipitation in the South Atlantic region .....22

Figure 3-6. Influence diagram for Balduina atropurpurea showing relationships between factors and species’ viability...... 26

Figure 4-1. Example of delineating populations using the 2-km separation rule ...... 27

Figure 4-2. Representation units for Balduina atropurpurea ...... 32

Figure 4-3. Distribution of 140 Balduina atropurpurea occurrence records across the species' range ...... 33

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Figure 4-4. Balduina atropurpurea current population resiliency scores across the species' range ...... 35

Figure 4-5. Balduina atropurpurea current population resiliency across representation units .....39

Figure 4-6. Balduina atropurpurea current population resiliency in the North Carolina representation unit ...... 40

Figure 4-7. Balduina atropurpurea current population resiliency in the South Carolina representation unit ...... 41

Figure 4-8. Balduina atropurpurea current population resiliency in the Georgia North representation unit ...... 42

Figure 4-9. Balduina atropurpurea current population resiliency in the Georgia South representation unit ...... 43

Figure 4-10. Balduina atropurpurea current population resiliency in the Georgia-Florida representation unit ...... 44

Figure 4-11. Balduina atropurpurea current population resiliency in the Alabama representation unit ...... 45

Figure 5-1. Balduina atropurpurea future population resiliency across the species' range at 2040 under three future scenarios ...... 52

Figure 5-2. Balduina atropurpurea future population resiliency across the species' range at 2060 under three future scenarios ...... 53

Figure 5-3. Balduina atropurpurea population resiliency across representation units for Scenario 1 (status quo management with high urbanization) at two time steps (a) 2040 and (b) 2060 ...... 60

Figure 5-4. Balduina atropurpurea population resiliency across representation units for Scenario 2 (decreased management with high urbanization) at two time steps (a) 2040 and (b) 2060...... 61

Figure 5-5. Balduina atropurpurea population resiliency across representation units for Scenario 3 (increased management with high urbanization) at two time steps (a) 2040 and (b) 2060 ...... 62

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CHAPTER 1 – INTRODUCTION

In this chapter, we discuss the previous Federal actions including petition history for Balduina atropurpurea (purpledisk honeycombhead), hereafter Balduina, and the analytical framework used to evaluate the status of the species.

Balduina is a perennial herb found in pine savanna and flatwood ecosystems of Florida, Georgia, South Carolina, North Carolina, and with potential records in Alabama and North Carolina. Balduina was initially recognized as a candidate species in the U.S. Fish and Wildlife Service’s (Service) first plant notice on July 1, 1975 (40 FR 27823), which indicated that the original Smithsonian plant report that included Balduina had been accepted as a listing petition under the Act. It was removed from the candidate list in the Service’s plant notice on December 15, 1980 (45 FR 82480-82569), where it was listed as a 3C species-taxa that had proven to be more abundant or widespread than was previously believed and/or that are not subject to an identifiable threat. On September 30, 1993 (50 FR 51145), Balduina was again identified as a candidate species, a category 2(d) taxa- “taxa for which information now in the possession of the Service indicates that proposing to list as endangered or threatened is possibly appropriate” and the status trend was noted as declining (d). On February 28, 1996, the Service discontinued the designation of Category 2 species as candidates (61 FR 7596).

On April 20, 2010, the Service was petitioned to list 404 aquatic species, including Balduina, in the southeastern United States as an endangered or threatened species under the Endangered Species Act of 1973, as amended (16 U.S.C. 1531-1543; Act) and designate critical habitat (CBD 2010, entire). On September 27, 2011, we published a 90-day finding, which determined that the petition contained substantial information indicating the Balduina may warrant listing (76 FR 59836). Therefore, a review of the status of the species was initiated to determine if the petitioned action is warranted. Based on the status review, the Service will issue a 12-month finding for the Balduina. Thus, we conducted a Species Status Assessment (SSA) to compile the best available data regarding the species’ biology and factors that influence the species’ viability. The Balduina atropurpurea SSA Report is a summary of the information assembled and reviewed by the Service and incorporates the best scientific and commercial data available. This SSA Report documents the results of the comprehensive status review for the Balduina and serves as the biological underpinning of the Service’s forthcoming decision on whether the species warrants protection under the Act.

The SSA framework (Service 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 Ecological Services Program of the Service, from Candidate Assessment to Listing to Consultations to Recovery. As such, the SSA Report will be a living document that may be used to inform Endangered Species Act decision making, such as listing, recovery, Section 7, Section 10, and reclassification decisions (the latter four decision types are only relevant should the species warrant listing under the Act). Therefore, we have developed this SSA Report to summarize the most relevant information regarding life history, biology, and considerations of current and

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future risk factors facing the Balduina. In addition, we forecast the possible response of the species to various future risk factors and environmental conditions to formulate a complete risk profile for the Balduina. In conducting this analysis, we took into consideration the likely changes that are happening in the environment – past, current, and future – to help us understand what factors drive the viability of the species.

For the purpose of this assessment, we define viability as the ability of the Balduina to sustain populations in pine savanna and flatwood ecosystems over time. Viability is not a specific state, but rather a continuous measure of the likelihood that the species will sustain populations over time (Service 2016, p. 9). 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 (Service 2016, entire; Wolf et al. 2015, entire). To evaluate the viability of the Balduina, we estimated the current condition and predicted future condition of the species in terms of resiliency, redundancy, and representation.

• Resiliency describes the ability of a population to withstand stochastic disturbance. Stochastic events are those arising from random factors such as weather, flooding, or fluctuations in reproduction rates. Resiliency is positively related to population Figure 1-1. Species Status size and growth rate and may be influenced by connectivity among Assessment Framework. populations. Generally speaking, populations need enough individuals, within habitat patches of adequate area and quality, to maintain survival and reproduction in spite of disturbance. Resiliency is measured using metrics that describe population condition and habitat; in the case of the Balduina, we used habitat and population metrics to assess resiliency.

• Representation describes the ability of a species to adapt to changing environmental conditions over time. Representation can be measured by reviewing the genetic diversity within and among populations and the ecological diversity (also called environmental variation or diversity) of populations across the species’ range. Theoretically, the more representation the species has, the higher its potential of adapting to changes (natural or human caused) in its environment. In the absence of species-specific genetic information, we assessed representation by evaluating the number and distribution of Balduina populations across ecoregions.

• Redundancy describes the ability of a species to withstand catastrophic events. A catastrophic event is defined as a rare, destructive event or episode that may have impacts to multiple occurrences within a population or multiple populations. Redundancy is about spreading the risk among populations, and thus, is assessed by characterizing the number of resilient populations across the range of the species. The more higher resiliency populations the species has, distributed over a larger area, the better chances that the species can withstand catastrophic events. For Balduina, we determined the number and distribution of resilient populations across the range to measure redundancy.

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This SSA Report provides a thorough assessment of the biology and natural history of the Balduina and assesses demographic risks, stressors, and limiting factors in the context of determining the viability and risks of extinction for the species. Importantly, this SSA Report does not result in, nor predetermine, any decisions by the Service under the Act. In the case of the Balduina, this SSA Report does not determine whether the Balduina warrants protections of the Act, or whether it should be proposed for listing as an endangered or threatened species under the Act. That decision will be made by the Service after reviewing this document, along with the supporting analysis, any other relevant scientific information, and all applicable laws, regulations, and policies. The results of the decision will be announced in the Federal Register. The contents of this SSA Report provide an objective, scientific review of the available information related to the biological status of the Balduina.

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CHAPTER 2 – SPECIES BIOLOGY, HABITAT, AND RESOURCE NEEDS

In this chapter, we provide biological information about Balduina, including its taxonomic history, morphological description, historical and current distribution throughout its range, and known life history. Lastly, we outline the resource needs of individuals.

2.1 Taxonomy

Balduina atropurpurea is a member of the Asteraceae (Aster) family. The genus Balduina includes three species: B. atropurpurea, B. uniflora (Nutt), and B. angustifolia (Pursh) Robinson. All three species occur in the coastal plain in the southeastern United States from North Carolina westward into Alabama and southward into Florida (Parker and Jones 1975, p. 355). Both B. atropurpurea and B. uniflora have been found growing among each other and flowering concurrently at Fort Stewart in Georgia (Patrick 2018, pers. comm.). Further, both Balduina species were found flowering concurrently at Florida State Parks (Walls 2018, pers. comm.). As such, there is a chance that where both species, B. atropurpurea and B. uniflora, coexist, hybridization could occur; however, to date, research has not been done to determine if hybrids produce viable seed.

Balduina atropurpurea (Harper) was first described as a new species in 1901 from a collection made in Berrien County, Georgia (Harper 1901, p. 483). Endorima atropurpurea (Harper) Small is the only synonym ever used, but was not accepted because the name Balduina Nutt is one year older than Endorima (Wunderlin 1983, p. 5). Therefore, Balduina atropurpurea is currently recognized as a valid taxon (Weakley 2015, p. 1089), and the currently accepted classification is:

Class: Magnoliopsida – Dicotyledons Order: Asterales Family: Asteraceae Species: Balduina atropurpurea

2.2 Species Description

Balduina is a perennial herb distinguished from other species in the genus by its dark purple disk flowers (Figure 2-1; Figure 2-2; Table 2-1). The yellow ray flowers extend out to 3 centimeters (cm; 1.18 inches (in)) and have three-to-five teeth at the tip (Kral 1983, p. 1107). The showy, inflorescences emerge from long stiff, slightly hairy peduncles (flowering stem) that have characteristic longitudinal ridges along the stem (Chafin 2007, p. 59). The honeycomb-like receptacle, responsible for the common name, forms from connected receptacular bractlets (reduced leaflike structures at the base of each achene) that surround the achenes (small, indehiscent fruit with a single seed) (Weakley 2015, p. 1089).

The plant grows to a height of 6-8 decimeters (dm; 23.6-31.5 in). The basal linear-spatulate leaves (wider at the leaf apex and gradually tapering towards the leaf base) are 20 times as long as wide (Weakley 2015, p. 1089). Stem/cauline leaves ascend the stem, gradually reducing in size as they reach the peduncle (Kral 1983, p. 1107). The stems and leaves arise from a stout, shallow underground rhizome.

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(a) (b) (c)

Figure 2-1. Balduina atropurpurea basal rosette (a), flowers/inflorescences (b), and habitat (c). Photos by Richard Porcher, The Citadel (a, b) and Jenn Walls, Florida Forest Service (c).

Figure 2-2. Botanical illustration of Balduina atropurpurea. Illustration by Jean C. Putnam Hancock and courtesy of Linda Chafin, State Botanical Garden of Georgia.

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Table 2-1. The diagnostic morphological characters across three species within the genus Balduina (adapted from Parker and Jones 1975, pp. 359-360; Kral 1983, p. 1107; Wunderlin 1983, pp 5-6.; Weakley 2015, p. 1089). Balduina atropurpurea Balduina uniflora Balduina angustifolia Character Purpledisk Savanna Coastal Plain honeycombhead honeycombhead honeycombhead Duration Perennial Perennial Annual or Biennial

Yellow to reddish- Disk flowers Purple Yellow orange Cauline/stem Few, ascending Few, 5 to 10 cm long Numerous, linear leaves 3.8-6.2 cm long Flowering Late August-early Late July-September Mainly June-November Period November

2.3 Habitat

Balduina occurs in variety of habitat types where moisture and light are conducive for growth throughout the pine savanna and flatwood ecosystem. These areas typically have poorly drained soils that are acidic, sandy, and nutrient-poor (Lincicome 1998, p. 25). A clay hardpan exists close to the surface and inhibits water from infiltrating deep into the earth. Laterally, water slowly moves downslope across the pine uplands through sandy soil, which keeps the water from evaporating due to poor heat conduction, and eventually seeps out at lower levels (Plummer 1963, p. 728; Edwards et al. 2013, pp. 430-431). Balduina occurs in the low, wet areas where water emerges from the soil surface and creates either a seepage slope, pitcherplant bog, or wet to mesic savanna. Balduina also occurs in habitat created by artificial means such as powerline rights-of-way and wet ditches.

Large-scale or small-scale disturbance caused primarily by fire has shaped and characterized the wet pine savannas, seepage slopes, and pitcherplant bogs of the southeastern Coastal Plain (Brewer 1999, p. 159) where Balduina occurs. Fire reduces woody competition, reduces graminoid coverage, and releases minerals and nutrients from the soil (Plummer 1963, p. 730). Low-intensity, regular burn cycles (1 to 3 years) maintain the pine savannas’ diverse herb- dominated communities by bringing sufficient light to the surface soil/ground layer (Martin and Kirkman 2006, p. 907) and releasing nutrients bound in the duff layer. The released nutrients are carried by rainwater and sheet flow into the low, wet areas where Balduina occurs.

2.4 Range and Distribution

Balduina occurs within the Coastal Plain from North Carolina south to Florida. Historically, it occurred in Alabama (Figure 2-3).

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Figure 2-3. Balduina atropurpurea range map displaying extant and historical populations across counties within the species range.

North Carolina The species is known from Bladen and Brunswick counties in North Carolina. One historical Balduina population is known from the Green Swamp Preserve, a 15,722-acre preserve owned and managed by The Nature Conservancy (TNC), located in Brunswick County. The population was first identified on October 26, 1980 by Hervey McIver (NC Heritage Program 2016, p. 1). One B. atropurpurea specimen with visible maroon disk flowers was found and used for positive identification. The other Balduina individuals were too mature and/or senescing to key (McIver 2018, pers. comm.). Since that time, more surveys have been conducted to search for B. atropurpurea, but B. uniflora was only found at the occurrence (McIver 2018, pers. comm.). Prior to TNC acquisition of the Green Swamp Preserve, the occurrence was owned by a timber company and clearcut in 1977. From 1955 to 2017, fire prescription occurred intermittingly, with an average 5-year rotation (West 2018, pers. comm.). Despite several recent surveys (1994 and 2004) occurring in the area, B. atropurpurea has not been relocated.

One extant Balduina population occurs at Rosindale Longleaf Pine Forest on private property located in Bladen County, North Carolina. The population was first identified on September 13, 2004 and located underneath a powerline easement within a seepage slope savanna (NC Natural

SSA Report – B. atropurpurea 7 August 2019

Heritage Program 2016, p. 3). This population has not been seen since first discovered. No individuals were found during a 2018 October field survey, and the occurrence was overgrown with vegetation (Figure 2-4). The NC Heritage Program ranks this occurrence as “failed to find,” and it is still considered extant since it was discovered in 2004, 15 years ago (NC Heritage Program 2016, p. 3).

Figure 2-4. Dense vegetation along powerline right-of-way where Balduina atropurpurea was located in 2004 at the Rosindale Longleaf Pine Forest in Bladen County, North Carolina. Photo by Caroline Krom, U.S. Fish and Wildlife Service.

South Carolina Balduina is known from Darlington and Richland counties in South Carolina. One extirpated (Hartsville North) and one historical (Society Hill) population occurs in Darlington County. The Hartsville North Balduina population once occurred at the “edge of dense bottomland woods” (SC Department of Natural Resources 2018, geospatial data). This population was destroyed by industrial development (Brown 2018, pers. comm.). In 1857, a Balduina specimen was collected in Society Hill by Dr. M.A. Curtis. The herbarium record was first identified as B. angustifolia and later annotated as B. atropurpurea (Brown 2018, pers. comm.). There is no habitat information associated with this record.

The one extant Balduina population occurs on Fort Jackson, a Department of Defense Army installation, in Richland County. The population was first located on September 5, 1994, during an endangered species survey on Fort Jackson. The population was last surveyed and observed in 2017 (Bradley 2018, pers. comm.). The population occurs in an open Sandhill Seepage Savannah that is seasonally to permanently saturated. Although the number of individuals has not been reported for this population, it has been reported as a “small colony” (Bradley 2018, pers. comm.).

SSA Report – B. atropurpurea 8 August 2019

Georgia The greatest concentration of Balduina occurs in the Coastal Plain of Georgia. The slight topographic relief and low elevation across the Coastal Plain of Georgia results in soil saturation and shallow inundation when elevation varies (Edwards et al. 2013, p. 2), resulting in ideal habitat for Balduina. Further, due to the local relief (less than 30 meters (100 feet)), fire has played in an important role in this region, occurring approximately every 1 to 3 years presettlement (Frost 1988, p. 76).

There are 31 extant, 32 historical, and one extirpated population(s) occurring across 21 counties (Ben Hill, Brantley, Bulloch, Candler, Charlton, Coffee, Colquitt, Cook, Emanuel, Evans, Irwin, Jeff Davis, Jenkins, Liberty, Tattnall, Long, Tift, Toombs, Turner, Wayne, Worth) in the Coastal Plain of Georgia. Balduina has been extirpated from six of the counties listed above, including Brantley, Bulloch, Candler, Emanuel, Tift, and Wayne counties. On private lands, powerline rights-of-way (ROW) support many of the extant Balduina populations in Georgia. However, the majority of these populations contain only one occurrence with fewer than 50 individuals.

The largest Balduina population occurs on Fort Stewart, the largest Department of Defense Army installation east of the Mississippi River and covering 280,000 acres. The large amount of suitable habitat combined with the historical and current fire frequency on Fort Stewart has resulted in the largest population of Balduina across its range. Balduina occurs on a variety of habitat types within Fort Stewart, including, pine savannas, pine flatwoods, cypress domes, seepage slope, woodland borders near streams, and sandhill seeps (Helton 1995, p. A-13). Balduina populations on Fort Stewart were surveyed in 1994, 1995, and 2018 (Helton 1995, pp. 1-38; Lincicome 1998, pp. 1-131; Hattaway and Osborn 2018, pp. 1-71). The 2018 surveys revealed that populations that increased in size since 1995 were “located either in wide open, savannah-like areas with few trees and near seeps, springs, swales, or along the edge of a wetland with ample light coming in from the higher ground” (Figure 2-5) (Hattaway and Osborn 2018, p. 8).

Figure 2-5. Balduina atropurpurea population in open conditions at Fort Stewart, Georgia. Photo by April Punsalan, U.S. Fish and Wildlife Service.

SSA Report – B. atropurpurea 9 August 2019

Florida Balduina is known from Clay, Duval, and Nassau counties and historically occurred in Putnam county in Florida. There are five extant and four historical populations that occur across the Sea Island Flatwoods subecoregion of northeast Florida. All extant populations occur on protected land including Ralph E. Simmons Memorial State Forest, Cary State Forest, Jennings State Forest, and St. Mary’s River Ranch. The majority of historical populations occur along roadside rights-of-way and have been impacted by road widening activities.

The largest Balduina population (approximately 700 individuals) in Florida occurs on Ralph E. Simmons Memorial State Forest in northern Nassau County (Walls and Jenkins 2018, pp. 1-2). The State forest contains approximately 3,638 acres of Longleaf Pine/Wiregrass Flatwoods and Savanna Communities, Seepage Slopes, and Pitcherplant Bogs. The St. Johns River Water Management District owns the property and has a management agreement with the Florida Forest Service. Balduina individuals on the State forest occur along a powerline right-of-way and within a pine savanna and flatwood ecosystem. Management, e.g., mowing, within the powerline has kept the habitat completely open from woody encroachment and thereby supported the robust Balduina population (Figure 2-6). The last prescribed fire in the area occurred in 2015 and did not reach the population (Walls and Jenkins 2018, p.2).

Figure 2-6. Largest Balduina atropurpurea population in Florida located predominately in a powerline right-of-way in Ralph E. Simmons Memorial State Forest, northern Nassau County. Photo by Jenn Walls, Florida Forest Service.

The second largest Balduina population in Florida occurs in Cary State Forest, located near Jacksonville in Nassau and Duval counties. The State forest contains 13,385 acres of Longleaf Pine Flatwoods and Mesic Savannas, Sandhills, Basin Swamps, and Cypress Domes.

SSA Report – B. atropurpurea 10 August 2019

The Balduina population occurs in a powerline right-of-way along a 1.5-mile stretch and contains approximately 200 individuals (Walls and Jenkins 2018, pp.1-2). Mowing and prescribed fire has maintained the occurrence in an open condition thereby supporting a self- sustaining Balduina population. The population was last burned in 2011 and 2016.

Three Balduina populations occur on Jennings State Forest, located in northeastern Florida in the northern section of Clay County. One population with 32 individuals was found in 2018.

Alabama One historical Balduina population reportedly occurred in Geneva County, Alabama (Schotz 2018, pers. comm.); however, the herbarium specimen was lost. As such, the Alabama Plant Atlas has not listed Balduina as a species occurring in Alabama. Several surveys have been conducted to relocate the population, but no individuals have ever been rediscovered.

2.4.1 Explanation of Status Codes

For this SSA, we used the following status codes to define terms:

Element occurrence (occurrence) is an area of land where a species, i.e., Balduina, is, or was, present (NatureServe 2018, unpaginated).

A population is defined as Balduina occurrences that are separated by a 2-kilometer (km; 1.24 miles (mi)) distance or 1-km (0.62 mi) separation distance when occurrences are separated by artificial or fragmented habitat, e.g., roads, agriculture fields, etc.

Historical is an occurrence that has not been verified in over 20 years.

Extant is an occurrence that has been verified as still existing within the past 20 years and the habitat has not been destroyed.

Extirpated is defined as repeated surveys by experienced observers during the appropriate survey window indicate that the species no longer exists at the given location or that the habitat of the occurrence has been destroyed to such an extent that it can no longer support the species (NatureServe 2018, unpaginated).

2.5 Life History and Individual Resource Needs

Balduina is a facultative wetland species flowering from mid-August to mid-October and fruiting from October to November. The four life stages of Balduina include seeds, basal rosette, basal rosette with vegetative stalk, and reproductive individuals (Figure 2-6). Vegetative/asexual reproduction occurs from shallow, stout rootstocks. Sexual reproduction occurs from seed. All three species within the genus Balduina produce self-incompatible flowers, requiring cross- pollination for viable seed production (Parker and Jones 1975, p. 358).

The life history (Figure 2-7) and resource needs (Table 2-2) of Balduina appears tightly interlinked with high relative light levels (little to no shading at or near the ground surface)

SSA Report – B. atropurpurea 11 August 2019

(Lincicome 1998, pp. 26-71). Germination trials at Atlanta Botanical Gardens revealed that seeds may require light for germination (Lincicome 1998, p. 37). Germination was rapid in high light conditions, with 95% of seeds germinating after 14 days (9 days of 18 hours (hrs.) of light; 5 days of 24 hrs. of light). Similarly, seedling emergence in a greenhouse after 24 hrs. of light was rapid, with seedling emergence essentially complete by day 11 and ranging from day 8 to day 14. Vegetative health appears associated with light levels as well. Basal rosettes produce taller vegetative stalks in areas with relative high light in comparison with populations in low to moderate light conditions (Lincicome 1998, p. 71). Lastly, floral production appeared greater in high light conditions (Lincicome 1998, p. 74).

Moisture availability is important for all life stages of Balduina. Seed germination for many plant species, including Balduina, depends upon the constant imbibement of water through the seed coat. Further, for seeds as well as other life stages, percolating soil water from the uplands carries soluble nutrients to the system (Plummer 1963, p. 733) and habitat where Balduina occurs. Therefore, moisture availability is important to bring nutrients for Balduina seedlings and for other life stages and reproduction.

• Seedling • Seed dispersal may recruitment may occur from October- occur April-June April or October- • Resource needs for November germination- light, • Resource needs nutrients, and for seedlings- moisture light, nutrients, and moisture Seed Basal rosette

Reproductive Basal rosette with vegetative stalk • Flowering-mid- August-mid- • Bolting-mid-June October • Resource needs • Resource needs for for vegetation- flowers- nutrients, light, nutrients, light, moisture, and and moisture cross pollination

Figure 2-7. Life history stages of Balduina atropurpurea: seed, basal rosette, basal rosette with vegetative stalk, and reproductive individual.

SSA Report – B. atropurpurea 12 August 2019

Seed Balduina seeds germinate to high percentages (greater than 75%) after four weeks of cold stratification (Lincicome 1998, pp. 87-92). Balduina seeds may enter into an after-ripening or dormancy stage post dispersal and come out of dormancy during cold stratification. This seed germination strategy is referred to as a Type 2 response (Baskin and Baskin, 1993, p. 64) and allows seeds to mature or afterripen during the winter, which helps them germinate at lower temperatures in the spring. The after-ripening stage (winter dormancy period) allows Balduina to germinate when conditions, such as light, moisture, and nutrients, are favorable for plant growth in early spring (Baskin and Baskin, 1993, p. 65). Overall, Balduina seeds have high viability and germination occurs rapidly within 8 to 14 days (Lincicome 1998, p. 87).

Table 2-2. Balduina atropurpurea individual resource needs by life stage. H=Habitat, N=Nutrition, R=Reproduction, D=Dispersal. Key resource needs: high available light, high available moisture, nutrients, cross pollination, and limited vegetative competition. Resource Resource and/or circumstances needed for Life stage function individuals to complete life stage (HNRD) Cold stratification H Exposed bare mineral soil H

Seed Light Availability H Moisture H, N Nutrients N Wind to dislodge seeds from receptacle D Limited woody and graminoid competition H, N Seedling/Basal Summer precipitation and high available soil N Rosette moisture Sunlight for photosynthesis N Limited woody and graminoid vegetative H, N Basal Rosette competition with Vegetative Summer precipitation and high available soil N Stalk moisture Sunlight for photosynthesis N Cross pollination/pollinators R High available soil moisture N Reproductive Limited woody and graminoid vegetative Individual H, N competition Nutrients N, R

Population size and light availability appear to influence the seed set of Balduina (Lincicome 1998, pp. 85-88). On Fort Stewart, larger populations had greater seed set in comparison with small populations (less than 50 individuals) suggesting a positive relationship between population size and seed set (Lincicome 1998, p. 86). In addition, light availability appeared to

SSA Report – B. atropurpurea 13 August 2019

influence seed set, because populations in open conditions had higher seed set (Lincicome 1998, p. 85).

Seedling/Basal Rosette and Basal Rosette with Vegetative Stalk As discussed above, light availability plays an important role in the health of the basal rosette because of its small stature. Due to high plant species diversity and density in pine savannas, light competition is the limiting resource in this habitat. The majority of photosynthetic tissue for Balduina lies close to the ground surface with very few cauline (stem) leaves occurring along the stem. At Fort Stewart, occurrences with higher light availability produced robust individuals with taller stems in comparison to occurrences with low to moderate light conditions (Lincicome 1998, p. 53).

Reproductive Individual The reproductive success of Balduina relies upon cross-pollination. Balduina produces self- incompatible flowers that depend upon visitation by pollinators (Parker and Jones, p. 357). Pollinator activity can increase in high light areas due to the rise in temperature that supports the foraging of ectothermic pollinators (Kilkenny and Galloway, p. 247-248). Many plants in open areas often produce a greater floral display and taller stem lengths, which result in greater pollinator visitation (Kilkenny and Galloway, pp. 247-248). For Balduina, floral production has been found to be greater in high light conditions suggesting that light availability plays an important role in the reproductive success of the species by influencing both pollinator behavior, and floral display. (Lincicome 1998, p. 78; Kilkenny and Galloway 2008, p. 248).

2.6 Population and Species Level Needs

At the population level, resource needs include the key needs of individuals mentioned above (high available light, high available moisture, cross pollination, and limited vegetative competition), sustainable population size (more than 100 individuals) (Ellstrand and Elam 1993, p. 219), and connectivity between populations. Small population size (fewer than 100 individuals) can increase the risk of genetic drift (changes in allele frequency/disappearance of particular genes) and inbreeding depression (mating of related individuals) (Ellstrand and Elam 1993, pp. 218-219). Small population size and isolated populations can change pollinator behavior by causing less visitation due to the limited resources available to pollinators. Small, isolated populations of rare plant species often receive less pollinator visitation in comparison with larger or more widespread plant species (Ellstrand and Elam 1993, p. 227). Higher seed set was observed for large Balduina populations on Fort Stewart in comparison with small populations (Lincicome 1998, p. 88), suggesting that large Balduina populations receive greater pollinator visitation. Reduced seed set serves as an indicator of pollinator loss (Kearns and Inouye 1997, p. 298). Lastly, populations need suitable habitat within the population area and between populations to create connectivity. Habitat and population connectivity provides higher pollinator visitation rates to flowers in comparison with fragmented habitats or populations (Kearns and Inouye 1997, p. 299). Connected populations can have up to three times higher visitation rates in comparison with isolated populations. Overall, the Balduina populations on Fort Stewart in comparison with the small, isolated populations on private land in Georgia highlight the importance of population size and connectivity in regards to viability.

SSA Report – B. atropurpurea 14 August 2019

For the species’ viability, there must be adequate redundancy (suitable number of populations, distribution of populations, and connectivity between populations to allow the species to withstand catastrophic events) and representation (suitable genetic and environmental diversity to allow the species to adapt to changing environmental conditions). Redundancy improves with increasing numbers of populations at moderate and high resiliency conditions and connectivity (either natural or anthropogenic-influenced) among those populations to allow populations to “rescue” each other after catastrophic events. Representation improves with increased genetic diversity and environmental conditions within and among populations.

SSA Report – B. atropurpurea 15 August 2019

CHAPTER 3 – FACTORS INFLUENCING VIABILITY

The following discussion provides a summary of the factors that are affecting or could be affecting the current and future condition of Balduina throughout some or all of its range. Risks that are not known or not suspected to have effects on Balduina populations, such as disease, are not discussed in this SSA report.

3.1 Habitat Destruction and Modification

Historically, across the range of Balduina, longleaf pine-dominated woodland and savanna habitats occurred extensively across the landscape (Figure 3-1) (Frost 1993, p. 18; Edwards et al. 2013, p. 366). A long history of habitat destruction and modification exists across the longleaf pine savanna and flatwood ecosystem where Balduina occurs. During the early Colonial Period, 1564 to 1900, longleaf pine exploited for tar, pitch, turpentine, and rosin was disappearing from the landscape. Further, with the advent of railroads, 1865-1870, followed northern investors that operated logging companies in the Southeast. By 1890, the majority of longleaf virgin timber was gone from the landscape in the Southeast (Frost 1993, p. 28). Longleaf pine did not regenerate naturally in many areas and was largely replaced by loblolly pine (Frost 1993, p. 31).

Figure 3-1. Presettlement range and major divisions of the longleaf pine ecosystem. Figure from “Four Centuries of Changing Landscape Patterns in the Longleaf Pine Ecosystem” by Cecil Frost.

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In conjunction with the removal of virgin longleaf pine timber, many longleaf pine woodlands and savannas were converted to agriculture and industrial pine plantations (Frank and Platt 2006, p. 7; Edwards et al. 2013, p. 367). Agricultural conversion largely occurred across the Coastal Flatwoods of North Carolina, South Carolina, Georgia, and Florida, where Balduina occurs, due to the gentle topography and tillable land (Peet and Allard, 1993, p. 46). Further, acreage in cropland increased with the advent of center pivot irrigation systems. For example, in the Coastal Plain of Georgia, where the majority of Balduina populations occur, more than 17,000 pivot irrigation systems were in operation by 2001, irrigating 1,500,000 acres (Edwards et al. 2013, p. 358). Water for these irrigation systems comes from surface streams or groundwater and alters seasonally inundated or saturated environments (Edwards et al. p. 358). Further, many wetlands have been drained for cropland (Plummer 1963, p. 732) or converted to farm ponds (Baker 2019, pers. comm.). In Georgia, Balduina occurs in the Dougherty Plain and Tifton Uplands where many seepage slope bogs and isolated wetlands have been converted to farm ponds (Edwards et al. 2013, p. 358). This impact of land conversion due to agriculture production has largely occurred on private lands, while Balduina occurrences on protected, well managed land across Fort Stewart remains intact (Figure 3-2).

Over time, marginal productive crop land in the Coastal Plain has been converted to slash or loblolly pine plantations. Intensive silvicultural practices involved with commercial pine production, including site preparation, herbicide application, raised planting sites, and fast rotation cycles (12 to 25 years), has resulted in stands with reduced species diversity, including loss of suitable habitat for Balduina. Further, on bottomland sites, the removal of the most valuable trees, i.e., high-grading, has resulted in more shade-tolerant species, altering vegetative composition (Edwards et al. 2013, pp. 358-361), which can result in greater woody vegetation encroachment and competition to Balduina.

Another factor contributing to the heavily fragmented land in the Coastal Plain of the Southeast is urban sprawl and development. Terando et al. (2014, p. 4) estimates the amount of land in urban areas in the Southeast region was approximately 90,700 kilometers squared (or approximately 7.4% of the land area) in 2009. Urbanization results in the direct loss of Balduina habitat, fragmentation of habitat, and increases the wildland-urban interface. In South Carolina, one population has become extirpated due to industrial development (Brown 2018, pers.comm.). Future urban sprawl projections in the Southeast suggest that there would be a doubling or tripling of land devoted to urban and suburban uses (Terando et al. 2014, p. 4).

As pine savanna and flatwood ecosystems have been converted to other land uses, this results in either the complete loss of Balduina, its habitat, and the surrounding ecosystem or the habitat conditions become unsuitable and resource needs become unavailable for Balduina. Today, habitat destruction and modification from land conversion and development continues to be a major factor influencing the viability of Balduina. The longleaf pine-dominated woodland and savanna ecosystem where Balduina occurs is now considered one of the most globally imperiled ecosystems in North America due to agricultural conversion, industrial pine plantations, urban development and fire suppression (Edwards et al. 2013, pp. 366-367).

SSA Report – B. atropurpurea 17 August 2019

3.2 Lack of Fire Management

Balduina occurs throughout the Southeast in areas with a historical average fire return interval of two to four years (Guyette et al. 2012, p. 330) (Figure 3-3). The warm, humid climate of the Southeast combined with high growing season rainfall events created a condition where there was plenty of volatile fuel to ignite in the presence of a lightning strike (Finch et al. 2012, p. 105). Historical accounts suggest that the indigenous people of the Southeast Coastal Plain conducted wide ranging burns on an annual basis to clear the forest undergrowth (Edwards et al. 2013, p. 356). Native Americans reportedly had an annual custom of “setting the woods on fire many miles in extent” for hunting (Catesby 1731, p. 13). After the removal of indigenous people from the land, anthropogenic prescribed fires in the Southeast were still largely practiced on an annual rotation as many farmers used fire as a tool to “green up” the woodlands for cattle and other livestock (Frost 1993, p. 34).

Figure 3-2. This map displays agriculture production and land conversion on private land and protected land (Fort Stewart) across Long, Liberty, Evans, and Tattnall counties, GA.

The shift from free roaming cattle to enclosed livestock due to the “stock laws” likely resulted in more agriculture production and less livestock grazing. As, farmers shifted from cattle ranching to agriculture, the need to “green up” the woodlands for cattle decreased and fire suppression likely increased as a result of this shift across the Southeast (Frost 1993, p. 28). By 1920-1950, suppression of wildfires and anthropogenic fires due to modern fire laws resulted in a major shift causing most longleaf pine savanna and flatwood ecosystems to be overgrown with dense second-growth forests (Frost 1993, p. 38). The removal of spring and autumn fires resulted in a

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dense thicket of woody vegetation “where one could not see for fifteen paces” (Frost 1993, p. 34). In contrast, before widespread fire suppression, one could see for a distance of a half of mile (Frost 1993, p. 34).

Balduina cannot persist underneath a closed canopy with dense woody vegetation. This species occurs in open longleaf pine savanna and flatwood habitat maintained by prescribed fire, mowing, or hydrological conditions. Historically, across the Coastal Plain of the Southeast, there were glades and openings in the “woods of pine trees” (Catesby 1731, p. 13). Extirpated Balduina occurrences often appear overgrown with dense woody vegetation due to the removal or loss of prescribed fire in the system (Figure 2-4). Open habitats with sparse trees allow light to penetrate the ground and thereby support the herbaceous density necessary to provide fine fuels, regular fire, and herbaceous dominance (Martin and Martin 2009, p. 910). Prescribed fire can provide a ‘window of recruitment’ for herbaceous species, like Balduina, by reducing dominant shrub and grass species (Platt et al. 2006, p. 45). Maintaining an average fire return interval similar to historical intervals reduces woody vegetation and competition to Balduina as well as introduces nutrients needed by the species to grow and reproduce.

Approximate range of Balduina

Figure 3-3. “Historic (1650–1850 CE) mean fire interval estimates for the presence of fire in all or part of an average 1.2 km2 area. Classification intervals are in 2-year classes (1–30 years), 5-year classes (31–50 years), 25- year classes (50–200 years), and a single class for intervals greater than 200 years.” Source: Guyette et al. 2012, p. 330.

Although the majority of longleaf pine occurs on private land in the Southeast, well-managed, fire-maintained longleaf pine ecosystems with diverse understory vegetation predominantly occur on public lands, such as Federal and State forests, gamelands, and military bases (Peet and Allard 1993, p. 46). The fire frequency, 1 to 3 years, required to suppress hardwoods and

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maintain suitable habitat for Balduina needs a prescribed fire program. Protected Federal and State managed properties often have prescribed fire programs and land management plans. In fact, over 65% of robust (more than 100 individuals) extant Balduina populations occur on well- managed public land. Fire management and suppression continues to be one of the main factors influencing the current viability of this species. Land management actions performed at the appropriate intervals are helpful to create suitable habitat conditions for Balduina, which can result in more resilient populations.

3.3 Hydrological Impacts

Seasonally-wet longleaf pine savannas and flatwoods have been reduced in both size and number due to hydrological impacts from agriculture and development. Road-drainage ditches, field-tile piping, stream channelization, and farm ponds have all contributed to the drainage of seasonally- wet longleaf pine savannas and flatwoods (Plummer 1963, p. 733; Folkerts 1982, p. 266). Road- drainage ditches, as shallow as 2 dm (7.9 in), and field-tile piping can lower the water table and cause the natural plant community to shift from seasonally-wet to mesic longleaf pine savanna (Plummer 1963, p. 733; Folkerts 1982, p. 266). Since Balduina is a facultative wetland species, even a slight lowering of the water table could have lasting negative impacts, resulting in greater woody encroachment, reduced seedling recruitment, and reduced nutrients entering into the system. This slight lowering of the water table from road-drainage ditches or other anthropogenic features (e.g., large ruts) could decrease the “window of opportunity” for Balduina by reducing the number of days for seed germination. Seed germination for many plant species depends upon the constant imbibement of water through the seed coat. Further, percolating soil water from the uplands carries soluble nutrients, especially following a prescribed fire, to the system (Plummer 1963, p. 733). The nutrients post prescribed fire likely help and support healthy vegetative growth and reproduction.

On Fort Stewart, Balduina populations with apparent standing water and natural hydrological conditions tended to have greater herbaceous plant diversity and less ground disturbance by hogs (i.e., hogs do not seem to “root” with standing water present) (Osborn 2018, pers. comm.). In contrast, Balduina populations occurring in areas with no apparent standing water (e.g., flatwoods) had higher shrub density and greater hog disturbance. The soil disturbance caused by feral hogs creates the “perfect seedbed for pioneer plants like pines, which in turn, require mowing or burning at a young age to control” (Hattaway and Osborn 2018, p. 9). Young Balduina rosettes, potentially 1-to-2 year-old seedlings, have been observed in areas with mucky soil and small areas of standing water at Canoochee Bogs in Georgia (Punsalan 2018, unpublished data).

Impacts to the natural hydrological conditions in Balduina habitat and the surrounding ecosystem from anthropogenic disturbances and invasive species can lead to competition and woody encroachment, which reduces light, nutrients, and water availability for Balduina to grow and reproduce. These impacts have occurred at an individual and population-level for the species.

3.4 Non-Native Invasive Species

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Feral hogs (Sus scrofa) can negatively affect almost all aspects of ecosystem structure and function (Jolley et al. 2010, p. 519) and are known to have significant impacts to native plant communities both directly through consumption and indirectly through rooting and soil disturbance (Barrios-Garcia and Ballari, 2012, pp. 2284-2293). At this time, feral hogs have been documented to negatively impact “almost ever site on Forest Stewart” where Balduina occurs (Hattaway and Osborn, p. 9). The hogs root and upturn the soil resulting in exposed seedbed, damaged habitat, and loss of Balduina individuals (Osborn 2018, pers. comm.). In addition, depending on the time of year, the upturning of the soil can cause a community shift from high herbaceous forb diversity to more graminoids and young pine trees (Osborn 2018, pers. comm.). On Fort Stewart in Georgia, the hogs have been documented to impact Balduina populations and their habitat for over 20 years (Lincicome 1998, p. 50). Balduina rosettes have been upturned by hogs, with the rootstock completely exposed (Lincicome 1998, p. 50). To date, feral hog disturbance has been noted for Balduina populations in Georgia and Florida.

3.5 Climate Change

In the southeast United States, several climate change models have predicted an increase in drought frequency and intensity, air temperatures, and heavy precipitation along with increased storm events (Burkett and Kusler 2000, p. 314; Klos et al. 2009, p. 699; IPCC 2013, pp. 3-29). When taking into account future climate projections for temperature and precipitation in areas where Balduina occurs, warming is expected to be greatest in the summer, which is predicted to increase drought frequency, while annual mean precipitation is expected to increase slightly, leading to a slight increase flooding events (Figures 3-4 and 3-5) (IPCC 2013, entire; USGS 2019, unpaginated).

Figure 3-4. Time series of the seasonal average of maximum air temperature in the South Atlantic region with historical (black), RCP4.5 projection (blue), and RCP8.5 projection (red). “The historical period ends in 2005 and the future periods begin in 2006. The average of 30 CMIP5 models is indicated by the solid lines and their standard deviations are indicated by the respective shaded envelopes.” Source: USGS National Climate Change Viewer (Credit: Alder and Hostetler 2013, unpaginated; Hostetler and Adler 2016, entire; Thrasher et al. 2013, entire).

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Figure 3-5. Time series of the seasonal average of precipitation in the South Atlantic region with historical (black), RCP4.5 projection (blue), and RCP8.5 projection (red). “The historical period ends in 2005 and the future periods begin in 2006. The average of 30 CMIP5 models is indicated by the solid lines and their standard deviations are indicated by the respective shaded envelopes.” Source: USGS National Climate Change Viewer (Credit: Alder and Hostetler 2013, unpaginated; Hostetler and Adler 2016, entire; Thrasher et al. 2013, entire).

Although uncertainty exists within each model, results suggest that climate change will have an impact on wetlands due to an increase in temperature (Burkett and Kusler 2000, p. 319), which can lead to more drought conditions. Wetlands are more vulnerable to changes in surface and ground water hydrology because they occur in the transition zone between aquatic and terrestrial environments (Burkett and Kusler 2000, p. 315). Since Balduina is a facultative wetland species occurring in freshwater wetland ecosystems, sensitive to changes in surface and ground water, climate change may cause additional stress to the species in the future. For example, during drought periods, Balduina occurrences on Fort Stewart were not visible (i.e., remained dormant), and only one out of nine Balduina occurrences was visible during the 2010 drought (Mincey 2018, pers. comm.). However, once rainfall patterns returned to normal, the Balduina occurrences reappeared (Mincey 2018, pers. comm.). Survey efforts for this species are best timed during years with high rainfall (Patrick 2018, pers. comm.), highlighting the importance of precipitation and the species’ vegetative and sexual reproduction in a given year. This highlights the importance of soil moisture and precipitation for the growth and reproduction of Balduina. Changes in temperature and precipitation patterns may have a negative impact on Balduina in the future.

Balduina populations occurring in degraded and impacted wetlands may be more sensitive to fluctuations in temperature, precipitation, and evaporation resulting from climate change than populations in intact and managed wetland systems. In the lower 48 states, approximately one- half of the wetlands have been converted for other land uses. The function and structure of many wetlands have been degraded and impacted from ditching, diking, draining, and impounding, leaving many wetland systems vulnerable to changes in precipitation and temperature (Burkett and Kusler 2000, p. 315).

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At Fort Stewart, Balduina populations occurring in areas with apparent standing water appear to have higher resiliency (i.e., larger population size, can withstand longer periods of time without fire, less woody competition) than populations in drier habitat. Vegetative competition from graminoids and woody species tends to be reduced in wetter and standing water conditions. Further, hogs are less likely to “root” in standing water conditions (Osborn 2018, pers. comm.). Lastly, populations likely receive a nutrient flush from water infiltrating from surrounding uplands. These factors combined help make Balduina populations in more intact wetland systems more resilient and may buffer populations from fluctuations in precipitation and temperature in the future. In contrast, changes in precipitation and temperature will likely have a negative impact on populations occurring in fragmented wetland habitat, such as roadside rights- of-way near ditches.

In the future, fire prescription and frequency may decrease with increasing temperatures and occurrence of drought (Mitchell et al. 2014, p. 321). Constraints to managing habitat with prescribed fire is likely one of the most substantial risk factors associated with climate change for this species. Predicted changes in temperature and precipitation could limit the number of days with suitable conditions for prescribed burns. As the ability to implement prescribed fire becomes further constrained, the ability to reduce woody vegetation and maintain an open habitat will be limited, resulting in habitat conditions and resource needs becoming degraded for Balduina. Further, these changes will have a major negative impact on Balduina populations occurring in fragmented landscapes with degraded wetland systems. As mentioned above, Balduina populations occurring in drier habitat, without pockets of standing water, have less resiliency. These drier habitats depend upon frequent fire prescription to keep the habitat open, keep woody vegetation low, and help put nutrients back into the system.

In conclusion, in the future, climate change could have a negative impact on Balduina populations with low to very low resiliency due to the fact that these populations tend to occur in fragmented, degraded wetland habitat, which are vulnerable to changes in precipitation and temperature. Drought has been documented to have short-term impacts on Balduina populations in more intact wetlands as well (as discussed above). Lastly, management of habitat using prescribed fire could be constrained in the future, which could result in degraded habitat conditions and/or direct loss of Balduina populations.

3.6 Conservation Measures

Suitable habitat for Balduina is found within Department of Defense lands, State lands, and other conservation areas across the species’ range (Table 3-1). Many of these landowners implement beneficial land management practices (e.g., prescribed burning, mowing, mechanical removal of woody vegetation) to maintain the structure and function of wet pine savannas where Balduina occurs by mimicking natural disturbance regimes (Gilliam and Platt 2006, p. 7). Removing woody vegetation encroachment reduces vegetative competition thereby increasing the availability of water, nutrients, and light. The growth and reproduction of Balduina depends upon these natural resources. At present, the majority (80%) of high resiliency Balduina populations occur on protected lands, and over 65% of robust (more than 100 individuals) populations occur on well-managed public lands. Conservation lands owned by Federal and State agencies are expected to remain protected and managed for conservation purposes in the

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near future, which would eliminate the risk of direct habitat loss due to urbanization in these areas.

Table 3-1. Protected lands that provide suitable habitat for extant Balduina atropurpurea populations. State Owner/Manager Property South Carolina Depart. Of Defense Fort Jackson

Georgia Depart. Of Defense Fort Stewart* State Doerun Pitcherplant Bog

Florida State Cary State Forest* Ralph E. Simmons State Forest* Jennings State Forest* *Balduina is listed in an Integrated Natural Resource Management Plan or State Land Management Plan. Sources: Florida Department of Agriculture and Consumer Resources 2012, p. 12; Florida Department of Agriculture and Consumer Resources 2016, p. 11; Florida Department of Agriculture and Consumer Resources 2018, p. 12; Carlile 2019, pers. comm.; Hawkins 2019, pers. comm.

3.7 State Protections

Balduina atropurpurea is listed as State endangered in Florida and North Carolina. Balduina is considered state-protected in Georgia under the Georgia Wildflower Protection Act (GWPA) of 1973 and is listed as “Rare” indicating that is a “species which may not be endangered or threatened but which should be protected because of its scarcity.” This “rare” status confers the same protections as a species listed as Endangered, Threatened, or Unusual under the GWPA. In South Carolina, Balduina is not listed by the State, and South Carolina does not currently list any plant species as ‘state threatened’ or ‘state endangered’ (Brown 2019, pers. comm.). South Carolina has included Balduina in their State Wildlife Action Plant (SWAP) as a “high priority” plant (Smith 2019, pers. comm.).

3.8 Synergistic Effects

In addition to impacting Balduina individually, it is likely that several of the above summarized risk factors are acting synergistically or additively on the species. The combined impact of multiple stressors is likely more harmful than a single stressor acting alone. For example, at the known population in Rosindale Longleaf Pine Forest, the combination of fire suppression and lack of mowing has allowed woody plants to encroach into the seepage slope savannah where Balduina was previously found. In the future, drought projections could lead to further fire suppression activities, which would result in poorer habitat conditions for the population. The combination of these factors has probably impacted the Rosindale Longleaf Pine Forest population, in that the species has not been relocated despite survey efforts. On Fort Stewart,

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drought conditions and feral hogs have been documented as impacting Balduina populations. Prescribed fire and mechanical removal of woody vegetation in combination with normal precipitation patterns has resulted in populations becoming re-established following drought at Fort Stewart; this suggests habitat management around bogs and seeps where Balduina occurs is beneficial and can help higher resiliency populations “hold on” during drought conditions.

3.9 Summary of Factors Influencing Viability

We reviewed and summarized the factors that could be affecting the viability of Balduina (Figure 3-6). Concerns about Balduina’s viability focused on the following factors: (1) habitat destruction and modification, (2) fire suppression, (3) hydrological impacts, (4) non-native invasive species, and (5) climate change.

The longleaf pine-dominated woodland and savanna ecosystem where Balduina occurs is now considered one of the most globally imperiled ecosystems in North America due to agricultural conversion, industrial pine plantations, urban development, and fire suppression (Edwards et al. 2013, pp. 366-367). Across Balduina’s range, the transition zone between longleaf pine uplands and aquatic wetlands has been heavily impacted by habitat destruction and modification. Large tracts of land, containing both uplands and aquatic wetlands, are needed to protect these transitions zones. This fire-dependent ecosystem, “fire forest”, currently depends upon human application of prescribed fire due to the fragmentation of the Coastal Plain landscape and regional fire suppression (Edwards et al. 2014, p. 367-368). Balduina and its habitat requires frequent fire prescription to maintain the open conditions and facilitate the nutrient release needed for growth and reproduction. Across the Southeast, land is heavily fragmented, wetlands are degraded, and fire prescription is becoming increasingly harder for managers to implement. The habitat based factors seem to be abated on public lands due to intact and managed upland and aquatic wetlands that maintain the transition zone, where Balduina occurs. Further, fire prescription on public land is more stable due to larger tracts of intact land and availability of resources. However, feral hogs and availability of resources for fire prescription still affect the viability of Balduina on public land.

In summary, the primary factors impacting the viability of Balduina are habitat based: habitat degradation and loss resulting from fire suppression, lack of other management activities including mowing, and land conversion of wetlands to other land uses including pine plantations, agriculture, and development. In addition, the current constraints on the ability to manage habitat through prescribed fire, mowing, and mechanical treatments will be further exacerbated by urbanization and climate change in the future.

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Figure 3-6. Influence diagram for Balduina atropurpurea showing relationships between factors and species’ viability. Color meanings: Dark Pink – Land Management; Orange – Negative Factors; Light Pink – Resource Needs; Green – Population Metrics; Light Blue – Components of Viability (3 Rs); Dark Purple – Species Viability.

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CHAPTER 4 – POPULATION AND SPECIES NEEDS AND CURRENT CONDITION

In this chapter, we consider Balduina’s historical distribution, its current distribution, and the species’ needs for viability. First, we define what constitutes a population for Balduina. Next, we characterize the species’ needs in terms of resiliency (population-level), representation (species-level), and redundancy (species-level) (the 3Rs). Finally, we estimate the current condition of Balduina by using habitat and population metrics used to characterize the 3Rs.

4.1 Delineating Populations

We delineated Balduina populations using NatureServe’s Habitat-based Plant Element Occurrence Delimitation Guidance (NatureServe Explorer 2018, unpaginated). For each Balduina element occurrence, we used NatureServe’s decision tree to determine the separation distance between element occurrences. Since Balduina populations do not occur along riparian areas, we used a 2-kilometer (km; 1.24 miles (mi)) separation distance rule to separate populations. We buffered the element occurrence records by 2 km (Figure 4-1); element occurrence records were joined into a single Figure 4-1. Example of delineating population when the occurrences overlapped within populations using the 2-km separation rule. the buffer. When occurrences within 2 km were disjunct from one another for more than 1 km due to agriculture fields or pine plantations, we used a 1 km separation distance rule to separate populations.

Element occurrence data was obtained from State Natural Heritage Programs, including North Carolina Natural Heritage Program (2016), South Carolina Heritage Trust Program (2018), and Georgia Department of Natural Resources (2019), and from personal communication with Chafin (2018), Jenkins (2018), and Schotz (2018).

4.2 Methods for Estimating Current Condition

For the purpose of this assessment, we defined viability as the ability of Balduina to sustain populations in pine savanna and flatwood ecosystems over time. Using the SSA framework, we described viability of Balduina by estimating the current condition, and predicting the future condition (Chapter 5), of metrics used to assess resiliency, representation, and redundancy.

4.2.1 Population Resiliency

Each population of Balduina needs to be able to withstand, or be resilient to, stochastic events or disturbances (e.g., drought, flooding, fluctuations in seed viability, etc.). To have high resiliency, these populations need to have an adequate number of individuals, cover a large enough area (multiple occurrences within a population) that a localized event does not eliminate

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a population, and have connectivity among occurrences within a population. As outlined below, we used population and habitat metrics to assess current population resiliency of Balduina.

Resiliency-Habitat Metrics We revised a habitat and population metrics datasheet, designed for NatureServe’s 2018 Pond Cypress Savanna Classification Project, to assess the resiliency of Balduina populations across the species’ range (Table 4-1, Appendix A). The habitat metrics included five metric categories that were designed to assess the vegetation, hydrology, and fire management of each Balduina population (Appendix A). Three vegetation metrics (native herbaceous ground cover, vegetation structure/canopy cover, and shrub cover) were used to assess the habitat condition of each Balduina population. We used one hydrology metric that was scored based on the evidence of ruts, drainage ditches, and/or hog damage (Appendix A). Further, we used one fire metric to assess the fire return interval of each Balduina population.

As discussed above in Section 2.6, Balduina needs light, water, and nutrients to grow and reproduce. Therefore, limited competition from trees and shrubs is important to allow high availability of these resources to Balduina. In addition, unaltered hydrology and a regular fire return interval are important mechanisms to reduce woody encroachment and competition as well as introduce and flush nutrients needed by Balduina to grow and reproduce. Finally, the reproductive success of Balduina depends upon cross-pollination. High light conditions increases the floral display of Balduina (Lincicome 1998, p. 78), increases forb diversity, and attracts more pollinators (Kilkenny and Galloway 2008, p. 248).

Therefore, habitat components that resulted in high resiliency for a Balduina population included high native herbaceous ground cover (more than 50%), low canopy cover (less than 15%), low shrub cover (less than 10%), intact hydrology (no ruts, ditches, hog damage, etc.), and a 1-to-2 year fire-return interval (Table 4-1, Appendix A). Poor habitat or habitat with very low resiliency included less than 10% herbaceous groundcover, more than 60% canopy cover, more than 50% shrub density, a high evidence of hydrological impacts, and a fire return interval of more than 5 years (Table 4-1, Appendix A).

To summarize the current population habitat resiliency, each of the five habitat metrics were given a score from 1 to 4, with 4 being high and 1 being very low (Appendix A). Then, we computed a habitat resiliency score for each population by adding the scores across the five habitat metrics. Final scores were evaluated using the following scale: high resiliency is 20 to 16; moderate is 15 to 11; low is 10 to 6; very low is less than or equal to 5 (Appendix A).

Resiliency-Population Metrics In addition to the habitat metrics, we used two population metrics (number of individuals per population and number of occurrences within a population) to assess resiliency of Balduina populations (Table 4-1; Appendix A). Due to the fact that the reproductive success of Balduina depends upon cross-pollination and larger, connected populations receive more pollinator visitation (Ellstrand and Elam 1993, p. 227), the number of occurrences per population combined with the number of individuals per population effectively communicates population resiliency. The number of individuals per population metric included four condition categories: high resiliency (more than 500 to 1000 individuals), moderate resiliency (100 to 500 individuals), low

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resiliency (50 to 99 individuals), and very low resiliency (fewer than 50 individuals). The number of occurrences within a population metric included four condition categories, ranging from four occurrences per population as high resiliency to one occurrence per population as very low resiliency (Table 4-1; Appendix A).

To summarize the population metric resiliency scores, the two metrics were given a score from 2 to 10, with 10 being high and two being very low (Appendix A). Then, we computed a population metric resiliency score for each population by adding the scores across the two population metrics. Total scores were evaluated using the following scale: high resiliency is 20 to 17; moderate is 16 to 11; low is 10 to 5; very low is less than or equal to 4. In order to equally weight habitat (5) and population (2) metrics to summarize overall current population resiliency (see below), the population metric scores were double weighted (Appendix A).

We considered including percent of flowering individuals as a metric to assess population resiliency. This metric included four condition categories: high (more than 50% flowering), moderate (25 to 50%), fair (15 to 25%), and poor (less than 15%) flowering. However, we decided to omit this metric, because every element occurrence had over 50 percent of flowering individuals, despite the total number of individuals.

Current Population Resiliency To summarize the overall current population resiliency, we combined the habitat and population metrics for 10 extant Balduina population into a final current condition resiliency (High, Moderate, Low, and Very Low). The population and habitat metric scores were weighted equally. Datasheets were completed for every occurrence within a population. Therefore, we had to compute an average score for Balduina populations with multiple occurrences.

In the summer and fall of 2018, the habitat and population metric datasheet was completed for 10 Balduina populations (Appendix B, Table B-1). We do not have completed datasheets for historical and extirpated Balduina occurrences. For 28 extant Balduina populations without a completed habitat and population metrics datasheet, we compiled state heritage data and used the two population metrics (number of individuals per population and number occurrences within a population) to compute population resiliency (High, Moderate, Low, and Very Low; Appendix B, Table B-2). Data collected using the habitat and population metric datasheets illustrated that large (over 500 individuals) Balduina populations generally have higher quality habitat (high herbaceous diversity, low canopy, and low shrub cover). As such, the population metrics seem to reflect habitat suitability; meaning if the habitat is in suitable condition and managed appropriately, the population will have higher resiliency. Therefore, for populations without completed datasheets, we computed resiliency scores using the two populations metrics and recent state heritage data.

Six of the 28 Balduina populations did not have completed metric datasheets or state heritage data that included the number of individuals. Therefore, we used the best available science to assign a resiliency score based on the time since last seen, land protection status, and number of occurrences per population. Based on this information, the six populations were assigned a very low resiliency score, because the populations have not been seen for greater than five years, occur on private land, and only have one occurrence per population.

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Table 4-1. Population and habitat metrics used to assess and determine the current condition and population resiliency of Balduina atropurpurea.

Resiliency Population Factors Habitat Factors

Number of Number of Occurrences or Fire- Condition Individuals sub EO Native Herbaceous Vegetation Shrub Cover Hydrology Return Category per occurrences Ground Cover Structure Interval Population within a Population Savanna, seep, Less than No evidence of Over 50% forbs bog or >500-1000 10% low ruts, drainage, High ≥4 throughout depression 1-2 years individuals shrub cover ditches, hog depression/wetland meadow (< 15% throughout damage canopy cover) Little evidence 10-20% low- 25-50% forbs scattered Woodland (15- of ruts, 100-500 moderate Moderate 3 throughout 30% canopy drainage, 3 years individuals shrub cover depression/wetland cover) ditches, hog throughout damage

Moderate 25-50% 10-20% occasional Woodland (35- evidence of 50-99 moderate Low 2 forbs throughout 60% canopy ruts, drainage, 5 years individuals shrub cover depression/wetland cover) ditches, hog throughout damage High evidence Less than 10% forbs Over 50% Forest (greater of ruts, Less than 50 throughout the dense shrub Over 5 Very Low ≤1 than 60% drainage, individuals depression/wetland cover years canopy cover) ditches, hog area throughout damage

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4.2.2 Species Representation and Redundancy

Representation reflects a species’ adaptive capacity to respond to changing environmental conditions over time and can be characterized by the breadth of genetic and ecological diversity within and among populations. For Balduina, we do not have information on the genetic diversity within and among all of the populations across the species’ range. However, for Balduina populations at Fort Stewart, the genetic variation within and among populations was found to be similar (Halward et al. 1997, p. 290). Demonstrating that for an out-crossing species, populations in close proximity to each other and occurring in the same ecoregion will likely have similar genetic diversity.

In the absence of species-specific genetic and ecological diversity information, representation can be assessed based on the extent and variability of habitat characteristics across the geographical range. Balduina occurs in different ecoregions across its geographic range that could influence its genetic makeup and adaptive capacity to respond to environmental changes. Ecoregions are a system of classification based on physiography, where areas with similar characteristics of land formation, dominant soil and vegetation types, climate, air and sea currents, and distribution of flora and fauna are grouped into a single ecoregion (Bailey 1983, 365; Bailey et al. 1994, entire). Ecoregions have been used to reflect broad areas within which local adaptations and genetic coadaptation have likely occurred. Therefore, we used ecoregions as an appropriate proxy for factors likely to influence the adaptive capacity of Balduina across the landscape.

To understand the representation of Balduina, we mapped populations across ecoregions in the Southeast. We used the U.S. Environmental Protection Agency’s (EPA) Level IV Ecoregions spatial map to delineate Balduina representation units (EPA 2018, unpaginated). We delineated six representation units based on ecoregions and major river systems (Great Pee Dee River, Savannah River, Altamaha River, and Flint River) (Figure 4-2). We measured representation as the number of resilient populations within each of the delineated representation units.

Redundancy is characterized by having multiple, resilient populations across the species range. Having multiple, high resiliency populations can increase the species’ ability to persist after a catastrophic event. Species that are well-distributed across their historical range are considered less susceptible to extinction and more likely to be viable than species confined to a small portion of their range (Carroll et al. 2010, entire; Redford et al. 2011, entire). For Balduina, redundancy is characterized by having multiple resilient populations within ecoregions and across the species’ entire geographic range. In addition, these multiple, resilient populations should also maintain levels of connectivity among them. To determine redundancy, we assessed the number of extant and historical Balduina populations within defined ecoregions and across the species’ entire range (historical and current range).

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Figure 4-2. Representation units for Balduina atropurpurea. Six units were selected based on ecoregion groups and major river systems (Great Pee Dee River, Savannah River, Altamaha River, and Flint River).

4.3 Current Condition

4.3.1 Populations

Overall, 140 Balduina element occurrence records exist range-wide, from years 1902 to 2018 (Figure 4-3). Using the methods in Section 4.1 above, we delineated 79 Balduina populations from the 140 occurrence records. Of these 79 Balduina populations, 38 remain extant, 39 are historical, and two are extirpated. The majority (64) of Balduina populations occur in Georgia. Of the 38 exant pouplations, there are 31 in Georgia, five in Florida, one in North Carolina, and one in South Carolina. Seventeen extant populations occur on protected lands, while 21 occur on private property. Across this species’ range, approximately half of the Balduina populations are considered historical. Of the 39 historical Balduina populations, 37 occur on private property. Further, 12 of these historical populations occur within rights-of-way. This information demonstrates the importance of land protection and management to Balduina.

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Figure 4-3. Distribution of 140 Balduina atropurpurea occurrence records across the species' range.

4.3.2 Current Population Resiliency

Of the 38 extant Balduina populations, five presently have high resiliency: St. Mary’s River Ranch (FL), Ralph E. Simmons Memorial State Forest (FL), Cary State Forest (FL), Lake Creek Seep and Ward Property (GA), and Fort Stewart Main Population (GA) (Table 4-2; Figure 4-4). The common thread across all of the high resiliency populations is land management, either through a prescribed fire program or annual mowing. Further, the majority (80%) of high resiliency populations occur on protected land.

The largest Balduina population occurs on Fort Stewart. This population contains over 2,000 individuals distributed across nine occurrences (GADNR 2018, unpaginated; Hattaway and Osborn 2018a, pp. 1-13). The number of individuals per occurrence across Fort Stewart greatly fluctuates from 0 to more than 500 individuals per occurrence. Factors that appear to increase the abundance of Balduina at each occurrence include low canopy cover (less than 25%), low shrub density (less than 35%), intact hydrology, minor hog damage and few tank ruts, and frequent fire. For example, two occurrences (EO 83; site names: RCW 137 and Bobby’s Island), with more than 400 individuals, occur in areas near red-cockaded woodpecker clusters that

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receive frequent fire. Further, both of these sites contain approximately 20% canopy cover and more than 55% native forb diversity (Hattaway and Osborn 2018a, pp. 45-48). In contrast, factors that appeared to decrease the abundance of Balduina across Fort Stewart include hydrological degradation (EO 84, e.g. low water crossing), high canopy cover (more than 35%), fire impediment from old fire breaks or tank ruts, and hog damage. For example, one occurrence (EO 38, site name: Carol Helton’s N5) had more than 50% canopy cover and only one Balduina individual. This occurrence is “poor, due to shrub and pine competition and a lack of recent fire coverage” (Hattaway and Osborn 2018a, p. 10). Overall, Balduina occurrences in areas with intact hydrology, low canopy cover, low shrub density, high native forb diversity, and frequent fire have greater resiliency. These results mirror past survey and research efforts across Fort Stewart that illustrated an increase in Balduina individuals in high light conditions (Lincicome 1998, p. 73) and that “wet savanna soils are subject to rapid woody invasion in the absence of fire” and “require the historical fire frequency to prevent loss of fire dependent species”, such as Balduina (Frost 1988, p. 3).

The Lake Creek Seep and Ward Property is a high resiliency population that occurs on private land in Turner County, Georgia. This population has greater than 200 individuals, and has multiple occurrences within the population. However, the occurrences have not been surveyed since 2002, and the landowners are not interested in conservation (Baker 2019, pers. comm.).

Three high resiliency populations occur in Florida on protected lands: Ralph E. Simmons Memorial Sate Forest (Nassau County), St. Mary’s River Ranch (Nassau County), and Cary State Forest (Duvall County). The Ralph E. Simmons population contains approximately 700 individuals with three occurrences within the population. The majority of individuals (570) occur in one occurrence along a powerline right-of-way within the State Forest (Walls and Jenkins 2018, pp. 1-4). Management, e.g., mowing, within the powerline has kept the habitat completely open from woody encroachment, and thereby supporting the robust Balduina population by increasing light conditions and soil moisture. In contrast, adjacent to the powerline, a small occurrence with one individual occurs in an open, wooded area (Walls and Jenkins 2018, p. 2). The last prescribed fire near the powerline right-of-way occurred in 2015, but did not reach the population (Walls and Jenkins 2018, pp. 1-4). Although the prescribed fire may not sweep across the powerline where Balduina occurs due to soil moisture or fire breaks, the released nutrients may enter the seepage slope via water traveling downslope from burned upland areas.

The St. Mary’s River Ranch population has more than 500 Balduina individuals. This population occurs on protected (TNC conservation easement), private land, and the St. Mary’s River Ranch, which has a prescribed fire program with a 2 to 3 year fire return interval (Wilson Baker 2018, pers. comm.). The fire management program on this property likely supports the robust Balduina population.

The Cary State Forest population contains approximately 200 individuals across two occurrences. The Balduina population occurs in a powerline right-of-way along a 1.5-mile stretch. The habitat is maintained open by frequent mowing, which occurs approximately once a year. Further, prescribed fire may occasionally creep into the population. The population was

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last burned in 2016. Hog damage and signs of ruts from maintenance vehicles was noted during 2018 surveys (Walls and Jenkins 2018, pp. 1-2).

Table 4-2. Current resiliency of Balduina atropurpurea populations across the species’ range. See Table 4-1 for current condition metrics used to assess resiliency. Resiliency Class Number of populations in Percent of total populations each resiliency class (79) High 5 6 Moderate 4 5 Low 7 9 Very Low 22 28 Historical 39 50 Extirpated 2 2 Total 79 100

BALDUINA RANGE-WIDE RESILIENCY

20 22 15

10 7 5 4 5

0 High Moderate Low Very Low

Figure 4-4. Balduina atropurpurea current population resiliency classes for extant populations across the species' range.

Four Balduina populations have moderate resiliency; of these, two populations occur on protected land (Fort Stewart Tom’s Site, GA and Jennings State Forest, FL) and two occur on private land (Canoochee Bogs and Ashburn Hill Place/Paul Murphy Road, GA) (Table 4-2; Table 4-3). The Fort Stewart Tom’s Site population contains approximately 76 individuals. This population presently “appears healthy” but “there is too much shade from young pine

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regeneration” and will need to be thinned for the population to persist (Hattaway and Osborn 2018a, p. 42).

The Canoochee Bog population contains greater than 200 individuals spread across four occurrences along a powerline right-of-way. Each occurrence has a different property owner. One of the occurrences (EO 45) is on Rayonier property and is not being managed with fire or frequent mowing. This occurrence contains only 13 individuals. In contrast, one occurrence (EO 46) is being managed with a 2 to 3 year fire return interval and contains the greatest number of individuals (175) within the population. The Georgia Plant Conservation Alliance has worked diligently with the landowner, utility company, and State, to manage this occurrence with prescribed fire. The population has increased since the start of the partnership and management (Ceska 2018, pers. comm.).

The Ashburn Hill Place/Paul Murphy Road population has approximately 58 individuals with five occurrences distributed across two private properties (Baker 2019, pers. comm.). This population occurs in a seepage bog. We do not know the current management of this population. However, fire suppression has been noted as a potential threat (Baker 2019, pers. comm.).

Overall, the moderate resiliency populations occur on both private and protected land and contain approximately greater than 50 to 200 individuals. All of the high and moderate resiliency populations occur in Georgia and Florida (Figure 4-5).

Table 4-3. Current resiliency classes for protected and unprotected Balduina populations across the species’ range. Resiliency Class All Populations Protected Not Protected High 5 4 1 Moderate 4 2 2 Low 7 3 4 Very Low 22 8 14 Historical 39 2 37 Extirpated 2 0 2 Range-wide total 79 19 60

In contrast, Balduina populations with very low resiliency contain fewer than 50 individuals and remain isolated from other populations. Of the 38 extant Balduina populations, over half (22) have very low resiliency (Figure 4-4). Further, many (14) of these populations occur on private land that is heavily fragmented (Table 4-3). Seventy percent of the populations with very low resiliency have identified threats, such as destructive logging, beneath powerline-herbicide potential, fire suppression or lack of fire prescription, woody competition, silviculture practices, roadside maintenance, and gas line maintenance, within the State element occurrence data.

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These identified threats highlight the fact that many of these populations may have very low resiliency due to lack of management and impacts from incompatible land use practices. These isolated populations will likely remain small without management, such as prescribed fire and/or mowing. Further, without nearby occurrences, a reduction occurs in both cross pollination and pollinator visitation. Overtime, these combined factors can result in inbreeding depression, thereby decreasing the species’ resiliency and ability to respond to stochastic events.

In summary, of the 38 extant Balduina populations, five currently have high resiliency and four have moderate resiliency. All of these high and moderate resiliency populations occur on public and private lands being managed with prescribed fire or mowing in Georgia and Florida. Impacts from habitat destruction and modification and fire suppression do not appear to be affecting high or moderate resiliency Balduina populations, as the species has maintained robust populations in Georgia and Florida with active land management programs. Across the range, the remaining extant populations (29) have low (7) to very low (22) resiliency. Thirty-nine Balduina populations are considered in historical status and two are extirpated.

4.3.3 Current Species Representation

To understand representation, we summarized the number and resiliency of Balduina populations across six representation units in the Southeast (Table 4-4; Figure 4-5). Of the six representation units, three representation units contain high to moderate resiliency populations (Figure 4-5). The three representation units with high and moderate resiliency populations occur in Georgia and Florida (Table 4-4; Figure 4-5). On the periphery of the species’ range in North Carolina, South Carolina, and Alabama, the majority of the populations have a resiliency of low or very low, or are considered either historical or extirpated; therefore, these units are at a higher risk of unit-wide extirpation and indicates the potential for further loss of representation. At this time, there is no known genetic variation as this analysis has not been conducted range-wide for the species. At present, Balduina has 38 extant populations in varying levels of resiliency occurring in five representation units. Therefore, the species has some level of adaptive capacity, but given the low to very low resiliency of 29 Balduina populations, the current representation is reduced. See below for a full description of the species’ condition in each representation unit.

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Table 4-4. Balduina atropurpurea representation units with the number of records, number of populations, and current population resiliency scores. Population Resiliency Classes No. of No. of Representation unit High Moderate Low Very Low Historical Extirpated records populations North Carolina- Atlantic Southern Loam Plains and 2 2 1 1 Swamps/Peatlands South Carolina- Atlantic Southern Loam Plains and 3 3 1 1 1 Sandhills Georgia North- Atlantic Southern Loam Plains and Sea 61 18 1 2 1 2 12 Island Flatwoods Georgia South-Atlantic Southern Loam Plains, Bacon Terraces, 50 40 1 1 4 17 16 1 and Tifton Upland

Florida-Georgia-Sea Island 23 15 3 1 2 1 8 Flatwoods

Alabama-Dougherty Plains 1 1 1

Total 140 79 5 4 7 22 39 2

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Figure 4-5. Balduina atropurpurea current population resiliency across representation units.

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North Carolina-Atlantic Southern Loam Plains and Swamps/Peatlands The North Carolina representation unit contains one extant and one historical record (Figure 4-6; Table 4-4). The Rosindale Longleaf Pine Forest population has not been observed since 2004 and occurs within a poorly managed powerline right-of-way. This population currently has very low resiliency. The historical population is located on the Green Swamp Preserve, a TNC owned and managed property, and was last observed in 1980.

Figure 4-6. Balduina atropurpurea current population resiliency in the North Carolina representation unit.

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South Carolina-Atlantic Southern Loam Plains and Sandhills The South Carolina representation unit contains one extant population with very low resiliency in Richland County, one extirpated population in Darlington County, and one historical population in Darlington County (Figure 4-7; Table 4-4). The extirpated population, Hartsville North, was last observed in 1921 and was impacted by development (Brown 2019, pers. comm.). The historical population, Society Hill, is known from an old annotated herbarium specimen in 1972 (Brown 2019, pers. comm.).

The remaining extant population occurs on Fort Jackson, a Department of Defense Army installation, and was last observed in 2017. This population has very low resiliency, because it contains less than 50 individuals and is isolated from other extant populations. The Integrated Natural Resource Management Plan for Fort Jackson does not include Balduina. However, the population is managed with prescribed fire, and the biologists at Fort Jackson have “been actively burning it for years now” (Chadwick 2018, pers. comm.).

Figure 4-7. Balduina atropurpurea current population resiliency in the South Carolina representation unit.

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Georgia North Unit-Atlantic Southern Loam Plains and Sea Island Flatwoods The Georgia North representation unit, located north of the Altamaha River, contains one high resiliency population on Fort Stewart. Fort Stewart serves as a refuge for Balduina, supporting and safeguarding three populations, with one population containing greater than 2000 individuals. The current resiliency and distribution of Balduina across Fort Stewart illustrates the probable distribution of this species across the landscape before major development and agriculture production (Figure 3-2). Two moderate resiliency Balduina populations, Canoochee Bogs and Tom’s Occurrence on Fort Stewart, occur within this representation unit. For the populations located outside of Fort Stewart, the majority are historical (12) or have very low (2) resiliency, likely due to agriculture production (Figure 4-8; Table 4-4). Overall, within this representation unit, the distribution of Balduina has decreased, with the majority of moderate to high resiliency populations occurring within Fort Stewart.

Figure 4-8. Balduina atropurpurea current population resiliency in the Georgia North representation unit.

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Georgia South Unit-Atlantic Southern Loam Plains, Bacon Terraces, and Tifton Upland The Georgia South representation unit, located south of the Altamaha River, contains only one high resiliency population, the Lake Creek Seep and Ward Property population (Figure 4-9; Table 4-4). This population was last observed in 2002 and occurs on private land. The landowners do not have an interest in conservation, so the status of Balduina on this property remains uncertain (Patrick 2019, pers. comm.). The remaining extant populations in this unit have moderate (1), low (4), and very low (17) resiliency. This representation unit contains the highest concentration of historical (16) populations and has one extirpated population. The lack of land protection and management likely resulted in the decline in the species’ resiliency in this representation unit.

Figure 4-9. Balduina atropurpurea current population resiliency in the Georgia South representation unit.

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Georgia-Florida-Sea Island Flatwoods The Georgia-Florida representation unit contains the Sea Island Flatwoods and extends from the southern part of Georgia into Florida. The Georgia portion of this unit contains two historical populations (Figure 4-10; Table 4-4). The majority of Balduina populations within this unit occur in Florida. This representation unit contains the highest number (3) of high resiliency populations (Figure 4-10; Table 4-4). Two of the high resiliency populations occur on State lands: Cary State Forest and Ralph E. Simmons Memorial State Forest. The third high resiliency population, St. Mary’s River Ranch, occurs on private land, which is protected by a TNC conservation easement. One population with moderate resiliency occurs on Jennings State Forest. All of the high to moderate resiliency Balduina populations within this representation unit occur on public or protected land with a prescribed fire program. Several occurrences on Ralph E. Simmons and Cary State Forest occur within powerline right-of-way managed with mowing. Eight Balduina populations have not been in observed in at least 20 years within this unit and are now considered historical, likely due to development and fire suppression.

Figure 4-10. Balduina atropurpurea current population resiliency in the Georgia-Florida representation unit.

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Alabama-Dougherty Plains The Alabama representation unit contains one historical record in Geneva County (Figure 4-11; Table 4-4). This record was first and last observed in 1988 (Schotz 2018, pers. comm.).

Figure 4-11. Balduina atropurpurea current population resiliency in the Alabama representation unit.

4.3.4 Current Species Redundancy

There has been a reduction in redundancy for Balduina within each representation unit and across the range. This is highlighted by three representation units having the majority of populations in a low or very low resiliency condition or historical status. These three representation units have no high to moderate resiliency populations. Further, the Georgia North and Georgia South units have seen a reduction in redundancy with the majority of populations having very low resiliency and only one to three populations having high to moderate resiliency.

Across the species’ range, redundancy of Balduina has been reduced from historical levels. Of the 79 Balduina populations, 38 populations are currently extant with 39 being considered historical and two extirpated. Of those extant Balduina populations, five populations have high

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resiliency and four populations have moderate resiliency. All of the high and moderate resiliency populations occur in Georgia and Florida. The remaining 29 populations have low to very low resiliency. Multiple occurrences within the same population may allow occurrences to become re-established following a catastrophic event. Given the species’ current reduced redundancy and with the majority of populations having low to very low resiliency, it might be difficult to re-establish occurrences (or an entire population) affected by a catastrophic event without human intervention, as the connectivity among some of the populations (especially populations in North Carolina and South Carolina) is low.

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CHAPTER 5 – FUTURE CONDITIONS AND VIABILITY

In the previous chapters, we reviewed Balduina ecological and resource needs (Chapter 2), factors influencing the historical, current, and future viability of the species (Chapter 3), and the current condition of the species (Chapter 4). We now consider what the species’ future conditions are likely to be. We apply our future forecasts to the concepts of resiliency, representation, and redundancy to describe the future viability of Balduina.

5.1 Introduction to Methods for Estimating Future Condition

To assess the future condition of Balduina, we have forecasted what Balduina may have in terms of the resiliency, redundancy, and representation under three plausible future scenarios. As outlined above, the primary factors influencing the viability of Balduina are habitat-based and include habitat degradation and loss resulting from fire suppression and other lack of management activities including mowing; land conversion of wetlands to other land uses including pine plantations, agriculture, and development; and active land management (prescribed fire, mowing, and mechanical treatment), which is beneficial at the appropriate interval to the species. In addition, we considered how land management of Balduina habitats may be further hampered by urbanization and climate change in the future.

Since the main factors influencing the viability of Balduina are habitat-based, we performed spatial analyses to project changes in land cover under various levels of urbanization over time. We used these projections to develop our future scenarios to predict Balduina population resiliency as influenced by changes in habitat suitability due to management and urbanization.

Although the life span of Balduina is unknown, it is a perennial with high seed viability and reproduces sexually and asexually, and therefore, we considered a projection time frame of 40 years with a 20 year time step to account for normal variation in plant reproduction and environmental conditions. In addition, we were reasonably certain we could forecast the response of Balduina to varying levels of management efforts, as well as urbanization, within this time period. Therefore, we summarized predicted population resiliency conditions for each scenario at 2040 and 2060. The time steps began in 2018, as this was the end of our current condition timeframe.

5.2 Projections and Modeling

We used a habitat suitability model and urbanization (SLEUTH) model to project change in habitat suitability due to urbanization.

5.2.1 Habitat Suitability Model

We used species distribution modeling as a way to find potential suitable habitat for Balduina based on the environmental conditions associated with known extant populations (Ramirez- Reyes et al. 2019, in prep).

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Data Used To model the habitat suitability for Balduina, we used two sets of information: a) the location of known extant locations of the species, specifically the geographic coordinates associated to the presence of the species; and b) environmental information that influence the species in the form of digital environmental maps (Table 5-1).

We used the location of extant populations of the species as a measure of presence of the species. Our initial set of locations included 97 coordinate points, but we did a spatial filtering to avoid having a particular area over represented in the model. The spatial filtering consisted of randomly removing points within 500 meters (m; 0.31 mi) of another point.

We considered what Balduina needs to grow and survive, and then, we choose environmental variables (e.g., precipitation and temperature) that could influence the presence of the species. In our modeling, we included 10 variables that were not correlated to each other (Table 5-1).

Table 5-1. Environmental variables used in habitat suitability modeling for Balduina atropurpurea. Variables Description of Variables Data Source Google Earth Engine; USGS Landsat 8 Land Surface Temperature Temperature of the ground Surface Reflectance LST 2014-2017 Elevation Digital elevation model USGS SRTM Vegetation Continuous Field Google Earth Engine; National Land Percent tree canopy cover (VCF) Cover Database 2011 USFS Fraction of the sand content in Percent sand USDA SSURGO database soils Amount of organic matter Organic matter USDA SSURGO database present in soils pH pH in soils USDA SSURGO database Normalized Difference Google Earth Engine; Landsat 7 & 8, Plant productivity; mean value Vegetation Index (NDVI) * NDVI Compooccurrence 2012–2017 Bioclim WorldClim World Climate Total precipitation during annual Precipitation Annual Data – Bio 12 Annual Precipitation period; mean value (mm); 1950–2000 Google Earth Engine; MODIS Burned Burned Area Fire frequency for recent fires Area and LANDFIRE 2001-2016 Maximum temperature over Bioclim WorldClim World Climate Max Temperature of Winter coldest 3 month annual period; Data – Bio 6 Maximum Temperature mean value of Coldest Quarter; 1950–2000

Modelling Approach The presence and environmental data were used in three common algorithms to model habitat suitability: generalized additive model (GAM), generalized boosted model (GBM), and maximum entropy (MAXENT). Each of these approaches produced a habitat suitability score for every pixel in the map. To verify the accuracy of these predictions, we calculated the commonly used area under the receiver operating characteristic curve (AUC). Because each of these three approaches has different assumptions and strengths, and therefore a different AUC,

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we used a model ensemble approach to minimize uncertainties associated with each individual model. We built the ensemble model by averaging the results of each of the three approaches weighted by their performance. In this way, an approach that performed well will have higher weight in the ensemble. The final model ensemble produced a habitat suitability map for the species containing values ranging from 0 (lowest suitability) to 1 (highest suitability).

5.2.2 Urbanization Model

Approach and Scenarios We used SLEUTH urbanization projections (Terando et al. 2014, entire) for the Southeastern U.S. at two time periods: 2040 and 2060. Each of these datasets map a baseline of developed areas (circa 2009) and the expected probability for an area to be developed in that decade. We created a simulated habitat suitability Balduina under these development projections assuming two levels of urbanization probabilities:

(1) Under the High Urbanization scenario, we included all probabilities for an area to be developed.

(2) Under the Low Urbanization scenario, we only included areas with a very high probability, greater than or equal to 90%, of being developed.

The equation to calculate this future habitat suitability is: Future habitat suitability = Current habitat suitability * (1-Probability of development)

We also calculated the percent habitat loss with the equation: Percent habitat loss = 1- (Future habitat suitability/ Current habitat suitability)

Analysis Results Predicted habitat losses over the entire modeled area suggests a fairly steady linear pattern over time. Under the Low Urbanization scenario, habitat losses were 5.7% in 2040 and 10.1% in 2060. Similarly, under the High Urbanization scenario, habitat losses were 7.4% in 2040 and 12.6% in 2060. Whereas this pattern of steady habitat loss over time appeared to hold for Balduina representation units and populations, the magnitude of habitat loss was much more variable at these smaller spatial scales. For the six representation units, habitat loss by 2060 ranged from 8.9% to 13.8% under the Low Urbanization Scenario and 10.7% to 17.0% under the High Urbanization scenario (Table 5-2). For example, total predicted habitat loss under the high urbanization was 13.8% by 2060. For the 79 Balduina populations, habitat loss by 2060 ranged 0.0% to 73.9% under the Low Urbanization Scenario and 0.0% to 85.9% under the High Urbanization scenario (Appendix C).

Considering Urbanization in Future Scenario Development After reviewing the results for the High and Low Urbanization scenarios, we decided to select one urbanization risk scenario for inclusion in our future condition scenarios. The difference in habitat loss between the High and Low Urbanization scenarios was minimal, ranging from a 1 to 10% difference. Therefore, we selected the High Urbanization scenario for our future urbanization projection modeling.

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Table 5-2. Percent habitat loss due to urbanization within each Balduina atropurpurea representation unit. Low Urbanization High Urbanization Representation Unit 2040 2060 2040 2060 North Carolina- Atlantic Southern Loam Plains and 4.6% 8.9% 5.9% 10.7% Swamps/Peatlands South Carolina- Atlantic Southern 8.6% 13.8% 10.8% 17.0% Loam Plains and Sandhills Georgia North- Atlantic Southern Loam Plains and Sea Island 6.2% 10.8% 7.9% 13.3% Flatwoods Georgia South-Atlantic Southern Loam Plains, Bacon Terraces, and 5.5% 10.2% 7.3% 12.7% Tifton Upland Florida-Georgia-Sea Island 6.1% 10.3% 8.2% 13.1% Flatwoods

Alabama-Dougherty Plains 7.0% 12.8% 9.7% 16.1%

Under the High Urbanization scenario, we included all probabilities for an area to be developed and determined percent habitat loss within a 5 km (3.1 mi; 2 km (1.2 mi) population + 3 km (1.9 mi) buffer zone) buffer for each population. We considered that populations projected to lose greater than 25% of their suitable habitat within the 5 km buffer at 2040 and 2060 have a higher risk of development. Conversely, we considered that populations projected to lose less than 25% of their suitable habitat within the 5 km buffer at 2040 and 2060 have a low risk of development.

5.3 Incorporating Management into the Future Scenarios

We developed three plausible scenarios to assess future resiliency conditions of Balduina. All three scenarios predicted future conditions under varying levels of management effort with a High Urbanization scenario component.

5.3.1 Scenario 1 – Status Quo, Management for Stability

Under the Scenario 1, we considered a future where the amount of protected lands for each population will remain constant through time with no new protected areas being acquired and no new Balduina populations are established. Management effort to achieve population stability (e.g., prescribed fire at 2 to 4 year fire return interval, mowing, etc.) for all currently managed populations is continues and management would not be hindered by funding, political issues, climate change, or other factors. As a result, population resiliency scores remain stable unless

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there is a high risk of development (25% or greater habitat loss) that could hinder management at 2040 or 2060 (Table 5-3).

5.3.2 Scenario 2 – Decreased Management

Under the Scenario 2, we considered a future where management effort for all Balduina populations decreases due to a change in priorities or resources. In addition, management efforts are hindered by increased temperatures and drought as future climate change predictions are realized. The amount of protected lands for each population will remain constant through time and no new populations are established. In this scenario, with decreases in management effort, population resiliency scores decrease (Table 5-3).

5.2.3 Scenario 3 – Increased Management and Conservation

Under the Scenario 3, we considered a future where resources are readily available and additional management efforts are focused on improving habitat suitability for all Balduina populations. All populations are receiving a regular fire return interval of 1 to 3 years. Further, populations within rights-of ways are being maintained by manual removal of woody vegetation and mowing. The regular fire return interval maintains the high and moderate resiliency populations. Management effort is not hindered by projected climate change impacts, and under increased management, some effects of climate change to the species are alleviated by regular prescribed fire, mowing, and mechanical removal of woody is occurring. In this scenario, with increases in management, population resiliency increases when there is low risk of development impacts, but stays the same when there is a high risk (Table 5-3).

Summary of Scenarios The three future scenarios that included varying levels of management effort: decreased management (a decrease in fire return interval and a decrease in manual removal of woody vegetation), management for stability (current management for populations on protected land, 2- to-4 year fire-return interval, and/or manual removal of woody vegetation), and increased management (increase in woody vegetation removal and increase in fire prescription at 1-to-3 year fire return interval). Using the High Urbanization scenario component, we developed the two categories: (1) low risk of development and (2) high risk of development. We developed a rule set to determine the change in resiliency of Balduina populations from current to future conditions based on the interaction between risk of development and management for the three scenarios (Table 5-3). Then, we determined future population resiliency at 2040 and 2060.

Table 5-3. Interaction between risk of development and management in three plausible future scenarios to determine future resiliency of Balduina atropurpurea populations Decreased Management for Increased Management Stability Management Development-Low Resiliency decrease No Change Resiliency increase Risk Development-High Resiliency decrease Resiliency decrease No Change Risk

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5.4 Future Condition

5.4.1 Future Population Resiliency

Using the Balduina habitat suitability and urbanization model, we predicted future habitat loss given a High Urbanization scenario and then determined the change in resiliency for 38 extant Balduina populations from current to future conditions based on the interaction between risk of development and management under three scenarios at two time steps: 2040 (Table 5-4; Figure 5-1) and 2060 (Table 5-5; Figure 5-2). See below for a summary of the population resiliency condition under each scenario at 2040 and 2060.

Table 5-4. Future resiliency for Balduina atropurpurea populations at 2040 under three future scenarios.

Future Scenarios Management Management Resiliency Current Status Quo Decrease Increase High 5 5 0 9 Moderate 4 4 5 7 Low 7 7 4 20 Very Low 22 20 7 2 Extirpated 2 4 24 2 Historical 39 39 39 39 Total 79

45 40 35 30 25 20 15 10 5 0 Current Status Quo Management Management Decrease Increase

High Moderate Low Very Low Extirpated

Figure 5-1. Balduina atropurpurea future population resiliency across the species' range at 2040 under three future scenarios.

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Table 5-5. Future resiliency for Balduina atropurpurea populations at 2060 under three future scenarios. Future Scenarios Management Management Resiliency Current Status Quo Decrease Increase High 5 4 0 8 Moderate 4 4 5 8 Low 7 8 4 18 Very Low 22 18 7 4 Extirpated 2 6 24 2 Historical 39 39 39 39 Total 79

45 40 35 30 25 20 15 10 5 0 Current Status Quo Management Management Decrease Increase High Moderate Low Very Low Extirpated

Figure 5-2. Balduina atropurpurea future population resiliency across the species' range at 2060 under three future scenarios.

Scenario 1 Under Scenario 1 (status quo management with high risk of development), the projected change in resiliency from current to future would result in two additional Balduina populations being lost (resiliency changes from current – very low to future – extirpated) due to a high risk of development (greater than 25% of suitable habitat loss) in 2040, resulting in 36 extant populations (Table 5-2; Figure 5-1). Of the remaining 36 extant Balduina populations, five are predicted to continue to have high resiliency and four continue have moderate resiliency. All of these high and moderate resiliency populations will continue to occur on public and private lands being managed with prescribed fire in Georgia and Florida. Across the range, the remaining extant populations (27) are predicted to continue to have low to very low resiliency. Thirty-nine Balduina populations will remain in historical status.

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At 2060, the projected change in resiliency from current to future would result in an additional four Balduina populations being lost (resiliency changes from current – very low to future – extirpated) due to a high risk of development (greater than 25% of suitable habitat loss) at 2060, resulting in 34 extant populations (Table 5-3; Figure 5-2). In addition, one Balduina population with current high resiliency is predicted to change to moderate resiliency and one with current moderate resiliency will change to low resiliency. Lastly, 26 Balduina populations are predicted to have low to very low resiliency at 2060. These changes in resiliency are attributed to the high risk of development resulting in greater than 25% suitable habitat loss within these populations.

Scenario 2 Under Scenario 2 (decreased management with high risk of development), the projected change in resiliency from current to future would result in an additional 22 Balduina populations being lost (resiliency changes from current – very low to future – extirpated) due to a high risk of development and decreased management at 2040 and 2060 (Table 5-2; Figure 5-1; Table 5-3; Figure 5-2). Of the remaining 16 extant Balduina populations, no populations are predicted to have high resiliency in the future as all of these populations (5) will change to having moderate resiliency at 2040 and 2060. Eleven extant populations are predicted to have low to very low resiliency at 2040 and 2060. Thirty-nine Balduina populations will remain in historical status.

Scenario 3 Under Scenario 3 (increased management with high risk of development), the projected change in resiliency from current to future would result in no additional Balduina populations being lost (resiliency changes from current – very low to future – extirpated) at 2040 and 2060 (Table 5-2; Figure 5-1; Table 5-3; Figure 5-2). In addition, two of 38 extant populations are projected to have very low resiliency in 2040 and four populations will have very low resiliency at 2060. Further, there would be an increase in the number of high (9) and moderate (7) resiliency populations at 2040. At 2060, there is still predicted to be an increase in the number of high (8) and moderate (8) resiliency populations from current (5 and 4, respectively). However, there is a higher risk of development at 2060 resulting in one fewer population at high and moderate resiliency at this time step. Thirty-nine Balduina populations will remain in historical status.

In summary, if management continues as today as under Scenario 1, the projected change in resiliency results in losing two Balduina populations at 2040 and four populations at 2060 due to development. Across the range, more than half (27 and 26) of the remaining extant populations (36 and 34) are predicted to continue to have low to very low resiliency under Scenario 1 at 2040 and 2060, respectively. In contrast, if management efforts increase across the species’ range as in Scenario 3, the projected change would result in no additional Balduina populations being lost and four populations would have very low resiliency in 2060. Further, nine populations are predicted to have high resiliency at 2040, an increase from five populations having high resiliency currently. If management efforts increase in the future, the majority of Balduina populations are predicted to have an increase in resiliency, resulting in 20 and 18 populations with low resiliency at 2040 and 2060, respectively. If management efforts decrease in the future as under Scenario 2, over half (22) of the currently extant Balduina populations (38) are predicted to be extirpated in 2040 and 2060. Currently, many of the Balduina populations (22) have very low resiliency and occur in fragmented habitat. As such, if management efforts

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decrease in the future, they will be lost from the landscape. Further, with a decrease in management, no high resiliency populations are predicted to occur in 2040 and 2060.

5.4.2 Future Species Representation

To understand future species representation, we summarized the number and resiliency of Balduina populations across six representation units under the three future scenarios at 2040 and 2060 (Table 5-6; Figure 5-3; Figure 5-4; Figure 5-5).

Scenario 1 Under Scenario 1 (status quo management with high risk of development), future representation is predicted to remain the same for all Balduina populations within representation units except for two populations in the Georgia South and Florida-Georgia units at 2040 (Table 5-6; Figure 5- 3). Both populations have very low resiliency and are predicted to be extirpated due to greater than 25% of their suitable habitat being developed 2040. At 2060, two populations are predicted to be extirpated in the Georgia South unit, one population in the Georgia North unit, and one population in the Florida-Georgia unit.

Of the six representation units, three representation units are predicted to have high to moderate resiliency populations at 2040 and 2060 (Figure 5-3). The three representation units with high and moderate resiliency populations occur in Georgia and Florida. In the North Carolina, South Carolina, and Alabama units, the populations are predicted to maintain a condition of low, very low, historical, or extirpated at 2040 and 2060. These units are predicted to remain at a higher risk of unit-wide extirpation and indicate a potential for continued loss of representation.

At 2040, Balduina is predicted to have 36 extant populations (38 populations currently) in varying levels of resiliency occurring in five ecoregions. At 2060, the species is predicted to have 34 extant populations occurring in five ecoregions. Therefore, the species has some level of adaptive capacity, but given the low to very low resiliency to extirpated status of 31 Balduina populations at 2040 and 32 populations at 2060, the future representation is similar to current levels of representation.

Scenario 2 Under Scenario 2 (decreased management with high risk of development), future representation is predicted to decline for Balduina (Table 5-6; Figure 5-4). At 2040, the one current population in South Carolina is predicted to be lost. As such, there would be no species representation in the South Carolina representation unit. In addition, the one extant population in North Carolina is predicted to be lost. A decrease in management would result in the loss of both North Carolina and South Carolina representation units by 2040. The Georgia South representation unit is predicted to experience a contraction since many of these populations have very low resiliency (17) currently and are predicted to be extirpated (17) at 2040 and 2060; in addition, no population is predicted to have high resiliency, one population is predicted to have low resiliency, and one population is predicted to have moderate resiliency at 2040 and 2060.

The Georgia North and Florida-Georgia representation units are predicted to no longer contain high resiliency populations; while under current conditions, these units have one and 3

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populations, respectively. Both units are predicted to have very low (1, 2), low (2, 1), and moderate (1, 3) resiliency populations, respectively; while the remaining populations in both (2, 1) are predicted to become extirpated.

Of the six representation units, no representation units are predicted to have high resiliency populations at 2040 and 2060 (Figure 5-4). Three representation units are predicted to have moderate resiliency populations, and these occur in Georgia and Florida. Four representation units are predicted to have some extirpated populations at 2040 and 2060 (Figure 5-4).

At 2040 and 2060, Balduina is predicted to have 16 extant populations (38 populations currently) in varying levels of resiliency occurring in three ecoregions. Given the low to very low resiliency to extirpated status of 35 Balduina populations at 2040 and 2060, the future representation is greatly reduced.

Scenario 3 Under Scenario 3 (increased management with high risk of development), future representation is predicted to increase for Balduina (Table 5-6; Figure 5-5). One population in the North Carolina unit is predicted to change from very low to low resiliency at 2040 and 2060. The one population in the South Carolina unit is predicted to change from very low to low resiliency at both time steps. The Georgia South representation unit is predicted to have 16 populations (17 currently) change from very low to low resiliency at 2040 and 15 from very low to low resiliency at 2060 (Table 5-6). In addition, the high and moderate resiliency populations will increase from one population each to two and four populations, respectively, at 2040 and 2060. In the Georgia North and Florida units, high and moderate resiliency populations are predicted to increase (Figure 5-5). No currently extant populations are predicted to be extirpated under this scenario.

At 2040 and 2060, Balduina is predicted to have 38 extant populations (38 populations currently) in varying levels of resiliency occurring in five ecoregions. Given the low to very low resiliency of 22 Balduina populations at 2040 and 2060, the future representation under Scenario 3 has increased from current representation levels.

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Table 5-6. Balduina atropurpurea population resiliency across for the six representation units under three future scenarios at 2040 and 2060. 2040 2060 Scenario Scenario Scenario Scenario Population Resiliency Scores Current Scenario 2 Scenario 3 1 2 3 1 North Carolina- Atlantic Southern Loam Plains and Swamps/Peatlands – Number of Populations: 2 High Moderate Low 1 1 Very Low 1 1 1 Extirpated 1 1

Historical 1 1 1 1 1 1 1

South Carolina- Atlantic Southern Loam Plains and Sandhills – Number of Populations: 3

High

Moderate

Low 1 1

Very Low 1 1 1

Extirpated 1 1 2 1 1 2 1

Historical 1 1 1 1 1 1 1

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2040 2060 Scenario Scenario Scenario Scenario Population Resiliency Scores Current Scenario 2 Scenario 3 1 2 3 1 Georgia North- Atlantic Southern Loam Plains and Sea Island Flatwoods – Number of Populations: 18

High 1 1 3 1 3

Moderate 2 2 1 1 1 1 1

Low 1 1 2 2 2 2 2

Very Low 2 2 1 1 1

Extirpated 2 1 2

Historical 12 12 12 12 12 12 12

Georgia South-Atlantic Southern Loam Plains, Bacon Terraces, and Tifton Upland – Number of Populations: 40

High 1 1 2 1 2

Moderate 1 1 1 4 1 1 4

Low 4 4 1 16 4 1 15

Very Low 17 16 4 1 15 4 2

Extirpated 1 2 18 1 3 18 1

Historical 16 16 16 16 16 16 16

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2040 2060 Scenario Scenario Scenario Scenario Population Resiliency Scores Current Scenario 2 Scenario 3 1 2 3 1 Florida-Georgia-Sea Island Flatwoods – Number of Populations: 15

High 3 3 4 2 4

Moderate 1 1 3 2 2 3 2

Low 2 2 1 2 1

Very Low 1 1 2 1 2 1

Extirpated 1 1 1

Historical 8 8 8 8 8 8 8

Alabama-Dougherty Plains – Number of Populations: 1

High

Moderate

Low

Very Low

Extirpated

Historical 1 1 1 1 1 1 1

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(a)

(b)

Figure 5-3. Balduina atropurpurea population resiliency across representation units for Scenario 1 (status quo management with high urbanization) at two time steps (a) 2040 and (b) 2060.

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(a)

(b)

Figure 5-4. Balduina atropurpurea population resiliency across representation units for Scenario 2 (decreased management with high urbanization) at two time steps (a) 2040 and (b) 2060.

SSA Report – B. atropurpurea 61 August 2019

(a)

(b)

Figure 5-5. Balduina atropurpurea population resiliency across representation units for Scenario 3 (increased management with high urbanization) at two time steps (a) 2040 and (b) 2060.

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5.3.3 Future Species Redundancy

As outlined above in Chapter 4, there has been a reduction in current redundancy for Balduina with 36 populations predicted to be extant in 2040 and 34 in 2060 across the species range. Under Scenario 1, the future species redundancy is predicted to remain similar to current redundancy. Three representation units are predicted to have the majority of populations in a low or very low resiliency and historical condition at 2040 and 2060. As under current condition, these three representation units are predicted to have no high to moderate resiliency populations in the future. Further, the Georgia North and Georgia South units is predicted to have a reduction in redundancy with the majority of populations in a very low condition and only one to three populations having high to moderate resiliency. Under current condition, 38 populations are extant with 39 being considered historical. Due to projected development, two popuations become extirpated in 2040 and four in 2060. In 2040 and 2060, nine and eight populations, respectively, have high to moderate resiliency. All of the high and moderate resiliency populations occur in Georgia and Florida.

Under Scenario 2, future redundancy at 2040 and 2060 is predicted to be further reduced from current conditions. In 2040 and 2060, 24 populations are predicted to be extirpated due to a decrease in management. In addition, the South Carolina and North Carolina ecoregion would no longer contain any populations at 2040 and 2060. In addition to the 24 extirpated populations, 11 Balduina populations are predicted to be in low to very low resiliency at 2040 and 2060. This would greatly reduce the redundancy of this species across its range. Given the lack of connectivity and amount of isolated populations under this scenario, without human intervention and cultivated plants, it may be difficult to re-establish occurrences affected by a catastrophic event.

In contrast, under Scenario 3, there is an increase in management for Balduina populations, and therefore, the number of high and moderate resiliency populations is predicted to increase at 2040 and 2060. Given this increase from current condition, future redundancy is predicted to increase for Balduina. Under the increase in management scenario, at 2040, there would be nine highly resilient and seven moderately resilient populations (Figure 5.3). In 2060, under an increase management scenario, we would have eight highly resilient populations (Figure 5.4). Because the majority of highly resilient populations occur on public land, only one highly resilient population, St. Mary’s River Ranch, would be impacted by development in 2060.

Under Scenario 3 (increased management with high risk of development), future redundancy is predicted to increase for Balduina (Table 5-6; Figure 5-5). One population in the North Carolina unit is predicted to change from very low to low resiliency at 2040 and 2060. Further, no currently extant populations are predicted to be extirpated under this scenario. Having higher resiliency populations spread across the range and representation units may allow occurrences within a population and low to very low resiliency populations to become re-established following a catastrophic event.

SSA Report – B. atropurpurea 63 August 2019

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APPENDIX A

Balduina atropurpurea-Habitat Metrics Data Sheet Recorded Measured RANK/SCORE Value of Metric Vegetation Metrics High= 4.0 Moderate= 3.0 Low = 2.0 Very Low= 1.0 VEG1. Native >50% 25-50% 10-25% <10% %Native Herbaceous Throughout Scattered Occasional Throughout herbaceous

Ground Cover depression or throughout throughout depression or plant cover:

wetland area depression or depression or wetland area wetland area wetland area VEG2. Vegetation Savanna, seep, Woodland (15- Woodland (30- Forest (>=60% % canopy Structure bog or 30% canopy 60% canopy canopy cover) cover: depression cover) cover) meadow (<=15% canopy cover) VEG3.Shrub <10% Low 10-25% Low- 25-50% >50% Dense % shrub cover: Cover shrub cover moderate shrub Moderate shrub shrub cover throughout cover cover throughout throughout throughout ecotone

Hydrology and Fire Metrics HYD1. Hydrology No evidence of Little evidence Moderate High evidence ruts/drainage of ruts/drainage evidence of of ruts/drainage ditches/hog ditches/hog ruts/drainage ditches/hog damage. damage. ditches/hog damage. damage. FIRE1. Fire- Fire-return Fire-return Fire-return Fire-return return interval interval 1-2 interval-average interval- interval > 5 years 3 years average 5 years years Final Score is: Total Evaluation Scale: High= 20 to 15; Moderate=15 to 10; Low=10 to 5; Very Low = ≤5 Score =

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Balduina atropurpurea-Population Metrics Data Sheet

High= 10.0 Moderate= Low=5.0 Very Rank/Score: 8.0 Low=2.0 POP1. >500-1000 100-500 50-99 <50 Number of Number of individuals individuals individuals individuals Individuals: Individuals

POP2. % No. ≥4 3 2 ≤1 Number of of Occurrences occurrences per per population: population

Final Score is: Total Evaluation Scale: High: 20 to 17; Moderate 16 to 11; Low=10 to 5; Very Score = Low=≤4

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APPENDIX B

Table B-1. Current resiliency computed from completed Habitat and Population Metric Datasheets for extant Balduina atropurpurea populations in 2018.

Element Habitat Metrics Population State Population Ownership Resiliency Occurrence Score Metrics Score NC Rosindale 0002-26035 Low Very Low Private Very Low Longleaf 0 individuals Pine Forest (ROW) GA Canoochee 0033-3909, Moderate High Private Moderate Bogs, 0045-0974, 250 (Powerline 0046-7186, individuals ROW) 0047-2309 GA Fort Stewart, 0088- High High Public High Main 22111, >2000 Population 0083- individuals 21294, 0084- 21295, 0028-1632, 0087- 22109, 0036-8583, 0042-2953, 0038-8876, 0037-2710 GA Fort Stewart, 0082-21293 Moderate Low Public Moderate Tom's 76 individuals Occurrence GA Fort Stewart, 0086-22107 Moderate Very Low Public Low Carol 5 individuals Helton's N4 GA Race Pond 0080-19930 Moderate Very Low Private Low Bog 15 individuals (Powerline ROW) FL Cary State 0008, 0012 High Moderate Public High Forest 210 individuals FL Jennings 0006, 0009 Moderate Moderate Public Moderate State Forest 112 individuals FL Ralph E. 0003, 0011, High High Public High Simmons 0013 723 Memorial individuals State Forest

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Table B-2. Current resiliency computed using population metric data for Balduina atropurpurea populations from State heritage programs.

# of Date Occurre Land Date Last # of occurrenc of Last State nce EO # Protection/ Resiliency Observed individuals es per Known Name Ownership population Survey FL St. 14 6/30/1905 >500 3 Private, CE High Mary's River Ranch FL Jenning 7 11/3/2003 7 1 11/3/2 State Very Low s State 003 Forest GA Arrowh 0076- 9/27/2007 195 (2007), 1 9/27/2 Private, CE Very Low ead 15691 <50 (2015- 007 Farms 2018) GA Ashbur 0060- 9/20/2013 58 4 9/20/2 Private Moderate n Hill 6357, 013 Place 0061- and 2804 Tree Farm Rd GA Brooker 0071- 9/16/2003 unknown 1 9/16/2 Private Very Low Bog 4608 003 (powerli ne row) GA Cotting 0068- 10/17/2001 unknown 1 10/17/ Private Very Low ham 4438 2001 Propert y GA Doe 0044- 9/1/2018 <50 1 9/1/20 State Very Low Run 11089 18 Pitcherp lant Bog GA Douglas 0069- 9/16/2003 unknown 1 9/16/2 Private Very Low Radio 8739 003 Towers GA GA32 at 0026- 2010 unknown 1 7/2/19 Private Very Low Little 10518- 05 Sandy 31 Creek GA General 0074- 9/1/2016 <50 1 9/1/20 Private Very Low Coffee 12189 16 State Park GA Jeffords 0077- 9/11/2008 <50 3 9/11/2 Private Low Tract 15692 008

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GA Lake 0055- 8/16/2002 213 4 8/16/2 Private High Creek 7931, 002 Seep 0056- and 3276, Ward 0057- Propert 4132, y 0058- 11170 GA Marietta 0064- 9/15/2001 40 1 9/15/2 Private Very Low Thomas 1274- 001 Bog 47 GA Mary 0049- 11/10/1998? unknown 1 na Private Very Low Bluff 3922 Rd (Powerl ine ROW) GA McClen 0043- 8/25/2018 19 1 8/25/2 Private Very Low don Bog 2457 018 (Powerl ine ROW) GA Mitchell 0008- 10/1/2018 <50 1 10/1/2 Private Very Low Brother' 5628- 018 s Bog 15 GA N Side 0054- 10/11/2001 <50 1 10/11/ Private Very Low SR 112 9421 2001 (powerli ne ROW) GA NE of 0070- 9/16/2003 Unknown 1 9/16/2 Private Very Low Douglas 8740- 003 (Powerl 52H ine row) GA NW of 0031- 2000 unknown 1 2000 Private Very Low Ashbur 3710 n (betwee n road and railroad ) GA Page 0078- 7/11/2007 <50 1 7/11/2 Private Very Low Tract 15693 007 (Powerl ine row) GA Plant 0072- 9/16/2003 unknown 1 9/16/2 Private Very Low Hatch 1794, 003 Powerli 0050- ne 3923- (Powerl 37H ine row),

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West of Denton

GA Quail 0063- 9/7/2001 35 3 9/7/20 Private, Ridge 11363- 01 Tall Plantati 46 Timbers on CE Very Low GA Quail 0065- 9/6/2001 7 2 9/6/20 Private Ridge 9628, 01 Rd 0066- 3397 Low GA Spanish 0089- 10/14/2009 100-150 1 10/14/ Private, CE Creek 75439 plants 2009 North American Land Trust Low GA St. 0073- 11/13/2005 0 1 2018 Private Matthe 11969 ws Church Very Low GA Sylveste 0075- 9/26/2007 <50 1 9/26/2 Private r-Hwy 15690 007 82 (Gas line ROW) Very Low GA Turner 0053- 9/24/2015 <50 3 9/24/2 Private County 9421 015 Bog Low GA Wiley 0067- 10/1/18 20 1 10/1/2 Private Caves 9422 018 Bog Very Low GA Willis 0059- 10/15/2001 10 3 10/15/ Private Creek 6356 2001 Area

Low GA Yates 0062- 9/1/2018 <10 1 9/1/20 Private, Bog 2254 18 Tall Timbers CE Very Low

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APPENDIX C

Table C-1. Future resiliency for extant Balduina atropurpurea populations at 2040 under three future scenarios. Future Resiliency % Current Risk of Habitat Status Managemen Managemen State Populations Resilienc Urbanizatio Loss at Quo t Increase t Decrease y n 2040 FL Cary State High 8.20% Low High High Moderate Forest FL Jennings Moderate 9.90% Low Moderate High Low State Forest 1 FL Jennings Very Low 39.10% High Extirpated Very Low Extirpated State Forest 2 FL Ralph E. High 13.60% Low High High Moderate Simmons Memorial State Forest FL St. Mary's High 24.50% Low High High Moderate River Ranch GA Arrowhead Very Low 0.10% Low Very Low Low Extirpated Farms GA Ashburn Moderate 8.90% Low Moderate High Low Place, Tree Farm Rd, and Paul Murphy Rd. GA Brooker Bog Very Low 4.60% Low Very Low Low Extirpated GA Canoochee Moderate 16.80% Low Moderate High Low Bogs GA Cottingham Very Low 12.90% Low Very Low Low Extirpated Property GA Doerun Very Low 4.50% Low Very Low Low Extirpated Pitcherplant Bog GA Douglas Very Low 25.30% High Extirpated Very Low Extirpated Radio Towers GA Fort Stewart Low 0% Low Low Moderate Very Low Carol Helton's N4 GA Fort Stewart High 3.50% Low High High Moderate Main Population GA Fort Stewart Moderate 0% Low Moderate High Low Tom's Occurrence GA GA32 at Very Low 0.10% Low Very Low Low Extirpated Little Sandy Creek

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GA General Very Low 7.90% Low Very Low Low Extirpated Coffee State Park GA Jeffords Tract Low 1.40% Low Low Moderate Very Low GA Lake Creek High 2.30% Low High High Moderate Seep and Ward Property GA Marietta Very Low 0.70% Low Very Low Low Extirpated Thomas Bog GA McClendon Very Low 19.20% Low Very Low Low Extirpated Bog GA Mitchell Very Low 0.20% Low Very Low Low Extirpated Brother's Bog GA N Side SR Very Low 2.20% Low Very Low Low Extirpated 112 (Powerline ROW) GA NW of Very Low 8.40% Low Very Low Low Extirpated Ashburn GA Page Tract Very Low 10.70% Low Very Low Low Extirpated (Powerline ROW) GA Plant Hatch Very Low 0.70% Low Very Low Low Extirpated Powerline (Powerline ROW) GA Quail Ridge Very Low 0% Low Very Low Low Extirpated Plantation GA Quail Ridge Low 0% Low Low Moderate Very Low Road GA Race Pond Low 8.50% Low Low Moderate Very Low Bog GA Spanish Low 6.40% Low Low Moderate Very Low Creek GA St. Matthews Very Low 0.80% Low Very Low Low Extirpated Church GA Sylvester Very Low 17.20% Low Very Low Low Extirpated Hwy 82 GA Turner Low 4.80% Low Low Moderate Very Low County Bog GA Wiley Caves Very Low 0.80% Low Very Low Low Extirpated Bog GA Willis Creek Low 2.20% Low Low Moderate Very Low Area GA Yates Bog Very Low 0% Low Very Low Low Extirpated NC Rosindale Low 1.40% Low Low Moderate Very Low Longleaf Pine Forest SC Fort Jackson Very Low 0% Low Very Low Low Extirpated

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Table C-2. Future resiliency for extant Balduina atropurpurea populations at 2060 under three future scenarios. Future Resiliency % Current Risk of Habitat Status Managemen Managemen State Populations Resilienc Urbanizatio Loss at Quo t Increase t Decrease y n 2040 FL Cary State High 14.10% Low High High Moderate Forest FL Jennings Moderate 13.60% Low Moderate High Low State Forest 1 FL Jennings Very Low 47.50% High Extirpated Very Low Extirpated State Forest 2 FL Ralph E. High 24.50% Low High High Moderate Simmons Memorial State Forest FL St. Mary's High 40.60% High Moderate High Moderate River Ranch GA Arrowhead Very Low 0.40% Low Very Low Low Extirpated Farms GA Ashburn Moderate 15.90% Low Moderate High Low Place, Tree Farm Rd, and Paul Murphy Rd. GA Brooker Bog Very Low 9.10% Low Very Low Low Extirpated GA Canoochee Moderate 28.30% High Low Moderate Low Bogs GA Cottingham Very Low 21.20% Low Very Low Low Extirpated Property GA Doerun Very Low 9% Low Very Low Low Extirpated Pitcherplant Bog GA Douglas Very Low 36.50% High Extirpated Very Low Extirpated Radio Towers GA Fort Stewart Low 0% Low Low Moderate Very Low Carol Helton's N4 GA Fort Stewart High 7% Low High High Moderate Main Population GA Fort Stewart Moderate 0% Low Moderate High Low Tom's Occurrence GA GA32 at Very Low 0.40% Low Very Low Low Extirpated Little Sandy Creek GA General Very Low 13.40% Low Very Low Low Extirpated Coffee State Park GA Jeffords Low 3% Low Low Moderate Very Low Tract

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GA Lake Creek High 5.50% Low High High Moderate Seep and Ward Property GA Marietta Very Low 1.30% Low Very Low Low Extirpated Thomas Bog GA McClendon Very Low 33.30% High Extirpated Very Low Extirpated Bog GA Mitchell Very Low 1% Low Very Low Low Extirpated Brother's Bog GA N Side SR Very Low 3.90% Low Very Low Low Extirpated 112 (Powerline ROW) GA NW of Very Low 16.50% Low Very Low Low Extirpated Ashburn GA Page Tract Very Low 18.10% Low Very Low Low Extirpated (Powerline ROW) GA Plant Hatch Very Low 1.90% Low Very Low Low Extirpated Powerline (Powerline ROW) GA Quail Ridge Very Low 0.40% Low Very Low Low Extirpated Plantation GA Quail Ridge Low 0% Low Low Moderate Very Low Road GA Race Pond Low 14.10% Low Low Moderate Very Low Bog GA Spanish Low 9.50% Low Low Moderate Very Low Creek GA St. Matthews Very Low 2% Low Very Low Low Extirpated Church GA Sylvester Very Low 28.70% High Extirpated Very Low Extirpated Hwy 82 GA Turner Low 10.80% Low Low Moderate Very Low County Bog GA Wiley Caves Very Low 5.80% Low Very Low Low Extirpated Bog GA Willis Creek Low 3.70% Low Low Moderate Very Low Area GA Yates Bog Very Low 0.10% Low Very Low Low Extirpated NC Rosindale Low 2.90% Low Low Moderate Very Low Longleaf Pine Forest SC Fort Jackson Very Low 0% Low Very Low Low Extirpated

SSA Report – B. atropurpurea 79 August 2019

APPENDIX D

GLOSSARY

achene – a small, dry, indehiscent fruit with a single locule and a single seed (ovule), and with the seed attached to the ovary wall at a single point, as in the sunflower.

after-ripening – the process that must be undergone by certain seeds after harvest before germination can take place.

basal linear-spatulate – basal - positioned at or arising from the base, as leaves arising from the base of the stem; linear – resembling a line; long and narrow with more or less parallel side; spatulate – like a spatula in shape, with a rounded blade above gradually tapering to the base.

cauline leaves – of, on, or pertaining to the stem, as leaves arising from the stem above ground level.

cold stratification – the process of placing seeds between layers of moist peat or sand and exposing them to low temperatures in order to encourage germination.

disk – an enlargement or outgrowth of the receptacle around the base of the ovary; in the Compositae the central portion of the involucrate head bearing the tubular or disk flowers.

disk flowers – a regular flower of the Compositae.

facultative wetland plants – usually occur in wetlands, but may occur in non-wetlands. These plants predominately occur with hydric soils, often in geomorphic settings where water saturates the soils or floods the soil surface at least seasonally.

graminoid – of grasses (Gramineae) and grass-like plants, including edges and rushes (marsh plants). imbibition (imbibment) – the absorption of liquid, usually water, into the ultramicroscopic spaces or pores found in materials such as cellulose, pectin and cytoplasmic proteins in seeds. ligulate (ligule) – strap-shaped; a strap-shaped organ; the flattened part of the ray corolla. out-crossing species (outbreeding) – the crossing of plants that are not closely related genetically fitness (adaptive value) – the balance of genetic advantages or disadvantages that determines the ability of an individual organism to survive and reproduce in a given environment. peduncle – the inflorescence stalk of a plant; the stalk of a solitary flower of an inflorescence. ray flowers – a ligulate flower of the Compositae.

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receptacle – the portion of the pedicel upon which the flower parts are borne. In the Compositae (Asteraceae), the part of the peduncle where the flowers of the head are borne.

receptacular bractlets – reduced leaf or leaflike structure at the base of flower or inflorescence.

seed set – number of seeds per pollinated flower.

seepage slope – an open, grass-sedge dominated community kept continuously moist by groundwater seepage. It occurs in dissected topography, with 30 to 50-foot elevation differences, and is usually bordered by well-drained sandhill or upland pine communities. self-incompatible – the failure of gametes from the same plant to form a viable embryo. senescence (senescing) – the complex deteriorative processes that terminate naturally the functional life of an organ/organism. In plants, this is associated with flowering and fruiting, and is an important contributing factor in the decrease in chlorophyll content an in the ability of the plant to photosynthesize.

SSA Report – B. atropurpurea 81 August 2019